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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Properties_and_characteristics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Properties_and_characteristics"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Properties and characteristics</span> </div> </a> <ul id="toc-Properties_and_characteristics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Classifications_and_types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Classifications_and_types"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Classifications and types</span> </div> </a> <button aria-controls="toc-Classifications_and_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 Classifications and types subsection</span> </button> <ul id="toc-Classifications_and_types-sublist" class="vector-toc-list"> <li id="toc-RNA_plasmids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#RNA_plasmids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>RNA plasmids</span> </div> </a> <ul id="toc-RNA_plasmids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Chromids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Chromids"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Chromids</span> </div> </a> <ul id="toc-Chromids-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Vectors" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Vectors"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Vectors</span> </div> </a> <button aria-controls="toc-Vectors-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 Vectors subsection</span> </button> <ul id="toc-Vectors-sublist" class="vector-toc-list"> <li id="toc-Cloning" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cloning"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Cloning</span> </div> </a> <ul id="toc-Cloning-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_Production" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Protein_Production"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Protein Production</span> </div> </a> <ul id="toc-Protein_Production-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gene_therapy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Gene_therapy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Gene therapy</span> </div> </a> <ul id="toc-Gene_therapy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Disease_models" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Disease_models"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Disease models</span> </div> </a> <ul id="toc-Disease_models-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Biogenetic_Gene_Cluster_(BGC)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Biogenetic_Gene_Cluster_(BGC)"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Biogenetic Gene Cluster (BGC)</span> </div> </a> <ul id="toc-Biogenetic_Gene_Cluster_(BGC)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Episomes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Episomes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Episomes</span> </div> </a> <ul id="toc-Episomes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plasmid_maintenance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Plasmid_maintenance"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Plasmid maintenance</span> </div> </a> <ul id="toc-Plasmid_maintenance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plasmids_in_nature" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Plasmids_in_nature"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Plasmids in nature</span> </div> </a> <button aria-controls="toc-Plasmids_in_nature-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 Plasmids in nature subsection</span> </button> <ul id="toc-Plasmids_in_nature-sublist" class="vector-toc-list"> <li id="toc-Yeast_plasmids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Yeast_plasmids"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Yeast plasmids</span> </div> </a> <ul id="toc-Yeast_plasmids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plant_mitochondrial_plasmids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plant_mitochondrial_plasmids"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Plant mitochondrial plasmids</span> </div> </a> <ul id="toc-Plant_mitochondrial_plasmids-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Study_of_plasmids" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Study_of_plasmids"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Study of plasmids</span> </div> </a> <button aria-controls="toc-Study_of_plasmids-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 Study of plasmids subsection</span> </button> <ul id="toc-Study_of_plasmids-sublist" class="vector-toc-list"> <li id="toc-Plasmid_DNA_extraction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plasmid_DNA_extraction"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Plasmid DNA extraction</span> </div> </a> <ul id="toc-Plasmid_DNA_extraction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Conformations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Conformations"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Conformations</span> </div> </a> <ul id="toc-Conformations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Software_for_bioinformatics_and_design" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Software_for_bioinformatics_and_design"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.3</span> <span>Software for bioinformatics and design</span> </div> </a> <ul id="toc-Software_for_bioinformatics_and_design-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Plasmid_collections" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Plasmid_collections"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.4</span> <span>Plasmid collections</span> </div> </a> <ul id="toc-Plasmid_collections-sublist" class="vector-toc-list"> </ul> </li> </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">9</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">10</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Further reading</span> </div> </a> <button aria-controls="toc-Further_reading-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 Further reading subsection</span> </button> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> <li id="toc-General_works" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#General_works"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.1</span> <span>General works</span> </div> </a> <ul id="toc-General_works-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Episomes_2" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Episomes_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.2</span> <span>Episomes</span> </div> </a> <ul id="toc-Episomes_2-sublist" class="vector-toc-list"> </ul> </li> </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">12</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"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Plasmid</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 62 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-62" 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">62 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%A8%D9%84%D8%A7%D8%B2%D9%85%D9%8A%D8%AF" title="بلازميد – Arabic" lang="ar" hreflang="ar" data-title="بلازميد" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Pl%C3%A1smidu" title="Plásmidu – Asturian" lang="ast" hreflang="ast" data-title="Plásmidu" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Plazmid" title="Plazmid – Azerbaijani" lang="az" hreflang="az" data-title="Plazmid" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AA%E0%A7%8D%E0%A6%B2%E0%A6%BE%E0%A6%B8%E0%A6%AE%E0%A6%BF%E0%A6%A1" title="প্লাসমিড – Bangla" lang="bn" hreflang="bn" data-title="প্লাসমিড" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4" title="Плазмид – Bulgarian" lang="bg" hreflang="bg" data-title="Плазмид" 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/Plazmid" title="Plazmid – Bosnian" lang="bs" hreflang="bs" data-title="Plazmid" 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/Plasmidi" title="Plasmidi – Catalan" lang="ca" hreflang="ca" data-title="Plasmidi" 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/Plazmid" title="Plazmid – Czech" lang="cs" hreflang="cs" data-title="Plazmid" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Plasmid" title="Plasmid – Danish" lang="da" hreflang="da" data-title="Plasmid" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Plasmid" title="Plasmid – German" lang="de" hreflang="de" data-title="Plasmid" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Plasmiid" title="Plasmiid – Estonian" lang="et" hreflang="et" data-title="Plasmiid" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A0%CE%BB%CE%B1%CF%83%CE%BC%CE%AF%CE%B4%CE%B9%CE%BF" title="Πλασμίδιο – Greek" lang="el" hreflang="el" data-title="Πλασμίδιο" 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/Pl%C3%A1smido" title="Plásmido – Spanish" lang="es" hreflang="es" data-title="Plásmido" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Plasmido" title="Plasmido – Esperanto" lang="eo" hreflang="eo" data-title="Plasmido" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Plasmido" title="Plasmido – Basque" lang="eu" hreflang="eu" data-title="Plasmido" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%BE%D9%84%D8%A7%D8%B3%D9%85%DB%8C%D8%AF" title="پلاسمید – Persian" lang="fa" hreflang="fa" data-title="پلاسمید" 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/Plasmide" title="Plasmide – French" lang="fr" hreflang="fr" data-title="Plasmide" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Plasmaid" title="Plasmaid – Irish" lang="ga" hreflang="ga" data-title="Plasmaid" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Pl%C3%A1smido" title="Plásmido – Galician" lang="gl" hreflang="gl" data-title="Plásmido" 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/%ED%94%8C%EB%9D%BC%EC%8A%A4%EB%AF%B8%EB%93%9C" title="플라스미드 – Korean" lang="ko" hreflang="ko" data-title="플라스미드" 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/%D5%8A%D5%AC%D5%A1%D5%A6%D5%B4%D5%AB%D5%A4" title="Պլազմիդ – Armenian" lang="hy" hreflang="hy" data-title="Պլազմիդ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A5%8D%E0%A4%B2%E0%A4%BE%E0%A4%B8%E0%A5%8D%E0%A4%AE%E0%A4%BF%E0%A4%A1" title="प्लास्मिड – Hindi" lang="hi" hreflang="hi" data-title="प्लास्मिड" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Plazmid" title="Plazmid – Croatian" lang="hr" hreflang="hr" data-title="Plazmid" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Plasmid" title="Plasmid – Indonesian" lang="id" hreflang="id" data-title="Plasmid" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Plasmide" title="Plasmide – Italian" lang="it" hreflang="it" data-title="Plasmide" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A4%D7%9C%D7%A1%D7%9E%D7%99%D7%93" title="פלסמיד – Hebrew" lang="he" hreflang="he" data-title="פלסמיד" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Plasmid" title="Plasmid – Javanese" lang="jv" hreflang="jv" data-title="Plasmid" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%9E%E1%83%9A%E1%83%90%E1%83%96%E1%83%9B%E1%83%98%E1%83%93%E1%83%94%E1%83%91%E1%83%98" title="პლაზმიდები – Georgian" lang="ka" hreflang="ka" data-title="პლაზმიდები" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4%D1%82%D0%B5%D1%80" title="Плазмидтер – Kazakh" lang="kk" hreflang="kk" data-title="Плазмидтер" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4%D0%B0%D0%BB%D0%B0%D1%80" title="Плазмидалар – Kyrgyz" lang="ky" hreflang="ky" data-title="Плазмидалар" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Plazmid" title="Plazmid – Hungarian" lang="hu" hreflang="hu" data-title="Plazmid" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4" title="Плазмид – Macedonian" lang="mk" hreflang="mk" data-title="Плазмид" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%AA%E0%B5%8D%E0%B4%B2%E0%B4%BE%E0%B4%B8%E0%B5%8D%E2%80%8C%E0%B4%AE%E0%B4%BF%E0%B4%A1%E0%B5%8D" title="പ്ലാസ്‌മിഡ് – Malayalam" lang="ml" hreflang="ml" data-title="പ്ലാസ്‌മിഡ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%9E%E1%83%9A%E1%83%90%E1%83%96%E1%83%9B%E1%83%98%E1%83%93%E1%83%94%E1%83%A4%E1%83%98" title="პლაზმიდეფი – Mingrelian" lang="xmf" hreflang="xmf" data-title="პლაზმიდეფი" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target"><span>მარგალური</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Plasmid" title="Plasmid – Malay" lang="ms" hreflang="ms" data-title="Plasmid" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4" title="Плазмид – Mongolian" lang="mn" hreflang="mn" data-title="Плазмид" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Plasmide" title="Plasmide – Dutch" lang="nl" hreflang="nl" data-title="Plasmide" 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/%E3%83%97%E3%83%A9%E3%82%B9%E3%83%9F%E3%83%89" title="プラスミド – Japanese" lang="ja" hreflang="ja" data-title="プラスミド" 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-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Plasmid" title="Plasmid – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Plasmid" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Plasmid" title="Plasmid – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Plasmid" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Plasmidi" title="Plasmidi – Occitan" lang="oc" hreflang="oc" data-title="Plasmidi" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Plazmid" title="Plazmid – Polish" lang="pl" hreflang="pl" data-title="Plazmid" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Plasm%C3%ADdeo" title="Plasmídeo – Portuguese" lang="pt" hreflang="pt" data-title="Plasmídeo" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Plasmid%C4%83" title="Plasmidă – Romanian" lang="ro" hreflang="ro" data-title="Plasmidă" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4%D1%8B" title="Плазмиды – Russian" lang="ru" hreflang="ru" data-title="Плазмиды" 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/Plasmid" title="Plasmid – Simple English" lang="en-simple" hreflang="en-simple" data-title="Plasmid" 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-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Plazmid" title="Plazmid – Slovak" lang="sk" hreflang="sk" data-title="Plazmid" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Plazmid" title="Plazmid – Slovenian" lang="sl" hreflang="sl" data-title="Plazmid" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%BE%D9%84%D8%A7%D8%B2%D9%85%DB%8C%D8%AF" title="پلازمید – Central Kurdish" lang="ckb" hreflang="ckb" data-title="پلازمید" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D0%B8%D0%B4" title="Плазмид – Serbian" lang="sr" hreflang="sr" data-title="Плазмид" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Plazmid" title="Plazmid – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Plazmid" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Plasmidi" title="Plasmidi – Finnish" lang="fi" hreflang="fi" data-title="Plasmidi" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Plasmid" title="Plasmid – Swedish" lang="sv" hreflang="sv" data-title="Plasmid" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tl badge-Q70893996 mw-list-item" title=""><a href="https://tl.wikipedia.org/wiki/Plasmido" title="Plasmido – Tagalog" lang="tl" hreflang="tl" data-title="Plasmido" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target"><span>Tagalog</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%95%E0%AE%A3%E0%AE%BF%E0%AE%AE%E0%AE%BF" title="கணிமி – Tamil" lang="ta" hreflang="ta" data-title="கணிமி" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%9E%E0%B8%A5%E0%B8%B2%E0%B8%AA%E0%B8%A1%E0%B8%B4%E0%B8%94" title="พลาสมิด – Thai" lang="th" hreflang="th" data-title="พลาสมิด" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Plazmid" title="Plazmid – Turkish" lang="tr" hreflang="tr" data-title="Plazmid" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9F%D0%BB%D0%B0%D0%B7%D0%BC%D1%96%D0%B4%D0%B0" title="Плазміда – Ukrainian" lang="uk" hreflang="uk" data-title="Плазміда" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Plasmid" title="Plasmid – Vietnamese" lang="vi" hreflang="vi" data-title="Plasmid" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-vls mw-list-item"><a href="https://vls.wikipedia.org/wiki/Plasmide" title="Plasmide – West Flemish" lang="vls" hreflang="vls" data-title="Plasmide" data-language-autonym="West-Vlams" data-language-local-name="West Flemish" class="interlanguage-link-target"><span>West-Vlams</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E8%B4%A8%E7%B2%92" title="质粒 – Wu" lang="wuu" hreflang="wuu" data-title="质粒" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%B4%A8%E7%B2%92" title="质粒 – Chinese" lang="zh" hreflang="zh" data-title="质粒" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q172778#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div 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.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">This article is about the DNA molecule. For the physics phenomenon, see <a href="/wiki/Plasmoid" title="Plasmoid">plasmoid</a>.</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Plasmid_(english).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Plasmid_%28english%29.svg/300px-Plasmid_%28english%29.svg.png" decoding="async" width="300" height="141" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Plasmid_%28english%29.svg/450px-Plasmid_%28english%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cf/Plasmid_%28english%29.svg/600px-Plasmid_%28english%29.svg.png 2x" data-file-width="320" data-file-height="150" /></a><figcaption>Illustration of a bacterium showing chromosomal DNA and plasmids (Not to scale)</figcaption></figure> <p>A <b>plasmid</b> is a small, <a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">extrachromosomal DNA</a> molecule within a cell that is physically separated from <a href="/wiki/GDNA" class="mw-redirect" title="GDNA">chromosomal DNA</a> and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in <a href="/wiki/Bacteria" title="Bacteria">bacteria</a>; however, plasmids are sometimes present in <a href="/wiki/Archaea" title="Archaea">archaea</a> and <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic organisms</a>.<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><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Plasmids often carry useful genes, such as <a href="/wiki/Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a> and <a href="/wiki/Virulence" title="Virulence">virulence</a>.<sup id="cite_ref-Smillie_2010_3-0" class="reference"><a href="#cite_note-Smillie_2010-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Carattoli_2013_4-0" class="reference"><a href="#cite_note-Carattoli_2013-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_5-0" class="reference"><a href="#cite_note-:1-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> While chromosomes are large and contain all the essential genetic information for living under normal conditions, plasmids are usually very small and contain additional genes for special circumstances. </p><p><a href="/wiki/Artificial_plasmids" class="mw-redirect" title="Artificial plasmids">Artificial plasmids</a> are widely used as <a href="/wiki/Vector_(molecular_biology)" title="Vector (molecular biology)">vectors</a> in <a href="/wiki/Molecular_cloning" title="Molecular cloning">molecular cloning</a>, serving to drive the replication of <a href="/wiki/Recombinant_DNA" title="Recombinant DNA">recombinant DNA</a> sequences within host organisms. In the laboratory, plasmids may be introduced into a cell via <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">transformation</a>. Synthetic plasmids are available for procurement over the internet by various vendors using submitted sequences typically designed with software, if a design does not work the vendor may make additional edits from the submission.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Plasmids are considered <i><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">replicons</a></i>, units of DNA capable of replicating autonomously within a suitable host. However, plasmids, like <a href="/wiki/Virus" title="Virus">viruses</a>, are not generally classified as <a href="/wiki/Life" title="Life">life</a>.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Plasmids are transmitted from one bacterium to another (even of another species) mostly through <a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">conjugation</a>.<sup id="cite_ref-Smillie_2010_3-1" class="reference"><a href="#cite_note-Smillie_2010-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> This host-to-host transfer of genetic material is one mechanism of <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>, and plasmids are considered part of the <a href="/wiki/Mobilome" title="Mobilome">mobilome</a>. Unlike viruses, which encase their genetic material in a protective protein coat called a <a href="/wiki/Capsid" title="Capsid">capsid</a>, plasmids are "naked" DNA and do not encode genes necessary to encase the genetic material for transfer to a new host; however, some classes of plasmids encode the <a href="/wiki/Pilus#Conjugative_pili" title="Pilus">conjugative "sex" pilus</a> necessary for their own transfer. Plasmids vary in size from 1 to over 400 k<a href="/wiki/Base_pair" title="Base pair">bp</a>,<sup id="cite_ref-ThomasSummers2008_10-0" class="reference"><a href="#cite_note-ThomasSummers2008-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> and the number of identical plasmids in a single <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> can range from one up to thousands. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term <i>plasmid</i> was coined in 1952 by the American <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biologist</a> <a href="/wiki/Joshua_Lederberg" title="Joshua Lederberg">Joshua Lederberg</a> to refer to "any extrachromosomal hereditary determinant."<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><sup id="cite_ref-Helinski_2022_12-0" class="reference"><a href="#cite_note-Helinski_2022-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The term's early usage included any bacterial genetic material that exists extrachromosomally for at least part of its replication cycle, but because that description includes bacterial viruses, the notion of plasmid was refined over time to refer to genetic elements that reproduce autonomously.<sup id="cite_ref-Hayes_2003_13-0" class="reference"><a href="#cite_note-Hayes_2003-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Later in 1968, it was decided that the term plasmid should be adopted as the term for extrachromosomal genetic element,<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> and to distinguish it from viruses, the definition was narrowed to genetic elements that exist exclusively or predominantly outside of the chromosome, can replicate autonomously, and contribute to transferring mobile elements between unrelated bacteria.<sup id="cite_ref-Smillie_2010_3-2" class="reference"><a href="#cite_note-Smillie_2010-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Carattoli_2013_4-1" class="reference"><a href="#cite_note-Carattoli_2013-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hayes_2003_13-1" class="reference"><a href="#cite_note-Hayes_2003-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Properties_and_characteristics">Properties and characteristics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=2" title="Edit section: Properties and characteristics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Plasmid_replication_(english).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Plasmid_replication_%28english%29.svg/400px-Plasmid_replication_%28english%29.svg.png" decoding="async" width="400" height="350" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Plasmid_replication_%28english%29.svg/600px-Plasmid_replication_%28english%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/42/Plasmid_replication_%28english%29.svg/800px-Plasmid_replication_%28english%29.svg.png 2x" data-file-width="800" data-file-height="700" /></a><figcaption>There are two types of plasmid integration into a host bacteria: Non-integrating plasmids replicate as with the top instance, whereas <a href="/wiki/Episomes" class="mw-redirect" title="Episomes">episomes</a>, the lower example, can integrate into the host <a href="/wiki/Chromosome" title="Chromosome">chromosome</a>.</figcaption></figure> <p>In order for plasmids to replicate independently within a cell, they must possess a stretch of DNA that can act as an <a href="/wiki/Origin_of_replication" title="Origin of replication">origin of replication</a>. The self-replicating unit, in this case, the plasmid, is called a <a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">replicon</a>. A typical bacterial replicon may consist of a number of elements, such as the gene for plasmid-specific replication initiation protein (Rep), repeating units called <a href="/wiki/Iteron" title="Iteron">iterons</a>, <a href="/wiki/DnaA" title="DnaA">DnaA</a> boxes, and an adjacent AT-rich region.<sup id="cite_ref-Hayes_2003_13-2" class="reference"><a href="#cite_note-Hayes_2003-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Smaller plasmids make use of the host replicative enzymes to make copies of themselves, while larger plasmids may carry genes specific for the replication of those plasmids. A few types of plasmids can also insert into the host chromosome, and these integrative plasmids are sometimes referred to as <a href="/wiki/Episome" title="Episome">episomes</a> in <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>.<sup id="cite_ref-brown_15-0" class="reference"><a href="#cite_note-brown-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Plasmids almost always carry at least one gene. Many of the genes carried by a plasmid are beneficial for the host cells, for example: enabling the host cell to survive in an environment that would otherwise be lethal or restrictive for growth. Some of these genes encode traits for antibiotic resistance or resistance to heavy metal, while others may produce <a href="/wiki/Virulence_factor" title="Virulence factor">virulence factors</a> that enable a bacterium to colonize a host and overcome its defences or have specific metabolic functions that allow the bacterium to utilize a particular nutrient, including the ability to degrade recalcitrant or toxic organic compounds.<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> Plasmids can also provide bacteria with the ability to <a href="/wiki/Nitrogen_fixation" title="Nitrogen fixation">fix nitrogen</a>. Some plasmids, called <a href="/wiki/Cryptic_plasmids" title="Cryptic plasmids">cryptic plasmids</a>, play a crucial role in <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal genes transfer</a>, since they carry <a href="/wiki/Antibiotic-resistance" class="mw-redirect" title="Antibiotic-resistance">antibiotic-resistance</a> genes. Thus they are important factors in spreading resistance, which can result in antibiotic treatment failures. <sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>Naturally occurring plasmids vary greatly in their physical properties. Their size can range from very small mini-plasmids of less than 1-kilobase pairs (kbp) to very large megaplasmids of several megabase pairs (Mbp). At the upper end, little differs between a megaplasmid and a <a href="/wiki/Minichromosome" title="Minichromosome">minichromosome</a>. Plasmids are generally circular, but examples of linear plasmids are also known. These linear plasmids require specialized mechanisms to replicate their ends.<sup id="cite_ref-Hayes_2003_13-3" class="reference"><a href="#cite_note-Hayes_2003-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Plasmids may be present in an individual cell in varying number, ranging from one to several hundreds. The normal number of copies of plasmid that may be found in a single cell is called the <a href="/wiki/Plasmid_copy_number" title="Plasmid copy number">plasmid copy number</a>, and is determined by how the replication initiation is regulated and the size of the molecule. Larger plasmids tend to have lower copy numbers.<sup id="cite_ref-brown_15-1" class="reference"><a href="#cite_note-brown-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Low-copy-number plasmids that exist only as one or a few copies in each bacterium are, upon <a href="/wiki/Cell_division" title="Cell division">cell division</a>, in danger of being lost in one of the segregating bacteria. Such single-copy plasmids have systems that attempt to actively distribute a copy to both daughter cells. These systems, which include the <a href="/wiki/ParABS_system" title="ParABS system">parABS system</a> and <a href="/wiki/ParMRC_system" title="ParMRC system">parMRC system</a>, are often referred to as the <a href="/wiki/Plasmid_partition_system" title="Plasmid partition system">partition system</a> or partition function of a plasmid.<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> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Regulatory_region_of_repBA_gene" title="Regulatory region of repBA gene">Regulatory region of repBA gene</a></div> <p>Plasmids of <i>linear</i> form are unknown among <a href="/wiki/Phytopathogen" class="mw-redirect" title="Phytopathogen">phytopathogens</a> with one exception, <i><a href="/wiki/Rhodococcus_fascians" title="Rhodococcus fascians">Rhodococcus fascians</a></i>.<sup id="cite_ref-Coup-d-Etat_19-0" class="reference"><a href="#cite_note-Coup-d-Etat-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Classifications_and_types">Classifications and types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=3" title="Edit section: Classifications and types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Conjugation.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Conjugation.svg/250px-Conjugation.svg.png" decoding="async" width="250" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Conjugation.svg/375px-Conjugation.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Conjugation.svg/500px-Conjugation.svg.png 2x" data-file-width="833" data-file-height="755" /></a><figcaption>Overview of bacterial conjugation</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:DNA_Under_electron_microscope_Image_3576B-PH.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/DNA_Under_electron_microscope_Image_3576B-PH.jpg/220px-DNA_Under_electron_microscope_Image_3576B-PH.jpg" decoding="async" width="220" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/DNA_Under_electron_microscope_Image_3576B-PH.jpg/330px-DNA_Under_electron_microscope_Image_3576B-PH.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ca/DNA_Under_electron_microscope_Image_3576B-PH.jpg/440px-DNA_Under_electron_microscope_Image_3576B-PH.jpg 2x" data-file-width="5614" data-file-height="3829" /></a><figcaption><a href="/wiki/Electron_micrograph" class="mw-redirect" title="Electron micrograph">Electron micrograph</a> of a DNA fiber bundle, presumably of a single bacterial chromosome loop</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Plasmid_em-en.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Plasmid_em-en.jpg/220px-Plasmid_em-en.jpg" decoding="async" width="220" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Plasmid_em-en.jpg/330px-Plasmid_em-en.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c5/Plasmid_em-en.jpg/440px-Plasmid_em-en.jpg 2x" data-file-width="600" data-file-height="391" /></a><figcaption>Electron micrograph of a bacterial DNA plasmid (chromosome fragment)</figcaption></figure> <p>Plasmids may be classified in a number of ways. Plasmids can be broadly classified into conjugative plasmids and non-conjugative plasmids. Conjugative plasmids contain a set of <a href="/wiki/Transfer_gene" title="Transfer gene">transfer genes</a> which promote sexual conjugation between different cells.<sup id="cite_ref-brown_15-2" class="reference"><a href="#cite_note-brown-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In the complex process of <a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">conjugation</a>, plasmids may be transferred from one bacterium to another via <a href="/wiki/Pilus#Conjugative_pili" title="Pilus">sex pili</a> encoded by some of the transfer genes (see figure).<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> Non-conjugative plasmids are incapable of initiating conjugation, hence they can be transferred only with the assistance of conjugative plasmids. An intermediate class of plasmids are mobilizable, and carry only a subset of the genes required for transfer. They can parasitize a conjugative plasmid, transferring at high frequency only in its presence.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Plasmids can also be classified into incompatibility groups. A microbe can harbour different types of plasmids, but different plasmids can only exist in a single bacterial cell if they are compatible. If two plasmids are not compatible, one or the other will be rapidly lost from the cell. Different plasmids may therefore be assigned to different incompatibility groups depending on whether they can coexist together. Incompatible plasmids (belonging to the same incompatibility group) normally share the same replication or partition mechanisms and can thus not be kept together in a single cell.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>Another way to classify plasmids is by function. There are five main classes: </p> <ul><li>Fertility <a href="/wiki/F-plasmid" title="F-plasmid">F-plasmids</a>, which contain <i>tra</i> genes. They are capable of <a href="/wiki/Bacterial_conjugation" title="Bacterial conjugation">conjugation</a> and result in the expression of <a href="/wiki/Pilus#Conjugative_pili" title="Pilus">sex pili</a>.<sup id="cite_ref-Helinski_2022_12-1" class="reference"><a href="#cite_note-Helinski_2022-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_24-0" class="reference"><a href="#cite_note-:0-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> F-plasmids are categorized as either (+) or (-) and contribute to the difference of being a donor or recipient during conjugation.<sup id="cite_ref-Helinski_2022_12-2" class="reference"><a href="#cite_note-Helinski_2022-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_24-1" class="reference"><a href="#cite_note-:0-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li> <li>Resistance (R) plasmids, which contain genes that provide resistance against <a href="/wiki/Antibiotic" title="Antibiotic">antibiotics</a> or antibacterial agents was first discovered in 1959.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> R-factors where seen as the contributing factor for the spread of <a href="/wiki/Multiple_drug_resistance" title="Multiple drug resistance">multidrug resistance</a> in bacteria, some R-plasmids assist in transmissibility of other specifically non- self transmissible R-factors.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Historically known as R-factors, before the nature of plasmids was understood.</li> <li>Col plasmids, which contain genes that code for <a href="/wiki/Bacteriocin" title="Bacteriocin">bacteriocins</a>, <a href="/wiki/Protein" title="Protein">proteins</a> that can kill other bacteria.</li> <li>Degradative plasmids, which enable the digestion of unusual substances, e.g. <a href="/wiki/Toluene" title="Toluene">toluene</a> and <a href="/wiki/Salicylic_acid" title="Salicylic acid">salicylic acid</a>.</li> <li>Virulence plasmids, which turn the bacterium into a <a href="/wiki/Pathogen" title="Pathogen">pathogen</a>. e.g. <a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti plasmid</a> in <i><a href="/wiki/Agrobacterium_tumefaciens" title="Agrobacterium tumefaciens">Agrobacterium tumefaciens</a>.</i> Bacteria under selective pressure will keep plasmids containing virulence factors as it is a cost - benefit for survival, removal of the selective pressure can lead to the loss of a plasmid due to the expenditure of energy needed to keep it is no longer justified.<sup id="cite_ref-:1_5-1" class="reference"><a href="#cite_note-:1-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup></li></ul> <p>Plasmids can belong to more than one of these functional groups. </p> <div class="mw-heading mw-heading3"><h3 id="RNA_plasmids">RNA plasmids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=4" title="Edit section: RNA plasmids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although most plasmids are double-stranded DNA molecules, some consist of <a href="/wiki/Single-stranded_DNA" class="mw-redirect" title="Single-stranded DNA">single-stranded DNA</a>, or predominantly <a href="/wiki/Double-stranded_RNA" title="Double-stranded RNA">double-stranded RNA</a>. RNA plasmids are non-infectious extrachromosomal linear RNA replicons, both <a href="/wiki/Virus-like_particle" title="Virus-like particle">encapsidated</a> and unencapsidated, which have been found in fungi and various plants, from algae to land plants. In many cases, however, it may be difficult or impossible to clearly distinguish RNA plasmids from RNA viruses and other infectious RNAs.<sup id="cite_ref-Brown_1989_30-0" class="reference"><a href="#cite_note-Brown_1989-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Chromids">Chromids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=5" title="Edit section: Chromids"><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/Chromid" class="mw-redirect" title="Chromid">Chromid</a></div> <p>Chromids are elements that exist at the boundary between a <a href="/wiki/Chromosome" title="Chromosome">chromosome</a> and a plasmid, found in about 10% of bacterial species sequenced by 2009. These elements carry core genes and have <a href="/wiki/Codon_usage" class="mw-redirect" title="Codon usage">codon usage</a> similar to the chromosome, yet use a plasmid-type replication mechanism such as the low copy number RepABC. As a result, they have been variously classified as minichromosomes or megaplasmids in the past.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> In <i><a href="/wiki/Vibrio" title="Vibrio">Vibrio</a></i>, the bacterium synchronizes the replication of the chromosome and chromid by a conserved genome size ratio.<sup id="cite_ref-Bruhn_32-0" class="reference"><a href="#cite_note-Bruhn-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Vectors">Vectors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=6" title="Edit section: Vectors"><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">Further information: <a href="/wiki/Vector_(molecular_biology)" title="Vector (molecular biology)">Vector (molecular biology)</a></div> <p>Artificially constructed plasmids may be used as <a href="/wiki/Vector_(molecular_biology)" title="Vector (molecular biology)">vectors</a> in <a href="/wiki/Genetic_engineering" title="Genetic engineering">genetic engineering</a>. These plasmids serve as important tools in genetics and biotechnology labs, where they are commonly used to clone and amplify (make many copies of) or <a href="/wiki/Gene_expression" title="Gene expression">express</a> particular genes.<sup id="cite_ref-Molecular_cloning_33-0" class="reference"><a href="#cite_note-Molecular_cloning-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> A wide variety of plasmids are commercially available for such uses. The gene to be replicated is normally inserted into a plasmid that typically contains a number of features for their use. These include a gene that confers resistance to particular antibiotics (<a href="/wiki/Ampicillin" title="Ampicillin">ampicillin</a> is most frequently used for bacterial strains), an <a href="/wiki/Origin_of_replication" title="Origin of replication">origin of replication</a> to allow the bacterial cells to replicate the plasmid DNA, and a suitable site for cloning (referred to as a <a href="/wiki/Multiple_cloning_site" title="Multiple cloning site">multiple cloning site</a>). </p><p>DNA structural instability can be defined as a series of spontaneous events that culminate in an unforeseen rearrangement, loss, or gain of genetic material. Such events are frequently triggered by the transposition of mobile elements or by the presence of unstable elements such as non-canonical (non-B) structures. Accessory regions pertaining to the bacterial backbone may engage in a wide range of structural instability phenomena. Well-known catalysts of <a href="/wiki/Genetic_instability" class="mw-redirect" title="Genetic instability">genetic instability</a> include direct, inverted, and tandem repeats, which are known to be conspicuous in a large number of commercially available cloning and expression vectors.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Insertion sequences can also severely impact plasmid function and yield, by leading to <a href="/wiki/Deletion_(genetics)" title="Deletion (genetics)">deletions</a> and rearrangements, activation, <a href="/wiki/Down-regulation" class="mw-redirect" title="Down-regulation">down-regulation</a> or inactivation of neighboring <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Therefore, the reduction or complete elimination of extraneous <a href="/wiki/Noncoding_DNA" class="mw-redirect" title="Noncoding DNA">noncoding</a> backbone sequences would pointedly reduce the propensity for such events to take place, and consequently, the overall recombinogenic potential of the plasmid.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:PBR322.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/PBR322.svg/220px-PBR322.svg.png" decoding="async" width="220" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/PBR322.svg/330px-PBR322.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5b/PBR322.svg/440px-PBR322.svg.png 2x" data-file-width="820" data-file-height="846" /></a><figcaption>A schematic representation of the <a href="/wiki/PBR322" title="PBR322">pBR322</a> plasmid, one of the first plasmids to be used widely as a <a href="/wiki/Cloning_vector" title="Cloning vector">cloning vector</a>. Shown on the plasmid diagram are the genes encoded (<i>amp</i> and <i>tet</i> for <a href="/wiki/Ampicillin" title="Ampicillin">ampicillin</a> and <a href="/wiki/Tetracycline" title="Tetracycline">tetracycline</a> resistance respectively), its origin of replication (<i>ori</i>), and various <a href="/wiki/Restriction_site" title="Restriction site">restriction sites</a> (indicated in blue).</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Cloning">Cloning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=7" title="Edit section: Cloning"><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/Cloning_vector" title="Cloning vector">Cloning vector</a></div> <p>Plasmids are the most-commonly used bacterial cloning vectors.<sup id="cite_ref-uldis_38-0" class="reference"><a href="#cite_note-uldis-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> These cloning vectors contain a site that allows DNA fragments to be inserted, for example a <a href="/wiki/Multiple_cloning_site" title="Multiple cloning site">multiple cloning site</a> or polylinker which has several commonly used <a href="/wiki/Restriction_sites" class="mw-redirect" title="Restriction sites">restriction sites</a> to which DNA fragments may be <a href="/wiki/Ligation_(molecular_biology)" title="Ligation (molecular biology)">ligated</a>. After the gene of interest is inserted, the plasmids are introduced into bacteria by a process called <a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">transformation</a>. These plasmids contain a <a href="/wiki/Selectable_marker" title="Selectable marker">selectable marker</a>, usually an antibiotic resistance gene, which confers on the bacteria an ability to survive and proliferate in a selective growth medium containing the particular antibiotics. The cells after transformation are exposed to the selective media, and only cells containing the plasmid may survive. In this way, the antibiotics act as a filter to select only the bacteria containing the plasmid DNA. The vector may also contain other <a href="/wiki/Marker_gene" title="Marker gene">marker genes</a> or <a href="/wiki/Reporter_gene" title="Reporter gene">reporter genes</a> to facilitate selection of plasmids with cloned inserts. Bacteria containing the plasmid can then be grown in large amounts, harvested, and the plasmid of interest may then be isolated using various methods of <a href="/wiki/Plasmid_preparation" title="Plasmid preparation">plasmid preparation</a>. </p><p>A plasmid cloning vector is typically used to clone DNA fragments of up to 15 <a href="/wiki/Base_pair" title="Base pair">kbp</a>.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> To clone longer lengths of DNA, <a href="/wiki/Lambda_phage" title="Lambda phage">lambda phage</a> with lysogeny genes deleted, <a href="/wiki/Cosmid" title="Cosmid">cosmids</a>, <a href="/wiki/Bacterial_artificial_chromosome" title="Bacterial artificial chromosome">bacterial artificial chromosomes</a>, or <a href="/wiki/Yeast_artificial_chromosome" title="Yeast artificial chromosome">yeast artificial chromosomes</a> are used. </p> <div class="mw-heading mw-heading3"><h3 id="Protein_Production">Protein Production</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=8" title="Edit section: Protein Production"><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/Expression_vector" title="Expression vector">Expression vector</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_insulin_100IU-ml_vial_white_background.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4e/Human_insulin_100IU-ml_vial_white_background.jpg/220px-Human_insulin_100IU-ml_vial_white_background.jpg" decoding="async" width="220" height="338" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4e/Human_insulin_100IU-ml_vial_white_background.jpg/330px-Human_insulin_100IU-ml_vial_white_background.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4e/Human_insulin_100IU-ml_vial_white_background.jpg/440px-Human_insulin_100IU-ml_vial_white_background.jpg 2x" data-file-width="882" data-file-height="1356" /></a><figcaption>Insulin</figcaption></figure> <p>Another major use of plasmids is to make large amounts of proteins. In this case, researchers grow bacteria containing a plasmid harboring the gene of interest. Just as the bacterium produces proteins to confer its antibiotic resistance, it can also be induced to produce large amounts of proteins from the inserted gene. This is a cheap and easy way of mass-producing the protein, for example, utilizing the rapid reproduction of E.coli with a plasmid containing the <a href="/wiki/Insulin" title="Insulin">insulin</a> gene leads to a large production of insulin. <sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Gene_therapy">Gene therapy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=9" title="Edit section: Gene therapy"><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/Vectors_in_gene_therapy" title="Vectors in gene therapy">Vectors in gene therapy</a></div> <p>Plasmids may also be used for gene transfer as a potential treatment in <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a> so that it may express the protein that is lacking in the cells. Some forms of <a href="/wiki/Gene_therapy" title="Gene therapy">gene therapy</a> require the insertion of therapeutic <a href="/wiki/Gene" title="Gene">genes</a> at pre-selected <a href="/wiki/Chromosome" title="Chromosome">chromosomal</a> target sites within the human <a href="/wiki/Genome" title="Genome">genome</a>. Plasmid vectors are one of many approaches that could be used for this purpose. <a href="/wiki/Zinc_finger_nuclease" class="mw-redirect" title="Zinc finger nuclease">Zinc finger nucleases</a> (ZFNs) offer a way to cause a site-specific <a href="/wiki/Double-strand_break" class="mw-redirect" title="Double-strand break">double-strand break</a> to the DNA genome and cause <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a>. Plasmids encoding ZFN could help deliver a therapeutic gene to a specific site so that <a href="/wiki/Cell_damage" title="Cell damage">cell damage</a>, cancer-causing mutations, or an <a href="/wiki/Immune_response" title="Immune response">immune response</a> is avoided.<sup id="cite_ref-Kandavelou_43-0" class="reference"><a href="#cite_note-Kandavelou-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Disease_models">Disease models</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=10" title="Edit section: Disease models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Plasmids were historically used to genetically engineer the embryonic stem cells of rats to create rat genetic disease models. The limited efficiency of plasmid-based techniques precluded their use in the creation of more accurate human cell models. However, developments in <a href="/wiki/Adeno-associated_virus" title="Adeno-associated virus">adeno-associated virus</a> recombination techniques, and <a href="/wiki/Zinc_finger_nucleases" class="mw-redirect" title="Zinc finger nucleases">zinc finger nucleases</a>, have enabled the creation of a new generation of <a href="/wiki/Isogenic_human_disease_models" title="Isogenic human disease models">isogenic human disease models</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Biogenetic_Gene_Cluster_(BGC)"><span id="Biogenetic_Gene_Cluster_.28BGC.29"></span>Biogenetic Gene Cluster (BGC)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=11" title="Edit section: Biogenetic Gene Cluster (BGC)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Plasmids assist in transporting biogenetic gene clusters - a set of gene that contain all the necessary enzymes that lead to the production of special metabolites (formally known as <a href="/wiki/Secondary_metabolite" title="Secondary metabolite">secondary metabolite)</a>.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> A benefit of using plasmids to transfer BGC is demonstrated by using a suitable host that can mass produce specialized metabolites, some of these molecules are able to control microbial population.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Plasmids can contain and express several BGCs with a few plasmids known to be exclusive for transferring BGCs.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> BGC's can also be transfers to the host organism's chromosome, utilizing a plasmid vector, which allows for studies in gene knockout experiments.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> By using plasmids for the uptake of BGCs, microorganisms can gain an advantage as production is not limited to antibiotic resistant biosynthesis genes but the production of <a href="/wiki/Toxin" title="Toxin">toxin</a>s/antitoxins.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Episomes">Episomes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=12" title="Edit section: Episomes"><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/Episome" title="Episome">Episome</a></div> <p>The term <i>episome</i> was introduced by <a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> and <a href="/wiki/%C3%89lie_Wollman" title="Élie Wollman">Élie Wollman</a> in 1958 to refer to extra-chromosomal genetic material that may replicate autonomously or become integrated into the chromosome.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Since the term was introduced, however, its use has changed, as <i>plasmid</i> has become the preferred term for autonomously replicating extrachromosomal DNA. At a 1968 symposium in London some participants suggested that the term <i>episome</i> be abandoned, although others continued to use the term with a shift in meaning.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>Today, some authors use <i>episome</i> in the context of prokaryotes to refer to a plasmid that is capable of integrating into the chromosome. The integrative plasmids may be replicated and stably maintained in a cell through multiple generations, but at some stage, they will exist as an independent plasmid molecule.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> In the context of eukaryotes, the term <i>episome</i> is used to mean a non-integrated extrachromosomal closed circular DNA molecule that may be replicated in the nucleus.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Viruses are the most common examples of this, such as <a href="/wiki/Herpesviridae" title="Herpesviridae">herpesviruses</a>, <a href="/wiki/Adenoviruses" class="mw-redirect" title="Adenoviruses">adenoviruses</a>, and <a href="/wiki/Polyomavirus" class="mw-redirect" title="Polyomavirus">polyomaviruses</a>, but some are plasmids. Other examples include aberrant chromosomal fragments, such as <a href="/wiki/Double_minute" title="Double minute">double minute chromosomes</a>, that can arise during artificial gene amplifications or in pathologic processes (e.g., cancer cell transformation). Episomes in eukaryotes behave similarly to plasmids in prokaryotes in that the DNA is stably maintained and replicated with the host cell. Cytoplasmic viral episomes (as in <a href="/wiki/Poxvirus" class="mw-redirect" title="Poxvirus">poxvirus</a> infections) can also occur. Some episomes, such as herpesviruses, replicate in a <a href="/wiki/Rolling_circle" class="mw-redirect" title="Rolling circle">rolling circle</a> mechanism, similar to <a href="/wiki/Bacteriophage" title="Bacteriophage">bacteriophages</a> (bacterial phage viruses). Others replicate through a bidirectional replication mechanism (<i>Theta type</i> plasmids). In either case, episomes remain physically separate from host cell chromosomes. Several cancer viruses, including <a href="/wiki/Epstein-Barr_virus" class="mw-redirect" title="Epstein-Barr virus">Epstein-Barr virus</a> and <a href="/wiki/Kaposi%27s_sarcoma-associated_herpesvirus" title="Kaposi's sarcoma-associated herpesvirus">Kaposi's sarcoma-associated herpesvirus</a>, are maintained as latent, chromosomally distinct episomes in cancer cells, where the viruses express <a href="/wiki/Oncogenes" class="mw-redirect" title="Oncogenes">oncogenes</a> that promote cancer cell proliferation. In cancers, these episomes passively replicate together with host chromosomes when the cell divides. When these viral episomes initiate <a href="/wiki/Lytic_cycle" title="Lytic cycle">lytic replication</a> to generate multiple virus particles, they generally activate cellular <a href="/wiki/Innate_immunity" class="mw-redirect" title="Innate immunity">innate immunity</a> defense mechanisms that kill the host cell. </p> <div class="mw-heading mw-heading2"><h2 id="Plasmid_maintenance">Plasmid maintenance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=13" title="Edit section: Plasmid maintenance"><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/Addiction_module" title="Addiction module">Addiction module</a></div> <p>Some plasmids or microbial hosts include an <a href="/wiki/Addiction_module" title="Addiction module">addiction system</a> or postsegregational killing system (PSK), such as the <a href="/wiki/Hok/sok_system" title="Hok/sok system">hok/sok</a> (host killing/suppressor of killing) system of plasmid R1 in <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i>.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> This variant produces both a long-lived <a href="/wiki/Poison" title="Poison">poison</a> and a short-lived <a href="/wiki/Antidote" title="Antidote">antidote</a>. Several types of plasmid addiction systems (toxin/ antitoxin, metabolism-based, ORT systems) were described in the <a href="/wiki/Literature" title="Literature">literature</a><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> and used in biotechnical (fermentation) or biomedical (vaccine therapy) applications. Daughter cells that retain a copy of the plasmid survive, while a daughter cell that fails to inherit the plasmid dies or suffers a reduced growth-rate because of the lingering poison from the parent cell. Finally, the overall productivity could be enhanced.<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The productivity of what? Enhanced by what? Is this sentence placed here by mistake? (October 2024)">clarification needed</span></a></i>]</sup> </p><p>In contrast, plasmids used in biotechnology, such as pUC18, pBR322 and derived vectors, hardly ever contain toxin-antitoxin addiction systems, and therefore need to be kept under antibiotic pressure to avoid plasmid loss. </p> <div class="mw-heading mw-heading2"><h2 id="Plasmids_in_nature">Plasmids in nature</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=14" title="Edit section: Plasmids in nature"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Yeast_plasmids">Yeast plasmids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=15" title="Edit section: Yeast plasmids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Yeast" title="Yeast">Yeasts</a> naturally harbour various plasmids. Notable among them are 2 μm plasmids—small circular plasmids often used for <a href="/wiki/Genetic_engineering" title="Genetic engineering">genetic engineering</a> of yeast—and linear pGKL plasmids from <i><a href="/wiki/Kluyveromyces_lactis" title="Kluyveromyces lactis">Kluyveromyces lactis</a></i>, that are responsible for <a href="/w/index.php?title=Killer_phenotypes&action=edit&redlink=1" class="new" title="Killer phenotypes (page does not exist)">killer phenotypes</a>.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p><p>Other types of plasmids are often related to yeast cloning vectors that include: </p> <ul><li><i>Yeast integrative plasmid (YIp)</i>, yeast vectors that rely on integration into the host chromosome for survival and replication, and are usually used when studying the functionality of a solo gene or when the gene is toxic. Also connected with the gene URA3, that codes an enzyme related to the biosynthesis of pyrimidine nucleotides (T, C);</li> <li><i>Yeast Replicative Plasmid (YRp)</i>, which transport a sequence of chromosomal DNA that includes an origin of replication. These plasmids are less stable, as they can be lost during budding.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Plant_mitochondrial_plasmids">Plant mitochondrial plasmids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=16" title="Edit section: Plant mitochondrial plasmids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The mitochondria of many higher plants contain <a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">self-replicating</a>, extra-chromosomal linear or circular DNA molecules which have been considered to be plasmids. These can range from 0.7 kb to 20 kb in size. The plasmids have been generally classified into two categories- circular and linear.<sup id="cite_ref-Gualberto-2014_59-0" class="reference"><a href="#cite_note-Gualberto-2014-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Circular plasmids have been isolated and found in many different plants, with those in <i><a href="/wiki/Vicia_faba" title="Vicia faba">Vicia faba</a> and <a href="/wiki/Chenopodium_album" title="Chenopodium album">Chenopodium album</a></i> being the most studied and whose mechanism of replication is known. The circular plasmids can replicate using the θ model of replication (as in <i>Vicia faba</i>) and through <a href="/wiki/Rolling_circle_replication" title="Rolling circle replication">rolling circle replication</a> (as in <i>C.album</i>).<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> Linear plasmids have been identified in some plant species such as <i><a href="/wiki/Beta_vulgaris" title="Beta vulgaris">Beta vulgaris</a></i>, <i><a href="/wiki/Brassica_napus" class="mw-redirect" title="Brassica napus">Brassica napus</a>, <a href="/wiki/Maize" title="Maize">Zea mays</a></i>, etc. but are rarer than their circular counterparts. </p><p>The function and origin of these plasmids remains largely unknown. It has been suggested that the circular plasmids share a common ancestor, some genes in the mitochondrial plasmid have counterparts in the nuclear DNA suggesting inter-compartment exchange. Meanwhile, the linear plasmids share structural similarities such as invertrons with viral DNA and fungal plasmids, like fungal plasmids they also have low GC content, these observations have led to some hypothesizing that these linear plasmids have viral origins, or have ended up in plant mitochondria through <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a> from pathogenic fungi.<sup id="cite_ref-Gualberto-2014_59-1" class="reference"><a href="#cite_note-Gualberto-2014-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Study_of_plasmids">Study of plasmids</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=17" title="Edit section: Study of plasmids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Plasmid_DNA_extraction">Plasmid DNA extraction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=18" title="Edit section: Plasmid DNA extraction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Plasmids are often used to purify a specific sequence, since they can easily be purified away from the rest of the genome. For their use as vectors, and for <a href="/wiki/Cloning#Molecular_cloning" title="Cloning">molecular cloning</a>, plasmids often need to be isolated. </p><p>There are several methods to <a href="/wiki/Plasmid_preparation" title="Plasmid preparation">isolate plasmid DNA</a> from bacteria, ranging from the plasmid extraction kits (<a href="/wiki/Plasmid_preparation#Preparations_by_size" title="Plasmid preparation">miniprep to the maxiprep or bulkprep)</a>, <a href="/wiki/Alkaline_lysis" title="Alkaline lysis">alkaline lysis</a>, enzymatic lysis, and mechanical lysis .<sup id="cite_ref-Molecular_cloning_33-1" class="reference"><a href="#cite_note-Molecular_cloning-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The former can be used to quickly find out whether the plasmid is correct in any of several bacterial clones. The yield is a small amount of impure plasmid DNA, which is sufficient for analysis by <a href="/wiki/Restriction_digest" title="Restriction digest">restriction digest</a> and for some cloning techniques. </p><p>In the latter, much larger volumes of bacterial suspension are grown from which a maxi-prep can be performed. In essence, this is a scaled-up miniprep followed by additional purification. This results in relatively large amounts (several hundred micrograms) of very pure plasmid DNA. </p><p>Many commercial kits have been created to perform plasmid extraction at various scales, purity, and levels of automation. </p> <div class="mw-heading mw-heading3"><h3 id="Conformations">Conformations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=19" title="Edit section: Conformations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Plasmid DNA may appear in one of five conformations, which (for a given size) run at different speeds in a gel during <a href="/wiki/Agarose_gel_electrophoresis" title="Agarose gel electrophoresis">electrophoresis</a>. The conformations are listed below in order of electrophoretic mobility (speed for a given applied voltage) from slowest to fastest: </p> <ul><li><i><a href="/wiki/Nick_(DNA)" title="Nick (DNA)">Nicked open-circular</a></i> DNA has one strand cut.</li> <li><i>Relaxed circular</i> DNA is fully intact with both strands uncut but has been enzymatically <i>relaxed</i> (supercoils removed). This can be modeled by letting a twisted extension cord unwind and relax and then plugging it into itself.</li> <li><i>Linear</i> DNA has free ends, either because both strands have been cut or because the DNA was linear <i>in vivo</i>. This can be modeled with an electrical extension cord that is not plugged into itself.</li> <li><i><a href="/wiki/DNA_supercoil" title="DNA supercoil">Supercoiled</a></i> (or <i>covalently closed-circular</i>) DNA is fully intact with both strands uncut, and with an integral twist, resulting in a compact form. This can be modeled by twisting an <a href="/wiki/Extension_cord" title="Extension cord">extension cord</a> and then plugging it into itself.</li> <li><i>Supercoiled <a href="/wiki/Denaturation_(biochemistry)" title="Denaturation (biochemistry)">denatured</a></i> DNA is similar to <i>supercoiled DNA</i>, but has unpaired regions that make it slightly less compact; this can result from excessive alkalinity during plasmid preparation.</li></ul> <p>The rate of migration for small linear fragments is directly proportional to the voltage applied at low voltages. At higher voltages, larger fragments migrate at continuously increasing yet different rates. Thus, the resolution of a gel decreases with increased voltage. </p><p>At a specified, low voltage, the migration rate of small linear DNA fragments is a function of their length. Large linear fragments (over 20 kb or so) migrate at a certain fixed rate regardless of length. This is because the molecules 'respirate', with the bulk of the molecule following the leading end through the gel matrix. <a href="/wiki/Restriction_digest" title="Restriction digest">Restriction digests</a> are frequently used to analyse purified plasmids. These enzymes specifically break the DNA at certain short sequences. The resulting linear fragments form 'bands' after <a href="/wiki/Gel_electrophoresis" title="Gel electrophoresis">gel electrophoresis</a>. It is possible to purify certain fragments by cutting the bands out of the gel and dissolving the gel to release the DNA fragments. </p><p>Because of its tight conformation, supercoiled DNA migrates faster through a gel than linear or open-circular DNA. </p> <div class="mw-heading mw-heading3"><h3 id="Software_for_bioinformatics_and_design">Software for bioinformatics and design</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=20" title="Edit section: Software for bioinformatics and design"><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/List_of_genetic_engineering_software" title="List of genetic engineering software">List of genetic engineering software</a></div> <p>The use of plasmids as a technique in <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a> is supported by <a href="/wiki/Bioinformatics" title="Bioinformatics">bioinformatics</a> <a href="/wiki/Software" title="Software">software</a>. These programs record the <a href="/wiki/DNA" title="DNA">DNA</a> sequence of plasmid vectors, help to predict cut sites of <a href="/wiki/Restriction_enzymes" class="mw-redirect" title="Restriction enzymes">restriction enzymes</a>, and to plan manipulations. Examples of software packages that handle plasmid maps are ApE, <a href="/wiki/Clone_manager" title="Clone manager">Clone Manager</a>, GeneConstructionKit, Geneious, <a href="/wiki/Genome_Compiler" class="mw-redirect" title="Genome Compiler">Genome Compiler</a>, LabGenius, Lasergene, <a href="/wiki/MacVector" title="MacVector">MacVector</a>, pDraw32, Serial Cloner, <a href="/wiki/UGENE" title="UGENE">UGENE</a>, VectorFriends, <a href="/wiki/Vector_NTI" title="Vector NTI">Vector NTI</a>, and WebDSV. These pieces of software help conduct entire experiments in silico before doing wet experiments.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Plasmid_collections">Plasmid collections</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=21" title="Edit section: Plasmid collections"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many plasmids have been created over the years and researchers have given out plasmids to plasmid databases such as the non-profit organisations <a rel="nofollow" class="external text" href="https://www.addgene.org">Addgene</a> and <a rel="nofollow" class="external text" href="https://bccm.belspo.be/about-GeneCorner">BCCM/GeneCorner</a>. One can find and request plasmids from those databases for research. Researchers also often upload plasmid sequences to the <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/nuccore/">NCBI database</a>, from which sequences of specific plasmids can be retrieved. </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=Plasmid&action=edit&section=22" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 30em;"> <ul><li><a href="/wiki/Bacterial_artificial_chromosome" title="Bacterial artificial chromosome">Bacterial artificial chromosome</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Recombinant_DNA" title="Recombinant DNA">DNA recombination</a></li> <li><a href="/wiki/Plasmidome" title="Plasmidome">Plasmidome</a></li> <li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li> <li><a href="/wiki/Segrosome" title="Segrosome">Segrosome</a></li> <li><a href="/wiki/Transposon" class="mw-redirect" title="Transposon">Transposon</a></li> <li><a href="/wiki/Triparental_mating" title="Triparental mating">Triparental mating</a></li> <li><a href="/wiki/VectorDB" title="VectorDB">VectorDB</a></li></ul></div> <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=Plasmid&action=edit&section=23" 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 data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFEsserKückLang-HinrichsLemke1986" class="citation book cs1">Esser K, Kück U, Lang-Hinrichs C, Lemke P, Osiewacz HD, Stahl U, et al. (1986). <i>Plasmids of Eukaryotes: fundamentals and Applications</i>. 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(identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/7045080">7045080</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+Bacteriology&rft.atitle=Transformation+of+Saccharomyces+cerevisiae+with+linear+DNA+killer+plasmids+from+Kluyveromyces+lactis&rft.volume=151&rft.issue=1&rft.pages=462-464&rft.date=1982-07&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC220260%23id-name%3DPMC&rft_id=info%3Apmid%2F7045080&rft_id=info%3Adoi%2F10.1128%2FJB.151.1.462-464.1982&rft.aulast=Gunge&rft.aufirst=N&rft.au=Murata%2C+K&rft.au=Sakaguchi%2C+K&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC220260&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></span> </li> <li id="cite_note-Gualberto-2014-59"><span class="mw-cite-backlink">^ <a href="#cite_ref-Gualberto-2014_59-0"><sup><i><b>a</b></i></sup></a> <a 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"The plant mitochondrial genome: dynamics and maintenance". <i>Biochimie</i>. <b>100</b>: 107–120. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.biochi.2013.09.016">10.1016/j.biochi.2013.09.016</a>. <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/24075874">24075874</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biochimie&rft.atitle=The+plant+mitochondrial+genome%3A+dynamics+and+maintenance&rft.volume=100&rft.pages=107-120&rft.date=2014-05&rft_id=info%3Adoi%2F10.1016%2Fj.biochi.2013.09.016&rft_id=info%3Apmid%2F24075874&rft.aulast=Gualberto&rft.aufirst=JM&rft.au=Mileshina%2C+D&rft.au=Wallet%2C+C&rft.au=Niazi%2C+AK&rft.au=Weber-Lotfi%2C+F&rft.au=Dietrich%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></span> </li> <li id="cite_note-60"><span class="mw-cite-backlink"><b><a href="#cite_ref-60">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBackertMeissnerBörner1997" class="citation journal cs1">Backert S, Meissner K, Börner T (February 1997). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC146482">"Unique features of the mitochondrial rolling circle-plasmid mp1 from the higher plant Chenopodium album (L.)"</a>. <i>Nucleic Acids Research</i>. <b>25</b> (3): 582–589. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fnar%2F25.3.582">10.1093/nar/25.3.582</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/PMC146482">146482</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/9016599">9016599</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nucleic+Acids+Research&rft.atitle=Unique+features+of+the+mitochondrial+rolling+circle-plasmid+mp1+from+the+higher+plant+Chenopodium+album+%28L.%29&rft.volume=25&rft.issue=3&rft.pages=582-589&rft.date=1997-02&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC146482%23id-name%3DPMC&rft_id=info%3Apmid%2F9016599&rft_id=info%3Adoi%2F10.1093%2Fnar%2F25.3.582&rft.aulast=Backert&rft.aufirst=S&rft.au=Meissner%2C+K&rft.au=B%C3%B6rner%2C+T&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC146482&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></span> </li> <li id="cite_note-61"><span class="mw-cite-backlink"><b><a href="#cite_ref-61">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHanda2008" class="citation journal cs1">Handa H (January 2008). "Linear plasmids in plant mitochondria: peaceful coexistences or malicious invasions?". <i>Mitochondrion</i>. <b>8</b> (1): 15–25. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.mito.2007.10.002">10.1016/j.mito.2007.10.002</a>. <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/18326073">18326073</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Mitochondrion&rft.atitle=Linear+plasmids+in+plant+mitochondria%3A+peaceful+coexistences+or+malicious+invasions%3F&rft.volume=8&rft.issue=1&rft.pages=15-25&rft.date=2008-01&rft_id=info%3Adoi%2F10.1016%2Fj.mito.2007.10.002&rft_id=info%3Apmid%2F18326073&rft.aulast=Handa&rft.aufirst=H&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></span> </li> <li id="cite_note-62"><span class="mw-cite-backlink"><b><a href="#cite_ref-62">^</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://vimeo.com/57923864">"Vector NTI feedback video"</a>. <i>The DNA Lab</i>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=The+DNA+Lab&rft.atitle=Vector+NTI+feedback+video&rft_id=http%3A%2F%2Fvimeo.com%2F57923864&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=24" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="General_works">General works</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=25" title="Edit section: General works"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKleinPrescottHarley1999" class="citation book cs1">Klein DW, Prescott LM, Harley J (1999). <i>Microbiology</i>. Boston: WCB/McGraw-Hill.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Microbiology&rft.place=Boston&rft.pub=WCB%2FMcGraw-Hill&rft.date=1999&rft.aulast=Klein&rft.aufirst=DW&rft.au=Prescott%2C+LM&rft.au=Harley%2C+J&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoatFosterSpector2002" class="citation book cs1">Moat AG, Foster JW, Spector MP (2002). <i>Microbial Physiology</i>. Wiley-Liss. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-471-39483-9" title="Special:BookSources/978-0-471-39483-9"><bdi>978-0-471-39483-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Microbial+Physiology&rft.pub=Wiley-Liss&rft.date=2002&rft.isbn=978-0-471-39483-9&rft.aulast=Moat&rft.aufirst=AG&rft.au=Foster%2C+JW&rft.au=Spector%2C+MP&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSmith2002" class="citation book cs1">Smith CU (2002). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=wey2dH3D0SkC&pg=PA110">"Chapter 5: Manipulating Biomolecules"</a>. <i>Elements of Molecular Neurobiology</i> (3rd ed.). Chichester, West Sussex, England: Wiley. pp. 101–11. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-470-85717-5" title="Special:BookSources/978-0-470-85717-5"><bdi>978-0-470-85717-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+5%3A+Manipulating+Biomolecules&rft.btitle=Elements+of+Molecular+Neurobiology&rft.place=Chichester%2C+West+Sussex%2C+England&rft.pages=101-11&rft.edition=3rd&rft.pub=Wiley&rft.date=2002&rft.isbn=978-0-470-85717-5&rft.aulast=Smith&rft.aufirst=CU&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dwey2dH3D0SkC%26pg%3DPA110&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading3"><h3 id="Episomes_2">Episomes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=26" title="Edit section: Episomes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239549316"><div class="refbegin refbegin-columns references-column-width" style="column-width: 32em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPiechaczekFetzerBaikerBode1999" class="citation journal cs1">Piechaczek C, Fetzer C, Baiker A, Bode J, Lipps HJ (January 1999). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC148196">"A vector based on the SV40 origin of replication and chromosomal S/MARs replicates episomally in CHO cells"</a>. <i>Nucleic Acids Research</i>. <b>27</b> (2): 426–428. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fnar%2F27.2.426">10.1093/nar/27.2.426</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/PMC148196">148196</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/9862961">9862961</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nucleic+Acids+Research&rft.atitle=A+vector+based+on+the+SV40+origin+of+replication+and+chromosomal+S%2FMARs+replicates+episomally+in+CHO+cells&rft.volume=27&rft.issue=2&rft.pages=426-428&rft.date=1999-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC148196%23id-name%3DPMC&rft_id=info%3Apmid%2F9862961&rft_id=info%3Adoi%2F10.1093%2Fnar%2F27.2.426&rft.aulast=Piechaczek&rft.aufirst=C&rft.au=Fetzer%2C+C&rft.au=Baiker%2C+A&rft.au=Bode%2C+J&rft.au=Lipps%2C+HJ&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC148196&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBodeFetzerNehlsenScinteie2001" class="citation journal cs1">Bode J, Fetzer CP, Nehlsen K, Scinteie M, Hinrichsen BH, Baiker A, et al. (January 2001). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20090530141241/http://www.gtmb.org/VOL6A/GTMBVOL6APDF/INDIVIDUAL/03%20%20Bode.pdf">"The Hitchhiking principle: Optimizing episomal vectors for the use in gene therapy and biotechnology"</a> <span class="cs1-format">(PDF)</span>. <i>Gene Therapy and Molecular Biology</i>. <b>6</b>: 33–46. Archived from <a rel="nofollow" class="external text" href="http://www.gtmb.org/VOL6A/GTMBVOL6APDF/INDIVIDUAL/03%20%20Bode.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 30 May 2009.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Gene+Therapy+and+Molecular+Biology&rft.atitle=The+Hitchhiking+principle%3A+Optimizing+episomal+vectors+for+the+use+in+gene+therapy+and+biotechnology&rft.volume=6&rft.pages=33-46&rft.date=2001-01&rft.aulast=Bode&rft.aufirst=J&rft.au=Fetzer%2C+CP&rft.au=Nehlsen%2C+K&rft.au=Scinteie%2C+M&rft.au=Hinrichsen%2C+BH&rft.au=Baiker%2C+A&rft.au=Piechazcek%2C+C&rft.au=Benham%2C+C&rft.au=Lipps%2C+HJ&rft_id=http%3A%2F%2Fwww.gtmb.org%2FVOL6A%2FGTMBVOL6APDF%2FINDIVIDUAL%2F03%2520%2520Bode.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNehlsenBrollBode2006" class="citation journal cs1">Nehlsen K, Broll S, Bode J (2006). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20090530141235/http://www.gtmb.org/VOL10B/INDIVIDUAL/25.%20Nehlsen%20et%20al%2C%20233-244.pdf">"Replicating minicircles: Generation of nonviral episomes for the efficient modification of dividing cells"</a> <span class="cs1-format">(PDF)</span>. <i>Gene Ther Mol Biol</i>. <b>10</b>: 233–44. Archived from <a rel="nofollow" class="external text" href="http://www.gtmb.org/VOL10B/INDIVIDUAL/25.%20Nehlsen%20et%20al,%20233-244.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 30 May 2009.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Gene+Ther+Mol+Biol&rft.atitle=Replicating+minicircles%3A+Generation+of+nonviral+episomes+for+the+efficient+modification+of+dividing+cells&rft.volume=10&rft.pages=233-44&rft.date=2006&rft.aulast=Nehlsen&rft.aufirst=K&rft.au=Broll%2C+S&rft.au=Bode%2C+J&rft_id=http%3A%2F%2Fwww.gtmb.org%2FVOL10B%2FINDIVIDUAL%2F25.%2520Nehlsen%2520et%2520al%2C%2520233-244.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEhrhardtHaaseSchepersDeutsch2008" class="citation journal cs1">Ehrhardt A, Haase R, Schepers A, Deutsch MJ, Lipps HJ, Baiker A (June 2008). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110926182142/http://www.benthamdirect.org/pages/content.php?CGT%2F2008%2F00000008%2F00000003%2F0001Q.SGM">"Episomal vectors for gene therapy"</a>. <i>Current Gene Therapy</i>. <b>8</b> (3): 147–161. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2174%2F156652308784746440">10.2174/156652308784746440</a>. <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/18537590">18537590</a>. Archived from <a rel="nofollow" class="external text" href="http://www.benthamdirect.org/pages/content.php?CGT/2008/00000008/00000003/0001Q.SGM">the original</a> on 26 September 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Gene+Therapy&rft.atitle=Episomal+vectors+for+gene+therapy&rft.volume=8&rft.issue=3&rft.pages=147-161&rft.date=2008-06&rft_id=info%3Adoi%2F10.2174%2F156652308784746440&rft_id=info%3Apmid%2F18537590&rft.aulast=Ehrhardt&rft.aufirst=A&rft.au=Haase%2C+R&rft.au=Schepers%2C+A&rft.au=Deutsch%2C+MJ&rft.au=Lipps%2C+HJ&rft.au=Baiker%2C+A&rft_id=http%3A%2F%2Fwww.benthamdirect.org%2Fpages%2Fcontent.php%3FCGT%2F2008%2F00000008%2F00000003%2F0001Q.SGM&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArgyrosWongNicetaWaddington2008" class="citation journal cs1">Argyros O, Wong SP, Niceta M, Waddington SN, Howe SJ, Coutelle C, et al. (December 2008). <a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fgt.2008.113">"Persistent episomal transgene expression in liver following delivery of a scaffold/matrix attachment region containing non-viral vector"</a>. <i>Gene Therapy</i>. <b>15</b> (24): 1593–1605. <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/10.1038%2Fgt.2008.113">10.1038/gt.2008.113</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/18633447">18633447</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Gene+Therapy&rft.atitle=Persistent+episomal+transgene+expression+in+liver+following+delivery+of+a+scaffold%2Fmatrix+attachment+region+containing+non-viral+vector&rft.volume=15&rft.issue=24&rft.pages=1593-1605&rft.date=2008-12&rft_id=info%3Adoi%2F10.1038%2Fgt.2008.113&rft_id=info%3Apmid%2F18633447&rft.aulast=Argyros&rft.aufirst=O&rft.au=Wong%2C+SP&rft.au=Niceta%2C+M&rft.au=Waddington%2C+SN&rft.au=Howe%2C+SJ&rft.au=Coutelle%2C+C&rft.au=Miller%2C+AD&rft.au=Harbottle%2C+RP&rft_id=https%3A%2F%2Fdoi.org%2F10.1038%252Fgt.2008.113&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWongArgyrosCoutelleHarbottle2009" class="citation journal cs1">Wong SP, Argyros O, Coutelle C, Harbottle RP (August 2009). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110917140204/http://www.biomedcentral.com/1464-8431/11/433/abstract">"Strategies for the episomal modification of cells"</a>. <i>Current Opinion in Molecular Therapeutics</i>. <b>11</b> (4): 433–441. <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/19649988">19649988</a>. Archived from <a rel="nofollow" class="external text" href="http://www.biomedcentral.com/1464-8431/11/433/abstract">the original</a> on 17 September 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Opinion+in+Molecular+Therapeutics&rft.atitle=Strategies+for+the+episomal+modification+of+cells&rft.volume=11&rft.issue=4&rft.pages=433-441&rft.date=2009-08&rft_id=info%3Apmid%2F19649988&rft.aulast=Wong&rft.aufirst=SP&rft.au=Argyros%2C+O&rft.au=Coutelle%2C+C&rft.au=Harbottle%2C+RP&rft_id=http%3A%2F%2Fwww.biomedcentral.com%2F1464-8431%2F11%2F433%2Fabstract&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHaaseArgyrosWongHarbottle2010" class="citation journal cs1">Haase R, Argyros O, Wong SP, Harbottle RP, Lipps HJ, Ogris M, et al. (March 2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2847955">"pEPito: a significantly improved non-viral episomal expression vector for mammalian cells"</a>. <i>BMC Biotechnology</i>. <b>10</b>: 20. <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/10.1186%2F1472-6750-10-20">10.1186/1472-6750-10-20</a></span>. <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/PMC2847955">2847955</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/20230618">20230618</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BMC+Biotechnology&rft.atitle=pEPito%3A+a+significantly+improved+non-viral+episomal+expression+vector+for+mammalian+cells&rft.volume=10&rft.pages=20&rft.date=2010-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2847955%23id-name%3DPMC&rft_id=info%3Apmid%2F20230618&rft_id=info%3Adoi%2F10.1186%2F1472-6750-10-20&rft.aulast=Haase&rft.aufirst=R&rft.au=Argyros%2C+O&rft.au=Wong%2C+SP&rft.au=Harbottle%2C+RP&rft.au=Lipps%2C+HJ&rft.au=Ogris%2C+M&rft.au=Magnusson%2C+T&rft.au=Vizoso+Pinto%2C+MG&rft.au=Haas%2C+J&rft.au=Baiker%2C+A&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2847955&rfr_id=info%3Asid%2Fen.wikipedia.org%3APlasmid" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Plasmid&action=edit&section=27" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://www.ispb.org/">International Society for Plasmid Biology and other Mobile Genetic Elements</a></li> <li><a rel="nofollow" class="external text" href="https://www.whatisbiotechnology.org/index.php/science/summary/plasmid/">What is Biotechnology</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090310010107/http://histmicro.yale.edu/mainfram.htm">History of Plasmids with timeline</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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style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></li> <li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> <ul><li><a href="/wiki/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li><a href="/wiki/Protist" title="Protist">Protista</a></li></ul></li> <li><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a> <ul><li><a href="/wiki/Giant_virus" title="Giant virus">Giant virus</a></li></ul></li> <li><a href="/wiki/DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li> <li><a href="/wiki/Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li> <li><a href="/wiki/RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li> <li><a href="/wiki/Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li> <li><a href="/wiki/DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="/wiki/Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br />agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="/wiki/Helper_virus" title="Helper virus">Helper-virus<br />dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="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%;text-align: center;;background:#CEDAF2;"><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</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>ssRNA satellite virus</li> <li>dsDNA satellite virus (<a href="/wiki/Virophage" title="Virophage">Virophage</a>)</li> <li>ssDNA satellite virus</li> <li>ssDNA satellite</li> <li>dsRNA satellite</li> <li>ssRNA satellite (<a href="/wiki/Virusoid" title="Virusoid">Virusoid</a>)</li> <li>Satellite-like nucleic acids <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</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/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a> <ul><li>RNA</li> <li>DNA</li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="/wiki/Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Prion" title="Prion">Mammalian prion</a></li> <li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br />self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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:7em;text-align: center;"><a href="/wiki/Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br />elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mobilome" title="Mobilome">Mobilome</a> <ul><li><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li> <li><a href="/wiki/Genomic_island" title="Genomic island">Genomic island</a></li></ul></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li> <li><a href="/wiki/DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li> <li><a class="mw-selflink selflink">Plasmid</a> <ul><li><a href="/wiki/Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li> <li><a href="/wiki/R-factor" class="mw-redirect" title="R-factor">Resistance</a></li> <li><a href="/wiki/Colicin" title="Colicin">Col</a></li> <li>Degradative</li> <li><a href="/wiki/Virulence_factor" title="Virulence factor">Virulence</a>/<a href="/wiki/Ti_plasmid" title="Ti plasmid">Ti</a></li> <li>Cryptic</li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a> <ul><li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li></ul></li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a href="/wiki/Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li> <li><a href="/wiki/Group_II_intron" title="Group II intron">Group II intron</a></li> <li><a href="/wiki/Retrozyme" title="Retrozyme">Retrozyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li> <li><a href="/wiki/Chromosome" title="Chromosome">Chromosome</a> <ul><li><a href="/wiki/Linear_chromosome" title="Linear chromosome">Linear</a></li> <li><a href="/wiki/Circular_chromosome" title="Circular chromosome">Circular</a></li> <li><a href="/wiki/Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li> <li><a href="/wiki/Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li> <li><a href="/wiki/Genome" title="Genome">Genome</a> <ul><li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Gene_duplication" title="Gene duplication">Gene duplication</a></li> <li><a href="/wiki/Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li> <li><a href="/wiki/Origin_of_replication" title="Origin of replication">Origin of replication</a> <ul><li><a href="/wiki/Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a> <ul><li><a href="/wiki/Provirus" title="Provirus">Provirus</a></li> <li><a href="/wiki/Prophage" title="Prophage">Prophage</a></li> <li><a href="/wiki/Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li> <li><a href="/wiki/Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a> <ul><li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a> <ul><li><a href="/wiki/Mitosome" title="Mitosome">Mitosome</a></li> <li><a href="/wiki/Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Kappa_organism" title="Kappa organism">Kappa organism</a></li> <li>Organs <ul><li><a href="/wiki/Bacteriome" title="Bacteriome">Bacteriome</a></li> <li><a href="/wiki/Trophosome" title="Trophosome">Trophosome</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li>?<a href="/wiki/RNA_world" title="RNA world">RNA life</a> <ul><li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li> <li>†<a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Coacervate" title="Coacervate">Coacervate</a></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a></li> <li><a href="/wiki/Sulphobes" title="Sulphobes">Sulphobe</a></li> <li>Research <ul><li><a href="/wiki/Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Organism" title="Organism">Organism</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a> <ul><li><a href="/wiki/Cell_division" title="Cell division">Cell division</a></li> <li><a href="/wiki/Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li> <li><a href="/wiki/Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li> <li><a href="/wiki/Synthetic_virology" title="Synthetic virology">Synthetic virus</a> <ul><li><a href="/wiki/Viral_vector" title="Viral vector">Viral vector</a></li> <li><a href="/wiki/Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li> <li>?<a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></li> <li>?<a href="/wiki/Nanobe" title="Nanobe">Nanobe</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Virome" title="Virome">Virome</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Life,_non-cellular_life,_and_comparable_structures" title="Template:Life, non-cellular life, and comparable structures"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life,_non-cellular_life,_and_comparable_structures" title="Template talk:Life, non-cellular life, and comparable structures"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life,_non-cellular_life,_and_comparable_structures" title="Special:EditPage/Template:Life, non-cellular life, and comparable structures"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Life,_non-cellular_life,_and_comparable_structures" style="font-size:114%;margin:0 4em"><a href="/wiki/Life" title="Life">Life</a>, <a href="/wiki/Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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/Prokaryote" title="Prokaryote">Prokaryota</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/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteria" title="Bacteria">Bacteria</a> <ul><li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a></li></ul></li> <li><a href="/wiki/LUCA" class="mw-redirect" title="LUCA">LUCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</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/Animal" title="Animal">Animalia</a></li> <li><a href="/wiki/Fungus" title="Fungus">Fungi</a></li> <li><a href="/wiki/Plant" title="Plant">Plantae</a></li> <li>'<a href="/wiki/Protist" title="Protist">Protista</a>'</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="/wiki/Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Parakaryon" title="Parakaryon">Parakaryon</a></i></li> <li><a href="/wiki/Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" 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/Virus" title="Virus">Virus</a> <hr /><a href="/wiki/Viroid" title="Viroid">Viroid</a> <hr /><a href="/wiki/Satellite_(biology)" title="Satellite (biology)">Satellite</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%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Adnaviria" title="Adnaviria">Adnaviria</a></i></li> <li><i><a href="/wiki/Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i></li> <li><i><a href="/wiki/Monodnaviria" title="Monodnaviria">Monodnaviria</a></i></li> <li><i><a href="/wiki/Riboviria" title="Riboviria">Riboviria</a></i></li> <li><i><a href="/wiki/Ribozyviria" title="Ribozyviria">Ribozyviria</a></i></li> <li><i><a href="/wiki/Varidnaviria" title="Varidnaviria">Varidnaviria</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</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%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><i><a href="/wiki/Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></i></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></i></li> <li><i><a href="/wiki/Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></i></li> <li><i><a href="/wiki/Anelloviridae" title="Anelloviridae">Anelloviridae</a></i></li> <li><i><a href="/wiki/Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li> <li><i><a href="/wiki/Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></i></li> <li><i><a href="/wiki/Clavaviridae" title="Clavaviridae">Clavaviridae</a></i></li> <li><i><a href="/wiki/Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></i></li> <li><i><a href="/wiki/Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></i></li> <li><i><a href="/wiki/Globuloviridae" title="Globuloviridae">Globuloviridae</a></i></li> <li><i><a href="/wiki/Guttaviridae" title="Guttaviridae">Guttaviridae</a></i></li> <li><i><a href="/wiki/Halspiviridae" title="Halspiviridae">Halspiviridae</a></i></li> <li><i><a href="/wiki/Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></i></li> <li><i><a href="/wiki/Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></i></li> <li><i><a href="/wiki/Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></i></li> <li><i><a href="/wiki/Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></i></li> <li><i><a href="/wiki/Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li> <li><i><a href="/wiki/Spiraviridae" title="Spiraviridae">Spiraviridae</a></i></li> <li><i><a href="/wiki/Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></i></li> <li><i><a href="/wiki/Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></i></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></i></li> <li><i><a href="/wiki/Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></i></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></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/Nanobe" title="Nanobe">Nanobe</a> (?)</li> <li><a href="/wiki/Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable<br />structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li> <li><a href="/wiki/Cancer_cell" title="Cancer cell">Cancer cell</a> <ul><li><a href="/wiki/HeLa" title="HeLa">HeLa</a></li> <li><a href="/wiki/Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li> <li><a href="/wiki/Cosmid" title="Cosmid">Cosmid</a></li> <li><a href="/wiki/Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li> <li><a href="/wiki/Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li> <li><a href="/wiki/Fosmid" title="Fosmid">Fosmid</a></li> <li><a href="/wiki/Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li> <li><a href="/wiki/Jeewanu" title="Jeewanu">Jeewanu</a></li> <li>"<i><a href="/wiki/Nanobacterium" title="Nanobacterium">Nanobacterium</a></i>"</li> <li><a href="/wiki/Phagemid" title="Phagemid">Phagemid</a></li> <li><a class="mw-selflink selflink">Plasmid</a></li> <li><a href="/wiki/Prion" title="Prion">Prion</a> <ul><li><a href="/wiki/Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li> <li><a href="/wiki/Proteinoid" title="Proteinoid">Proteinoid</a> <a href="/wiki/Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li> <li>Retroelements not elsewhere classified <ul><li><a href="/wiki/Retron" title="Retron">Retron</a></li> <li><a href="/wiki/Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li> <li><a href="/wiki/Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li> <li><a href="/wiki/Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li> <li><a href="/wiki/Ribozyme" title="Ribozyme">Ribozyme</a></li> <li><a href="/wiki/Spiegelman%27s_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li> <li><a href="/wiki/Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li> <li><a href="/wiki/Transposable_element" title="Transposable element">Transposable element</a> <ul><li><a href="/wiki/Retroposon" title="Retroposon">Retroposon</a></li></ul></li> <li><a href="/wiki/Transpoviron" title="Transpoviron">Transpoviron</a></li> <li><a href="/wiki/Xenobot" title="Xenobot">Xenobot</a></li></ul> </div></td></tr></tbody></table></div> <style data-mw-deduplicate="TemplateStyles:r1130092004">.mw-parser-output .portal-bar{font-size:88%;font-weight:bold;display:flex;justify-content:center;align-items:baseline}.mw-parser-output .portal-bar-bordered{padding:0 2em;background-color:#fdfdfd;border:1px solid #a2a9b1;clear:both;margin:1em auto 0}.mw-parser-output .portal-bar-related{font-size:100%;justify-content:flex-start}.mw-parser-output .portal-bar-unbordered{padding:0 1.7em;margin-left:0}.mw-parser-output .portal-bar-header{margin:0 1em 0 0.5em;flex:0 0 auto;min-height:24px}.mw-parser-output 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