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</li> <li id="toc-Infectious_disease_applications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Infectious_disease_applications"> <div class="vector-toc-text"> 3.4 Infectious disease applications </div> </a> <ul id="toc-Infectious_disease_applications-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Forensic_applications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Forensic_applications"> <div class="vector-toc-text"> 3.5 Forensic applications </div> </a> <ul id="toc-Forensic_applications-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Research_applications" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Research_applications"> <div class="vector-toc-text"> 3.6 Research applications </div> </a> <ul id="toc-Research_applications-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Advantages" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Advantages"> <div class="vector-toc-text"> 4 Advantages </div> </a> <ul id="toc-Advantages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Limitations" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Limitations"> <div class="vector-toc-text"> 5 Limitations </div> </a> <ul id="toc-Limitations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Variations" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Variations"> <div class="vector-toc-text"> 6 Variations </div> </a> <ul id="toc-Variations-sublist" class="vector-toc-list"> </ul> </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"> 7 History </div> </a> <button aria-controls="toc-History-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle History subsection </button> <ul id="toc-History-sublist" class="vector-toc-list"> <li id="toc-Patent_disputes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Patent_disputes"> <div class="vector-toc-text"> 7.1 Patent disputes </div> </a> <ul id="toc-Patent_disputes-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"> 8 See also </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"> 9 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 10 External links </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" title="Table of Contents" > <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" > Toggle the table of contents </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">Polymerase chain reaction</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 71 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-71" aria-hidden="true" > 71 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Polimerase_kettingreaksiemetode" title="Polimerase kettingreaksiemetode – Afrikaans" lang="af" hreflang="af" data-title="Polimerase kettingreaksiemetode" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D9%81%D8%A7%D8%B9%D9%84_%D8%A7%D9%84%D8%A8%D9%88%D9%84%D9%8A%D9%85%D8%B1%D8%A7%D8%B2_%D8%A7%D9%84%D9%85%D8%AA%D8%B3%D9%84%D8%B3%D9%84" title="تفاعل البوليمراز المتسلسل – Arabic" lang="ar" hreflang="ar" data-title="تفاعل البوليمراز المتسلسل" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/PCR" title="PCR – Aragonese" lang="an" hreflang="an" data-title="PCR" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Reaici%C3%B3n_en_cadena_de_la_polimerasa" title="Reaición en cadena de la polimerasa – Asturian" lang="ast" hreflang="ast" data-title="Reaición en cadena de la polimerasa" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Polimeraz_z%C9%99ncir_reaksiyas%C4%B1" title="Polimeraz zəncir reaksiyası – Azerbaijani" lang="az" hreflang="az" data-title="Polimeraz zəncir reaksiyası" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AA%E0%A6%B2%E0%A6%BF%E0%A6%AE%E0%A6%BE%E0%A6%B0%E0%A7%87%E0%A6%9C_%E0%A6%B6%E0%A7%83%E0%A6%99%E0%A7%8D%E0%A6%96%E0%A6%B2_%E0%A6%AC%E0%A6%BF%E0%A6%95%E0%A7%8D%E0%A6%B0%E0%A6%BF%E0%A6%AF%E0%A6%BC%E0%A6%BE" title="পলিমারেজ শৃঙ্খল বিক্রিয়া – Bangla" lang="bn" hreflang="bn" data-title="পলিমারেজ শৃঙ্খল বিক্রিয়া" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/To-th%C3%A9-m%C3%BBi_li%C3%A2n-s%C3%B3_ho%C3%A1n-%C3%A8ng" title="To-thé-mûi liân-só hoán-èng – Minnan" lang="nan" hreflang="nan" data-title="To-thé-mûi liân-só hoán-èng" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%9F%D0%B0%D0%BB%D1%96%D0%BC%D1%8D%D1%80%D0%B0%D0%B7%D0%BD%D0%B0%D1%8F_%D0%BB%D0%B0%D0%BD%D1%86%D1%83%D0%B3%D0%BE%D0%B2%D0%B0%D1%8F_%D1%80%D1%8D%D0%B0%D0%BA%D1%86%D1%8B%D1%8F" title="Палімэразная ланцуговая рэакцыя – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Палімэразная ланцуговая рэакцыя" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target">Беларуская (тарашкевіца)</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D0%BD%D0%B0_%D0%B2%D0%B5%D1%80%D0%B8%D0%B6%D0%BD%D0%B0_%D1%80%D0%B5%D0%B0%D0%BA%D1%86%D0%B8%D1%8F" title="Полимеразна верижна реакция – Bulgarian" lang="bg" hreflang="bg" data-title="Полимеразна верижна реакция" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Polimerazna_lan%C4%8Dana_reakcija" title="Polimerazna lančana reakcija – Bosnian" lang="bs" hreflang="bs" data-title="Polimerazna lančana reakcija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-ca badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ca.wikipedia.org/wiki/Reacci%C3%B3_en_cadena_de_la_polimerasa" title="Reacció en cadena de la polimerasa – Catalan" lang="ca" hreflang="ca" data-title="Reacció en cadena de la polimerasa" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Polymer%C3%A1zov%C3%A1_%C5%99et%C4%9Bzov%C3%A1_reakce" title="Polymerázová řetězová reakce – Czech" lang="cs" hreflang="cs" data-title="Polymerázová řetězová reakce" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/PCR" title="PCR – Danish" lang="da" hreflang="da" data-title="PCR" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://de.wikipedia.org/wiki/Polymerase-Kettenreaktion" title="Polymerase-Kettenreaktion – German" lang="de" hreflang="de" data-title="Polymerase-Kettenreaktion" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Pol%C3%BCmeraasi_ahelreaktsioon" title="Polümeraasi ahelreaktsioon – Estonian" lang="et" hreflang="et" data-title="Polümeraasi ahelreaktsioon" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%91%CE%BB%CF%85%CF%83%CE%B9%CE%B4%CF%89%CF%84%CE%AE_%CE%B1%CE%BD%CF%84%CE%AF%CE%B4%CF%81%CE%B1%CF%83%CE%B7_%CF%80%CE%BF%CE%BB%CF%85%CE%BC%CE%B5%CF%81%CE%AC%CF%83%CE%B7%CF%82" title="Αλυσιδωτή αντίδραση πολυμεράσης – Greek" lang="el" hreflang="el" data-title="Αλυσιδωτή αντίδραση πολυμεράσης" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Reacci%C3%B3n_en_cadena_de_la_polimerasa" title="Reacción en cadena de la polimerasa – Spanish" lang="es" hreflang="es" data-title="Reacción en cadena de la polimerasa" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Polimeraza_%C4%89en-reakcio" title="Polimeraza ĉen-reakcio – Esperanto" lang="eo" hreflang="eo" data-title="Polimeraza ĉen-reakcio" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Polimerasaren_kate-erreakzioa" title="Polimerasaren kate-erreakzioa – Basque" lang="eu" hreflang="eu" data-title="Polimerasaren kate-erreakzioa" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%88%D8%A7%DA%A9%D9%86%D8%B4_%D8%B2%D9%86%D8%AC%DB%8C%D8%B1%D9%87%E2%80%8C%D8%A7%DB%8C_%D9%BE%D9%84%DB%8C%D9%85%D8%B1%D8%A7%D8%B2" title="واکنش زنجیره‌ای پلیمراز – Persian" lang="fa" hreflang="fa" data-title="واکنش زنجیره‌ای پلیمراز" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/R%C3%A9action_en_cha%C3%AEne_par_polym%C3%A9rase" title="Réaction en chaîne par polymérase – French" lang="fr" hreflang="fr" data-title="Réaction en chaîne par polymérase" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Imoibri%C3%BA_slabhr%C3%BAil_polaim%C3%A9ar%C3%A1ise" title="Imoibriú slabhrúil polaiméaráise – Irish" lang="ga" hreflang="ga" data-title="Imoibriú slabhrúil polaiméaráise" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Reacci%C3%B3n_en_cadea_da_polimerase" title="Reacción en cadea da polimerase – Galician" lang="gl" hreflang="gl" data-title="Reacción en cadea da polimerase" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%A4%91%ED%95%A9%ED%9A%A8%EC%86%8C_%EC%97%B0%EC%87%84_%EB%B0%98%EC%9D%91" title="중합효소 연쇄 반응 – Korean" lang="ko" hreflang="ko" data-title="중합효소 연쇄 반응" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8A%D5%B8%D5%AC%D5%AB%D5%B4%D5%A5%D6%80%D5%A1%D5%A6%D5%A1%D5%B5%D5%AB%D5%B6_%D5%B7%D5%B2%D5%A9%D5%A1%D5%B5%D5%A1%D5%AF%D5%A1%D5%B6_%D5%BC%D5%A5%D5%A1%D5%AF%D6%81%D5%AB%D5%A1" title="Պոլիմերազային շղթայական ռեակցիա – Armenian" lang="hy" hreflang="hy" data-title="Պոլիմերազային շղթայական ռեակցիա" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A5%89%E0%A4%B2%E0%A4%BF%E0%A4%AE%E0%A4%B0%E0%A5%87%E0%A4%9C_%E0%A4%B6%E0%A5%83%E0%A4%82%E0%A4%96%E0%A4%B2%E0%A4%BE_%E0%A4%85%E0%A4%AD%E0%A4%BF%E0%A4%95%E0%A5%8D%E0%A4%B0%E0%A4%BF%E0%A4%AF%E0%A4%BE" title="पॉलिमरेज शृंखला अभिक्रिया – Hindi" lang="hi" hreflang="hi" data-title="पॉलिमरेज शृंखला अभिक्रिया" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Lan%C4%8Dana_reakcija_polimerazom" title="Lančana reakcija polimerazom – Croatian" lang="hr" hreflang="hr" data-title="Lančana reakcija polimerazom" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Reaksi_berantai_polimerase" title="Reaksi berantai polimerase – Indonesian" lang="id" hreflang="id" data-title="Reaksi berantai polimerase" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/PCR" title="PCR – Icelandic" lang="is" hreflang="is" data-title="PCR" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Reazione_a_catena_della_polimerasi" title="Reazione a catena della polimerasi – Italian" lang="it" hreflang="it" data-title="Reazione a catena della polimerasi" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/PCR" title="PCR – Hebrew" lang="he" hreflang="he" data-title="PCR" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%9E%E1%83%9D%E1%83%9A%E1%83%98%E1%83%9B%E1%83%94%E1%83%A0%E1%83%90%E1%83%96%E1%83%A3%E1%83%9A%E1%83%98_%E1%83%AF%E1%83%90%E1%83%AD%E1%83%95%E1%83%A3%E1%83%A0%E1%83%98_%E1%83%A0%E1%83%94%E1%83%90%E1%83%A5%E1%83%AA%E1%83%98%E1%83%90" title="პოლიმერაზული ჯაჭვური რეაქცია – Georgian" lang="ka" hreflang="ka" data-title="პოლიმერაზული ჯაჭვური რეაქცია" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D0%B4%D1%8B_%D1%82%D1%96%D0%B7%D0%B1%D0%B5%D0%BA%D1%82%D1%96_%D1%80%D0%B5%D0%B0%D0%BA%D1%86%D0%B8%D1%8F" title="Полимеразды тізбекті реакция – Kazakh" lang="kk" hreflang="kk" data-title="Полимеразды тізбекті реакция" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Polimer%C4%81zes_%C4%B7%C4%93des_reakcija" title="Polimerāzes ķēdes reakcija – Latvian" lang="lv" hreflang="lv" data-title="Polimerāzes ķēdes reakcija" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Polimerazin%C4%97_grandinin%C4%97_reakcija" title="Polimerazinė grandininė reakcija – Lithuanian" lang="lt" hreflang="lt" data-title="Polimerazinė grandininė reakcija" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Polimer%C3%A1z-l%C3%A1ncreakci%C3%B3" title="Polimeráz-láncreakció – Hungarian" lang="hu" hreflang="hu" data-title="Polimeráz-láncreakció" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D0%BD%D0%B0_%D0%B2%D0%B5%D1%80%D0%B8%D0%B6%D0%BD%D0%B0_%D1%80%D0%B5%D0%B0%D0%BA%D1%86%D0%B8%D1%98%D0%B0" title="Полимеразна верижна реакција – Macedonian" lang="mk" hreflang="mk" data-title="Полимеразна верижна реакција" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%AA%E0%B5%8B%E0%B4%B3%E0%B4%BF%E0%B4%AE%E0%B5%86%E0%B4%B1%E0%B5%87%E0%B4%AF%E0%B5%8D%E0%B4%B8%E0%B5%8D_%E0%B4%9A%E0%B5%86%E0%B4%AF%E0%B4%BF%E0%B5%BB_%E0%B4%B1%E0%B4%BF%E0%B4%AF%E0%B4%BE%E0%B4%95%E0%B5%8D%E0%B4%B7%E0%B5%BB" title="പോളിമെറേയ്സ് ചെയിൻ റിയാക്ഷൻ – Malayalam" lang="ml" hreflang="ml" data-title="പോളിമെറേയ്സ് ചെയിൻ റിയാക്ഷൻ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Tindak_balas_berantai_polimerase" title="Tindak balas berantai polimerase – Malay" lang="ms" hreflang="ms" data-title="Tindak balas berantai polimerase" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D1%8B%D0%BD_%D0%B3%D0%B8%D0%BD%D0%B6%D0%B8%D0%BD_%D1%83%D1%80%D0%B2%D0%B0%D0%BB" title="Полимеразын гинжин урвал – Mongolian" lang="mn" hreflang="mn" data-title="Полимеразын гинжин урвал" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target">Монгол</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Polymerasekettingreactie" title="Polymerasekettingreactie – Dutch" lang="nl" hreflang="nl" data-title="Polymerasekettingreactie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%9D%E3%83%AA%E3%83%A1%E3%83%A9%E3%83%BC%E3%82%BC%E9%80%A3%E9%8E%96%E5%8F%8D%E5%BF%9C" title="ポリメラーゼ連鎖反応 – Japanese" lang="ja" hreflang="ja" data-title="ポリメラーゼ連鎖反応" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/Polimeraase-k%C3%A4%C3%A4tereaksjoon" title="Polimeraase-käätereaksjoon – Northern Frisian" lang="frr" hreflang="frr" data-title="Polimeraase-käätereaksjoon" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target">Nordfriisk</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/PCR" title="PCR – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="PCR" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Reaccion_en_cadena_per_la_polimerasa" title="Reaccion en cadena per la polimerasa – Occitan" lang="oc" hreflang="oc" data-title="Reaccion en cadena per la polimerasa" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Polimeraza_zanjiri_reaktsiyasi" title="Polimeraza zanjiri reaktsiyasi – Uzbek" lang="uz" hreflang="uz" data-title="Polimeraza zanjiri reaktsiyasi" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AF_%D9%BE%D9%88%D9%84%DB%8C%D9%85%DB%8C%D8%B1%DB%90%D8%B2_%DA%81%D9%86%DA%81%DB%8C%D8%B1%D9%8A_%D8%AA%D8%B9%D8%A7%D9%85%D9%84" title="د پولیمیرېز ځنځیري تعامل – Pashto" lang="ps" hreflang="ps" data-title="د پولیمیرېز ځنځیري تعامل" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Reakcja_%C5%82a%C5%84cuchowa_polimerazy" title="Reakcja łańcuchowa polimerazy – Polish" lang="pl" hreflang="pl" data-title="Reakcja łańcuchowa polimerazy" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Rea%C3%A7%C3%A3o_em_cadeia_da_polimerase" title="Reação em cadeia da polimerase – Portuguese" lang="pt" hreflang="pt" data-title="Reação em cadeia da polimerase" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Reac%C8%9Bie_de_polimerizare_%C3%AEn_lan%C8%9B" title="Reacție de polimerizare în lanț – Romanian" lang="ro" hreflang="ro" data-title="Reacție de polimerizare în lanț" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D0%BD%D0%B0%D1%8F_%D1%86%D0%B5%D0%BF%D0%BD%D0%B0%D1%8F_%D1%80%D0%B5%D0%B0%D0%BA%D1%86%D0%B8%D1%8F" title="Полимеразная цепная реакция – Russian" lang="ru" hreflang="ru" data-title="Полимеразная цепная реакция" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/PCR" title="PCR – Albanian" lang="sq" hreflang="sq" data-title="PCR" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target">Shqip</a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%B4%E0%B7%9C%E0%B6%BD%E0%B7%92%E0%B6%B8%E0%B6%BB%E0%B7%9A%E0%B7%83%E0%B7%8A%E2%80%8C_%E0%B6%AF%E0%B7%8F%E0%B6%B8_%E0%B6%B4%E0%B7%8A%E2%80%8D%E0%B6%BB%E0%B6%AD%E0%B7%92%E0%B6%9A%E0%B7%8A%E2%80%8D%E0%B6%BB%E0%B7%92%E0%B6%BA%E0%B7%8F" title="පොලිමරේස්‌ දාම ප්‍රතික්‍රියා – Sinhala" lang="si" hreflang="si" data-title="පොලිමරේස්‌ දාම ප්‍රතික්‍රියා" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Polymerase_chain_reaction" title="Polymerase chain reaction – Simple English" lang="en-simple" hreflang="en-simple" data-title="Polymerase chain reaction" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Polymer%C3%A1zov%C3%A1_re%C5%A5azov%C3%A1_reakcia" title="Polymerázová reťazová reakcia – Slovak" lang="sk" hreflang="sk" data-title="Polymerázová reťazová reakcia" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Veri%C5%BEna_reakcija_s_polimerazo" title="Verižna reakcija s polimerazo – Slovenian" lang="sl" hreflang="sl" data-title="Verižna reakcija s polimerazo" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A0%D0%B5%D0%B0%D0%BA%D1%86%D0%B8%D1%98%D0%B0_%D0%BB%D0%B0%D0%BD%D1%87%D0%B0%D0%BD%D0%B5_%D0%BF%D0%BE%D0%BB%D0%B8%D0%BC%D0%B5%D1%80%D0%B8%D0%B7%D0%B0%D1%86%D0%B8%D1%98%D0%B5" title="Реакција ланчане полимеризације – Serbian" lang="sr" hreflang="sr" data-title="Реакција ланчане полимеризације" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Polimerazna_lan%C4%8Dana_reakcija" title="Polimerazna lančana reakcija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Polimerazna lančana reakcija" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Polymeraasiketjureaktio" title="Polymeraasiketjureaktio – Finnish" lang="fi" hreflang="fi" data-title="Polymeraasiketjureaktio" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/PCR" title="PCR – Swedish" lang="sv" hreflang="sv" data-title="PCR" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AA%E0%AE%BE%E0%AE%B2%E0%AE%BF%E0%AE%AE%E0%AE%B0%E0%AF%87%E0%AE%9A%E0%AF%81_%E0%AE%A4%E0%AF%8A%E0%AE%9F%E0%AE%B0%E0%AF%8D_%E0%AE%B5%E0%AE%BF%E0%AE%A9%E0%AF%88" title="பாலிமரேசு தொடர் வினை – Tamil" lang="ta" hreflang="ta" data-title="பாலிமரேசு தொடர் வினை" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-kab mw-list-item"><a href="https://kab.wikipedia.org/wiki/Tasedmirt_tanmezrit_n_Polimiraz" title="Tasedmirt tanmezrit n Polimiraz – Kabyle" lang="kab" hreflang="kab" data-title="Tasedmirt tanmezrit n Polimiraz" data-language-autonym="Taqbaylit" data-language-local-name="Kabyle" class="interlanguage-link-target">Taqbaylit</a></li><li class="interlanguage-link interwiki-te mw-list-item"><a href="https://te.wikipedia.org/wiki/%E0%B0%B5%E0%B1%87%E0%B0%A6%E0%B0%BF%E0%B0%95:%E0%B0%B5%E0%B0%B0%E0%B1%8D%E0%B0%A4%E0%B0%AE%E0%B0%BE%E0%B0%A8_%E0%B0%98%E0%B0%9F%E0%B0%A8%E0%B0%B2%E0%B1%81" title="వేదిక:వర్తమాన ఘటనలు – Telugu" lang="te" hreflang="te" data-title="వేదిక:వర్తమాన ఘటనలు" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target">తెలుగు</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%9B%E0%B8%8F%E0%B8%B4%E0%B8%81%E0%B8%B4%E0%B8%A3%E0%B8%B4%E0%B8%A2%E0%B8%B2%E0%B8%A5%E0%B8%B9%E0%B8%81%E0%B9%82%E0%B8%8B%E0%B9%88%E0%B8%9E%E0%B8%AD%E0%B8%A5%E0%B8%B4%E0%B9%80%E0%B8%A1%E0%B8%AD%E0%B9%80%E0%B8%A3%E0%B8%AA" title="ปฏิกิริยาลูกโซ่พอลิเมอเรส – Thai" lang="th" hreflang="th" data-title="ปฏิกิริยาลูกโซ่พอลิเมอเรส" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target">ไทย</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Polimeraz_zincir_reaksiyonu" title="Polimeraz zincir reaksiyonu – Turkish" lang="tr" hreflang="tr" data-title="Polimeraz zincir reaksiyonu" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9F%D0%BE%D0%BB%D1%96%D0%BC%D0%B5%D1%80%D0%B0%D0%B7%D0%BD%D0%B0_%D0%BB%D0%B0%D0%BD%D1%86%D1%8E%D0%B3%D0%BE%D0%B2%D0%B0_%D1%80%D0%B5%D0%B0%D0%BA%D1%86%D1%96%D1%8F" title="Полімеразна ланцюгова реакція – Ukrainian" lang="uk" hreflang="uk" data-title="Полімеразна ланцюгова реакція" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D9%BE%D9%88%D9%84%DB%8C%D9%85%D8%B1%D8%B2_%D8%B2%D9%86%D8%AC%DB%8C%D8%B1%DB%8C_%D8%AA%D8%B9%D8%A7%D9%85%D9%84" title="پولیمرز زنجیری تعامل – Urdu" lang="ur" hreflang="ur" data-title="پولیمرز زنجیری تعامل" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Ph%E1%BA%A3n_%E1%BB%A9ng_chu%E1%BB%97i_polymerase" 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class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Laboratory technique to multiply a DNA sample for study</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:PCR_tubes.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/PCR_tubes.png/220px-PCR_tubes.png" decoding="async" width="220" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/PCR_tubes.png/330px-PCR_tubes.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/PCR_tubes.png/440px-PCR_tubes.png 2x" data-file-width="800" data-file-height="656" /></a><figcaption>A strip of eight PCR tubes, each containing a 100 μL reaction mixture</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:PCR_masina_kasutamine.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/PCR_masina_kasutamine.jpg/220px-PCR_masina_kasutamine.jpg" decoding="async" width="220" height="146" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/PCR_masina_kasutamine.jpg/330px-PCR_masina_kasutamine.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/67/PCR_masina_kasutamine.jpg/440px-PCR_masina_kasutamine.jpg 2x" data-file-width="3008" data-file-height="2000" /></a><figcaption>Placing a strip of eight PCR tubes into a <a href="/wiki/Thermal_cycler" title="Thermal cycler">thermal cycler</a></figcaption></figure> The polymerase chain reaction (PCR) is a method widely used to make millions to billions of copies of a specific <a href="/wiki/DNA" title="DNA">DNA</a> sample rapidly, allowing scientists to amplify a very small sample of DNA (or a part of it) sufficiently to enable detailed study. PCR was invented in 1983 by American <a href="/wiki/Biochemist" title="Biochemist">biochemist</a> <a href="/wiki/Kary_Mullis" title="Kary Mullis">Kary Mullis</a> at <a href="/wiki/Cetus_Corporation" title="Cetus Corporation">Cetus Corporation</a>. Mullis and biochemist <a href="/wiki/Michael_Smith_(chemist)" title="Michael Smith (chemist)">Michael Smith</a>, who had developed other essential ways of manipulating DNA, were jointly awarded the <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a> in 1993.<a href="#cite_note-NobelPrize-1">[1]</a> PCR is fundamental to many of the procedures used in <a href="/wiki/Genetic_testing" title="Genetic testing">genetic testing</a> and research, including analysis of <a href="/wiki/Ancient_DNA" title="Ancient DNA">ancient samples of DNA</a> and identification of infectious agents. Using PCR, copies of very small amounts of <a href="/wiki/DNA_sequences" class="mw-redirect" title="DNA sequences">DNA sequences</a> are exponentially amplified in a series of cycles of temperature changes. PCR is now a common and often indispensable technique used in <a href="/wiki/Medical_laboratory" title="Medical laboratory">medical laboratory</a> research for a broad variety of applications including <a href="/wiki/Biomedical_research" class="mw-redirect" title="Biomedical research">biomedical research</a> and <a href="/wiki/Forensic_science" title="Forensic science">forensic science</a>.<a href="#cite_note-Saiki1-2">[2]</a><a href="#cite_note-Saiki2-3">[3]</a> The majority of PCR methods rely on <a href="/wiki/Thermal_cycler" title="Thermal cycler">thermal cycling</a>. Thermal cycling exposes reagents to repeated cycles of heating and cooling to permit different temperature-dependent reactions—specifically, <a href="/wiki/DNA_melting#Denaturation" class="mw-redirect" title="DNA melting">DNA melting</a> and <a href="/wiki/Enzyme" title="Enzyme">enzyme</a>-driven <a href="/wiki/DNA_replication" title="DNA replication">DNA replication</a>. PCR employs two main reagents—<a href="/wiki/Primer_(molecular_biology)" title="Primer (molecular biology)">primers</a> (which are short single strand DNA fragments known as <a href="/wiki/Oligonucleotide" title="Oligonucleotide">oligonucleotides</a> that are a <a href="/wiki/Complementary_DNA" title="Complementary DNA">complementary</a> sequence to the target DNA region) and a <a href="/wiki/Thermostable_DNA_polymerase" title="Thermostable DNA polymerase">thermostable DNA polymerase</a>. In the first step of PCR, the two strands of the DNA double helix are physically separated at a high temperature in a process called <a href="/wiki/Denaturation_(biochemistry)#Nucleic_acid_denaturation" title="Denaturation (biochemistry)">nucleic acid denaturation</a>. In the second step, the temperature is lowered and the primers bind to the complementary sequences of DNA. The two DNA strands then become <a href="/wiki/DNA#Polymerases" title="DNA">templates</a> for DNA polymerase to <a href="/wiki/Enzyme" title="Enzyme">enzymatically</a> assemble a new DNA strand from free <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a>, the building blocks of DNA. As PCR progresses, the DNA generated is itself used as a template for replication, setting in motion a <a href="/wiki/Chain_reaction" title="Chain reaction">chain reaction</a> in which the original DNA template is <a href="/wiki/Exponential_growth" title="Exponential growth">exponentially</a> amplified. Almost all PCR applications employ a <a href="/wiki/Thermostable_DNA_polymerase" title="Thermostable DNA polymerase">heat-stable DNA polymerase</a>, such as <a href="/wiki/Taq_polymerase" title="Taq polymerase">Taq polymerase</a>, an enzyme originally isolated from the <a href="/wiki/Thermophile" title="Thermophile">thermophilic</a> bacterium <a href="/wiki/Thermus_aquaticus" title="Thermus aquaticus">Thermus aquaticus</a>. If the polymerase used was heat-susceptible, it would denature under the high temperatures of the denaturation step. Before the use of Taq polymerase, DNA polymerase had to be manually added every cycle, which was a tedious and costly process.<a href="#cite_note-4">[4]</a> Applications of the technique include <a href="/wiki/DNA_cloning" class="mw-redirect" title="DNA cloning">DNA cloning</a> for <a href="/wiki/DNA_sequencing" title="DNA sequencing">sequencing</a>, gene cloning and manipulation, gene mutagenesis; construction of DNA-based <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogenies</a>, or functional analysis of <a href="/wiki/Gene" title="Gene">genes</a>; <a href="/wiki/Medical_diagnosis" title="Medical diagnosis">diagnosis</a> and <a href="/wiki/Monitoring_(medicine)" title="Monitoring (medicine)">monitoring</a> of <a href="/wiki/Genetic_disorder" title="Genetic disorder">genetic disorders</a>; amplification of ancient DNA;<a href="#cite_note-Ninfa-2009-5">[5]</a> analysis of genetic fingerprints for <a href="/wiki/DNA_profiling" title="DNA profiling">DNA profiling</a> (for example, in <a href="/wiki/Forensic_science" title="Forensic science">forensic science</a> and <a href="/wiki/Parentage_testing" class="mw-redirect" title="Parentage testing">parentage testing</a>); and detection of <a href="/wiki/Pathogen" title="Pathogen">pathogens</a> in <a href="/wiki/Nucleic_acid_test" title="Nucleic acid test">nucleic acid tests</a> for the diagnosis of <a href="/wiki/Infectious_disease" class="mw-redirect" title="Infectious disease">infectious diseases</a>. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Principles">Principles</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=1" title="Edit section: Principles">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Primitive_PCR_machine_for_scrap.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Primitive_PCR_machine_for_scrap.JPG/170px-Primitive_PCR_machine_for_scrap.JPG" decoding="async" width="170" height="192" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Primitive_PCR_machine_for_scrap.JPG/255px-Primitive_PCR_machine_for_scrap.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Primitive_PCR_machine_for_scrap.JPG/340px-Primitive_PCR_machine_for_scrap.JPG 2x" data-file-width="1122" data-file-height="1264" /></a><figcaption>An older, three-temperature <a href="/wiki/Thermal_cycler" title="Thermal cycler">thermal cycler</a> for PCR</figcaption></figure> PCR amplifies a specific region of a DNA strand (the DNA target). Most PCR methods amplify DNA fragments of between 0.1 and 10 <a href="/wiki/Kilo_base_pair" class="mw-redirect" title="Kilo base pair">kilo base pairs</a> (kbp) in length, although some techniques allow for amplification of fragments up to 40 kbp.<a href="#cite_note-6">[6]</a> The amount of amplified product is determined by the available substrates in the reaction, which becomes limiting as the reaction progresses.<a href="#cite_note-7">[7]</a> A basic PCR set-up requires several components and reagents,<a href="#cite_note-molecular_cloning-8">[8]</a> including: <ul><li>a DNA template that contains the DNA target region to amplify</li> <li>a <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a>; an enzyme that <a href="/wiki/Polymerization" title="Polymerization">polymerizes</a> new DNA strands; heat-resistant <a href="/wiki/Taq_polymerase" title="Taq polymerase">Taq polymerase</a> is especially common,<a href="#cite_note-9">[9]</a> as it is more likely to remain intact during the high-temperature DNA denaturation process</li> <li>two DNA <a href="/wiki/Primer_(molecular_biology)" title="Primer (molecular biology)">primers</a> that are <a href="/wiki/Complementarity_(molecular_biology)" title="Complementarity (molecular biology)">complementary</a> to the <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">3' (three prime) ends</a> of each of the <a href="/wiki/Sense_and_antisense" class="mw-redirect" title="Sense and antisense">sense and anti-sense</a> strands of the DNA target (DNA polymerase can only bind to and elongate from a double-stranded region of DNA; without primers, there is no double-stranded initiation site at which the polymerase can bind);<a href="#cite_note-learn.genetics.utah.edu-10">[10]</a> specific primers that are complementary to the DNA target region are selected beforehand, and are often custom-made in a laboratory or purchased from commercial biochemical suppliers</li> <li>deoxynucleoside triphosphates, or dNTPs (sometimes called "deoxynucleotide triphosphates"; <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a> containing triphosphate groups), the building blocks from which the DNA polymerase synthesizes a new DNA strand</li> <li>a <a href="/wiki/Buffer_solution" title="Buffer solution">buffer solution</a> providing a suitable chemical environment for optimum activity and stability of the DNA polymerase</li> <li><a href="/wiki/Bivalent_(chemistry)" class="mw-redirect" title="Bivalent (chemistry)">bivalent</a> <a href="/wiki/Cations" class="mw-redirect" title="Cations">cations</a>, typically <a href="/wiki/Magnesium" title="Magnesium">magnesium</a> (Mg) or <a href="/wiki/Manganese" title="Manganese">manganese</a> (Mn) ions; Mg2+ is the most common, but Mn2+ can be used for <a href="/wiki/PCR_mutagenesis" class="mw-redirect" title="PCR mutagenesis">PCR-mediated DNA mutagenesis</a>, as a higher Mn2+ concentration increases the error rate during DNA synthesis;<a href="#cite_note-11">[11]</a> and monovalent cations, typically <a href="/wiki/Potassium" title="Potassium">potassium</a> (K) ions[<a href="/wiki/Wikipedia:NOTRS" class="mw-redirect" title="Wikipedia:NOTRS">better source needed</a>]</li></ul> The reaction is commonly carried out in a volume of 10–200 <a href="/wiki/Microliter" class="mw-redirect" title="Microliter">μL</a> in small reaction tubes (0.2–0.5 mL volumes) in a <a href="/wiki/Thermal_cycler" title="Thermal cycler">thermal cycler</a>. The thermal cycler heats and cools the reaction tubes to achieve the temperatures required at each step of the reaction (see below). Many modern thermal cyclers make use of a <a href="/wiki/Thermoelectric_effect#Peltier_effect" title="Thermoelectric effect">Peltier device</a>, which permits both heating and cooling of the block holding the PCR tubes simply by reversing the device's electric current. Thin-walled reaction tubes permit favorable <a href="/wiki/Thermal_conductivity" class="mw-redirect" title="Thermal conductivity">thermal conductivity</a> to allow for rapid thermal equilibrium. Most thermal cyclers have heated lids to prevent <a href="/wiki/Condensation" title="Condensation">condensation</a> at the top of the reaction tube. Older thermal cyclers lacking a heated lid require a layer of oil on top of the reaction mixture or a ball of wax inside the tube.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading3"><h3 id="Procedure">Procedure</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=2" title="Edit section: Procedure">edit</a>]</div> Typically, PCR consists of a series of 20–40 repeated temperature changes, called thermal cycles, with each cycle commonly consisting of two or three discrete temperature steps (see figure below). The cycling is often preceded by a single temperature step at a very high temperature (>90 °C (194 °F)), and followed by one hold at the end for final product extension or brief storage. The temperatures used and the length of time they are applied in each cycle depend on a variety of parameters, including the enzyme used for DNA synthesis, the concentration of bivalent ions and dNTPs in the reaction, and the <a href="/wiki/DNA_melting" class="mw-redirect" title="DNA melting">melting temperature</a> (Tm) of the primers.<a href="#cite_note-12">[12]</a> The individual steps common to most PCR methods are as follows: <ul><li>Initialization: This step is only required for DNA polymerases that require heat activation by <a href="/wiki/Hot_start_PCR" title="Hot start PCR">hot-start PCR</a>.<a href="#cite_note-antibody_hot_start-13">[13]</a> It consists of heating the reaction chamber to a temperature of 94–96 °C (201–205 °F), or 98 °C (208 °F) if extremely thermostable polymerases are used, which is then held for 1–10 minutes.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>]</li> <li><a href="/wiki/Denaturation_(biochemistry)#Nucleic_acid_denaturation" title="Denaturation (biochemistry)">Denaturation</a>: This step is the first regular cycling event and consists of heating the reaction chamber to 94–98 °C (201–208 °F) for 20–30 seconds. This causes <a href="/wiki/DNA_melting" class="mw-redirect" title="DNA melting">DNA melting</a>, or denaturation, of the double-stranded DNA template by breaking the <a href="/wiki/Hydrogen_bond" title="Hydrogen bond">hydrogen bonds</a> between complementary bases, yielding two single-stranded DNA molecules.</li> <li><a href="/wiki/Annealing_(biology)" class="mw-redirect" title="Annealing (biology)">Annealing</a>: In the next step, the reaction temperature is lowered to 50–65 °C (122–149 °F) for 20–40 seconds, allowing annealing of the primers to each of the single-stranded DNA templates. Two different primers are typically included in the reaction mixture: one for each of the two single-stranded complements containing the target region. The primers are single-stranded sequences themselves, but are much shorter than the length of the target region, complementing only very short sequences at the 3' end of each strand.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>]</li></ul> <dl><dd>It is critical to determine a proper temperature for the annealing step because efficiency and specificity are strongly affected by the annealing temperature. This temperature must be low enough to allow for <a href="/wiki/DNA%E2%80%93DNA_hybridization" title="DNA–DNA hybridization">hybridization</a> of the primer to the strand, but high enough for the hybridization to be specific, i.e., the primer should bind only to a perfectly complementary part of the strand, and nowhere else. If the temperature is too low, the primer may bind imperfectly. If it is too high, the primer may not bind at all. A typical annealing temperature is about 3–5 °C below the Tm of the primers used. Stable hydrogen bonds between complementary bases are formed only when the primer sequence very closely matches the template sequence. During this step, the polymerase binds to the primer-template hybrid and begins DNA formation.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>]</dd></dl> <ul><li>Extension/elongation: The temperature at this step depends on the DNA polymerase used; the optimum <a href="/wiki/Enzyme" title="Enzyme">activity</a> temperature for the thermostable DNA polymerase of Taq polymerase is approximately 75–80 °C (167–176 °F),<a href="#cite_note-Chien_et_al.-14">[14]</a><a href="#cite_note-Lawyer_et_al.-15">[15]</a> though a temperature of 72 °C (162 °F) is commonly used with this enzyme. In this step, the DNA polymerase synthesizes a new DNA strand complementary to the DNA template strand by adding free dNTPs from the reaction mixture that is complementary to the template in the <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">5'-to-3' direction</a>, <a href="/wiki/Condensation_reaction" title="Condensation reaction">condensing</a> the 5'-<a href="/wiki/Phosphate_group" class="mw-redirect" title="Phosphate group">phosphate group</a> of the dNTPs with the 3'-<a href="/wiki/Hydroxy_group" title="Hydroxy group">hydroxy group</a> at the end of the nascent (elongating) DNA strand. The precise time required for elongation depends both on the DNA polymerase used and on the length of the DNA target region to amplify. As a rule of thumb, at their optimal temperature, most DNA polymerases polymerize a thousand bases per minute. Under optimal conditions (i.e., if there are no limitations due to limiting substrates or reagents), at each extension/elongation step, the number of DNA target sequences is doubled. With each successive cycle, the original template strands plus all newly generated strands become template strands for the next round of elongation, leading to exponential (geometric) amplification of the specific DNA target region.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>]</li></ul> <dl><dd>The processes of denaturation, annealing and elongation constitute a single cycle. Multiple cycles are required to amplify the DNA target to millions of copies. The formula used to calculate the number of DNA copies formed after a given number of cycles is 2n, where n is the number of cycles. Thus, a reaction set for 30 cycles results in 230, or 1,073,741,824 copies of the original double-stranded DNA target region.</dd></dl> <ul><li>Final elongation: This single step is optional, but is performed at a temperature of 70–74 °C (158–165 °F) (the temperature range required for optimal activity of most polymerases used in PCR) for 5–15 minutes after the last PCR cycle to ensure that any remaining single-stranded DNA is fully elongated.</li> <li>Final hold: The final step cools the reaction chamber to 4–15 °C (39–59 °F) for an indefinite time, and may be employed for short-term storage of the PCR products.</li></ul> <figure class="mw-halign-center" typeof="mw:File/Frameless"><a href="/wiki/File:Polymerase_chain_reaction-en.svg" class="mw-file-description" title="Schematic drawing of a complete PCR cycle"><img alt="Schematic drawing of a complete PCR cycle" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Polymerase_chain_reaction-en.svg/1024px-Polymerase_chain_reaction-en.svg.png" decoding="async" width="1024" height="438" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Polymerase_chain_reaction-en.svg/1536px-Polymerase_chain_reaction-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ab/Polymerase_chain_reaction-en.svg/2048px-Polymerase_chain_reaction-en.svg.png 2x" data-file-width="512" data-file-height="219" /></a><figcaption>Schematic drawing of a complete PCR cycle</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Roland_Gel.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Roland_Gel.JPG/220px-Roland_Gel.JPG" decoding="async" width="220" height="207" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Roland_Gel.JPG/330px-Roland_Gel.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Roland_Gel.JPG/440px-Roland_Gel.JPG 2x" data-file-width="518" data-file-height="488" /></a><figcaption><a href="/wiki/Ethidium_bromide" title="Ethidium bromide">Ethidium bromide</a>-stained PCR products after <a href="/wiki/Gel_electrophoresis" title="Gel electrophoresis">gel electrophoresis</a>. Two sets of primers were used to amplify a target sequence from three different tissue samples. No amplification is present in sample #1; DNA bands in sample #2 and #3 indicate successful amplification of the target sequence. The gel also shows a positive control, and a DNA ladder containing DNA fragments of defined length for sizing the bands in the experimental PCRs.</figcaption></figure> To check whether the PCR successfully generated the anticipated DNA target region (also sometimes referred to as the amplimer or <a href="/wiki/Amplicon" title="Amplicon">amplicon</a>), <a href="/wiki/Agarose_gel_electrophoresis" title="Agarose gel electrophoresis">agarose gel electrophoresis</a> may be employed for size separation of the PCR products. The size of the PCR products is determined by comparison with a <a href="/wiki/DNA_ladder" class="mw-redirect" title="DNA ladder">DNA ladder</a>, a molecular weight marker which contains DNA fragments of known sizes, which runs on the gel alongside the PCR products. <figure class="mw-default-size mw-halign-center" typeof="mw:File"><a href="/wiki/File:Tucker_PCR.png" class="mw-file-description" title="Tucker PCR"><img alt="Tucker PCR" src="//upload.wikimedia.org/wikipedia/commons/5/5a/Tucker_PCR.png" decoding="async" width="773" height="707" class="mw-file-element" data-file-width="773" data-file-height="707" /></a><figcaption>Tucker PCR</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Stages">Stages</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=3" title="Edit section: Stages">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Exponential_Amplification.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Exponential_Amplification.svg/290px-Exponential_Amplification.svg.png" decoding="async" width="290" height="107" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Exponential_Amplification.svg/435px-Exponential_Amplification.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Exponential_Amplification.svg/580px-Exponential_Amplification.svg.png 2x" data-file-width="512" data-file-height="189" /></a><figcaption>Exponential amplification</figcaption></figure> As with other chemical reactions, the reaction rate and efficiency of PCR are affected by limiting factors. Thus, the entire PCR process can further be divided into three stages based on reaction progress: <ul><li>Exponential amplification: At every cycle, the amount of product is doubled (assuming 100% reaction efficiency). After 30 cycles, a single copy of DNA can be increased up to 1,000,000,000 (one billion) copies. In a sense, then, the replication of a discrete strand of DNA is being manipulated in a tube under controlled conditions.<a href="#cite_note-Schochetman_1988_1154–1157-16">[16]</a> The reaction is very sensitive: only minute quantities of DNA must be present.</li> <li>Leveling off stage: The reaction slows as the DNA polymerase loses activity and as consumption of reagents, such as dNTPs and primers, causes them to become more limited.</li> <li>Plateau: No more product accumulates due to exhaustion of reagents and enzyme.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Optimization">Optimization</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=4" title="Edit section: Optimization">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/PCR_optimization" class="mw-redirect" title="PCR optimization">PCR optimization</a></div> In practice, PCR can fail for various reasons, such as sensitivity or contamination.<a href="#cite_note-17">[17]</a><a href="#cite_note-18">[18]</a> Contamination with extraneous DNA can lead to spurious products and is addressed with lab protocols and procedures that separate pre-PCR mixtures from potential DNA contaminants.<a href="#cite_note-molecular_cloning-8">[8]</a> For instance, if DNA from a crime scene is analyzed, a single DNA molecule from lab personnel could be amplified and misguide the investigation. Hence the PCR-setup areas is separated from the analysis or purification of other PCR products, disposable plasticware used, and the work surface between reaction setups needs to be thoroughly cleaned. Specificity can be adjusted by experimental conditions so that no spurious products are generated. Primer-design techniques are important in improving PCR product yield and in avoiding the formation of unspecific products. The usage of alternate buffer components or polymerase enzymes can help with amplification of long or otherwise problematic regions of DNA. For instance, Q5 polymerase is said to be ≈280 times less error-prone than Taq polymerase.<a href="#cite_note-19">[19]</a><a href="#cite_note-20">[20]</a> Both the running parameters (e.g. temperature and duration of cycles), or the addition of reagents, such as <a href="/wiki/Formamide" title="Formamide">formamide</a>, may increase the specificity and yield of PCR.<a href="#cite_note-21">[21]</a> Computer simulations of theoretical PCR results (<a href="/wiki/In_silico_PCR" title="In silico PCR">Electronic PCR</a>) may be performed to assist in primer design.<a href="#cite_note-ePCR-22">[22]</a> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=5" title="Edit section: Applications">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Selective_DNA_isolation">Selective DNA isolation</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=6" title="Edit section: Selective DNA isolation">edit</a>]</div> PCR allows isolation of DNA fragments from genomic DNA by selective amplification of a specific region of DNA. This use of PCR augments many ways, such as generating <a href="/wiki/Hybridization_probe" title="Hybridization probe">hybridization probes</a> for <a href="/wiki/Southern_blot" title="Southern blot">Southern</a> or <a href="/wiki/Northern_blot" title="Northern blot">northern</a> hybridization and <a href="/wiki/DNA_cloning" class="mw-redirect" title="DNA cloning">DNA cloning</a>, which require larger amounts of DNA, representing a specific DNA region. PCR supplies these techniques with high amounts of pure DNA, enabling analysis of DNA samples even from very small amounts of starting material.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Other applications of PCR include <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> to determine unknown PCR-amplified sequences in which one of the amplification primers may be used in <a href="/wiki/Sanger_sequencing" title="Sanger sequencing">Sanger sequencing</a>, isolation of a DNA sequence to expedite recombinant DNA technologies involving the insertion of a DNA sequence into a <a href="/wiki/Plasmid" title="Plasmid">plasmid</a>, <a href="/wiki/Phage" class="mw-redirect" title="Phage">phage</a>, or <a href="/wiki/Cosmid" title="Cosmid">cosmid</a> (depending on size) or the genetic material of another organism. Bacterial colonies (such as <a href="/wiki/Escherichia_coli" title="Escherichia coli">E. coli</a>) can be rapidly screened by PCR for correct DNA <a href="/wiki/Plasmid" title="Plasmid">vector</a> constructs.<a href="#cite_note-Hybrid-23">[23]</a> PCR may also be used for <a href="/wiki/Genetic_fingerprinting" class="mw-redirect" title="Genetic fingerprinting">genetic fingerprinting</a>; a forensic technique used to identify a person or organism by comparing experimental DNAs through different PCR-based methods.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Pcr_fingerprint.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/02/Pcr_fingerprint.png/170px-Pcr_fingerprint.png" decoding="async" width="170" height="256" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/02/Pcr_fingerprint.png/255px-Pcr_fingerprint.png 1.5x, //upload.wikimedia.org/wikipedia/commons/0/02/Pcr_fingerprint.png 2x" data-file-width="331" data-file-height="498" /></a><figcaption>Electrophoresis of PCR-amplified DNA fragments: <div><ol><li>Father</li><li>Child</li><li>Mother</li></ol></div> The child has inherited some, but not all, of the fingerprints of each of its parents, giving it a new, unique fingerprint.</figcaption></figure> Some PCR fingerprint methods have high discriminative power and can be used to identify genetic relationships between individuals, such as parent-child or between siblings, and are used in paternity testing (Fig. 4). This technique may also be used to determine evolutionary relationships among organisms when certain <a href="/wiki/Molecular_clock" title="Molecular clock">molecular clocks</a> are used (i.e. the <a href="/wiki/16S_rRNA" class="mw-redirect" title="16S rRNA">16S rRNA</a> and recA genes of microorganisms).<a href="#cite_note-24">[24]</a> <div class="mw-heading mw-heading3"><h3 id="Amplification_and_quantification_of_DNA">Amplification and quantification of DNA</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=7" title="Edit section: Amplification and quantification of DNA">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Use_of_DNA_in_forensic_entomology" title="Use of DNA in forensic entomology">Use of DNA in forensic entomology</a></div> Because PCR amplifies the regions of DNA that it targets, PCR can be used to analyze extremely small amounts of sample. This is often critical for <a href="/wiki/Forensic_analysis" class="mw-redirect" title="Forensic analysis">forensic analysis</a>, when only a trace amount of DNA is available as evidence. PCR may also be used in the analysis of <a href="/wiki/Ancient_DNA" title="Ancient DNA">ancient DNA</a> that is tens of thousands of years old. These PCR-based techniques have been successfully used on animals, such as a forty-thousand-year-old <a href="/wiki/Mammoth" title="Mammoth">mammoth</a>, and also on human DNA, in applications ranging from the analysis of Egyptian <a href="/wiki/Mummy" title="Mummy">mummies</a> to the identification of a Russian <a href="/wiki/Tsar" title="Tsar">tsar</a> and the body of English king <a href="/wiki/Richard_III" class="mw-redirect" title="Richard III">Richard III</a>.<a href="#cite_note-25">[25]</a> <a href="/wiki/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">Quantitative PCR</a> or Real Time PCR (qPCR,<a href="#cite_note-26">[26]</a> not to be confused with <a href="/wiki/Reverse_transcription_polymerase_chain_reaction" title="Reverse transcription polymerase chain reaction">RT-PCR</a>) methods allow the estimation of the amount of a given sequence present in a sample—a technique often applied to quantitatively determine levels of <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>. Quantitative PCR is an established tool for DNA quantification that measures the accumulation of DNA product after each round of PCR amplification. qPCR allows the quantification and detection of a specific DNA sequence in real time since it measures concentration while the synthesis process is taking place. There are two methods for simultaneous detection and quantification. The first method consists of using <a href="/wiki/Fluorophore" title="Fluorophore">fluorescent</a> dyes that are retained nonspecifically in between the double strands. The second method involves probes that code for specific sequences and are fluorescently labeled. Detection of DNA using these methods can only be seen after the hybridization of probes with its <a href="/wiki/Complementary_DNA" title="Complementary DNA">complementary DNA</a> (cDNA) takes place. An interesting technique combination is real-time PCR and reverse transcription. This sophisticated technique, called RT-qPCR, allows for the quantification of a small quantity of RNA. Through this combined technique, mRNA is converted to cDNA, which is further quantified using qPCR. This technique lowers the possibility of error at the end point of PCR,<a href="#cite_note-Garibyan,_Avashia_1–4-27">[27]</a> increasing chances for detection of genes associated with genetic diseases such as cancer.<a href="#cite_note-Ninfa-2009-5">[5]</a> Laboratories use RT-qPCR for the purpose of sensitively measuring gene regulation. The mathematical foundations for the reliable quantification of the PCR<a href="#cite_note-28">[28]</a> and RT-qPCR<a href="#cite_note-29">[29]</a> facilitate the implementation of accurate fitting procedures of experimental data in research, medical, diagnostic and infectious disease applications.<a href="#cite_note-30">[30]</a><a href="#cite_note-31">[31]</a><a href="#cite_note-32">[32]</a><a href="#cite_note-33">[33]</a> <div class="mw-heading mw-heading3"><h3 id="Medical_and_diagnostic_applications">Medical and diagnostic applications</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=8" title="Edit section: Medical and diagnostic applications">edit</a>]</div> Prospective parents can be tested for being <a href="/wiki/Genetic_carrier" class="mw-redirect" title="Genetic carrier">genetic carriers</a>, or their children might be tested for actually being affected by a <a href="/wiki/Cystic_fibrosis" title="Cystic fibrosis">disease</a>.<a href="#cite_note-Saiki1-2">[2]</a> DNA samples for <a href="/wiki/Prenatal_diagnosis" class="mw-redirect" title="Prenatal diagnosis">prenatal testing</a> can be obtained by <a href="/wiki/Amniocentesis" title="Amniocentesis">amniocentesis</a>, <a href="/wiki/Chorionic_villus_sampling" title="Chorionic villus sampling">chorionic villus sampling</a>, or even by the analysis of rare fetal cells circulating in the mother's bloodstream. PCR analysis is also essential to <a href="/wiki/Preimplantation_genetic_diagnosis" title="Preimplantation genetic diagnosis">preimplantation genetic diagnosis</a>, where individual cells of a developing embryo are tested for mutations. <ul><li>PCR can also be used as part of a sensitive test for <a href="/wiki/Tissue_typing" title="Tissue typing">tissue typing</a>, vital to <a href="/wiki/Organ_transplantation" title="Organ transplantation">organ transplantation</a>. As of 2008,<a class="external text" href="https://en.wikipedia.org/w/index.php?title=Polymerase_chain_reaction&action=edit">[update]</a> there is even a proposal to replace the traditional antibody-based tests for <a href="/wiki/Blood_type" title="Blood type">blood type</a> with PCR-based tests.<a href="#cite_note-34">[34]</a></li> <li>Many forms of cancer involve alterations to <a href="/wiki/Oncogene" title="Oncogene">oncogenes</a>. By using PCR-based tests to study these mutations, therapy regimens can sometimes be individually customized to a patient. PCR permits early diagnosis of <a href="/wiki/Malignant" class="mw-redirect" title="Malignant">malignant</a> diseases such as <a href="/wiki/Leukemia" title="Leukemia">leukemia</a> and <a href="/wiki/Lymphoma" title="Lymphoma">lymphomas</a>, which is currently the highest-developed in cancer research and is already being used routinely. PCR assays can be performed directly on genomic DNA samples to detect translocation-specific malignant cells at a sensitivity that is at least 10,000 fold higher than that of other methods.<a href="#cite_note-35">[35]</a> PCR is very useful in the medical field since it allows for the isolation and amplification of tumor suppressors. Quantitative PCR for example, can be used to quantify and analyze single cells, as well as recognize DNA, mRNA and protein confirmations and combinations.<a href="#cite_note-Garibyan,_Avashia_1–4-27">[27]</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Infectious_disease_applications">Infectious disease applications</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=9" title="Edit section: Infectious disease applications">edit</a>]</div> PCR allows for rapid and highly specific diagnosis of infectious diseases, including those caused by bacteria or viruses.<a href="#cite_note-Cai2014-36">[36]</a> PCR also permits identification of non-cultivatable or slow-growing microorganisms such as <a href="/wiki/Mycobacterium" title="Mycobacterium">mycobacteria</a>, <a href="/wiki/Anaerobic_organism" title="Anaerobic organism">anaerobic bacteria</a>, or <a href="/wiki/Virus" title="Virus">viruses</a> from <a href="/wiki/Tissue_culture" title="Tissue culture">tissue culture</a> assays and <a href="/wiki/Animal_model" class="mw-redirect" title="Animal model">animal models</a>. The basis for PCR diagnostic applications in microbiology is the detection of infectious agents and the discrimination of non-pathogenic from pathogenic strains by virtue of specific genes.<a href="#cite_note-Cai2014-36">[36]</a><a href="#cite_note-37">[37]</a> Characterization and detection of infectious disease organisms have been revolutionized by PCR in the following ways: <ul><li>The human immunodeficiency virus (or <a href="/wiki/HIV" title="HIV">HIV</a>), is a difficult target to find and eradicate. The earliest tests for infection relied on the presence of antibodies to the virus circulating in the bloodstream. However, antibodies don't appear until many weeks after infection, maternal antibodies mask the infection of a newborn, and therapeutic agents to fight the infection don't affect the antibodies. PCR <a href="/wiki/HIV_test" class="mw-redirect" title="HIV test">tests</a> have been developed that can detect as little as one viral genome among the DNA of over 50,000 host cells.<a href="#cite_note-38">[38]</a> Infections can be detected earlier, donated blood can be screened directly for the virus, newborns can be immediately tested for infection, and the effects of antiviral treatments can be <a href="/wiki/Viral_load" title="Viral load">quantified</a>.</li> <li>Some disease organisms, such as that for <a href="/wiki/Tuberculosis" title="Tuberculosis">tuberculosis</a>, are difficult to sample from patients and slow to be <a href="/wiki/Tuberculosis_diagnosis" class="mw-redirect" title="Tuberculosis diagnosis">grown</a> in the laboratory. PCR-based tests have allowed detection of small numbers of disease organisms (both live or dead), in convenient <a href="/wiki/Sputum" title="Sputum">samples</a>. Detailed genetic analysis can also be used to detect antibiotic resistance, allowing immediate and effective therapy. The effects of therapy can also be immediately evaluated.</li> <li>The spread of a <a href="/wiki/Pathogen" title="Pathogen">disease organism</a> through populations of <a href="/wiki/Domestication#Animals" title="Domestication">domestic</a> or <a href="/wiki/Wildlife" title="Wildlife">wild</a> animals can be monitored by PCR testing. In many cases, the appearance of new virulent sub-types can be detected and monitored. The sub-types of an organism that were responsible for <a href="/wiki/List_of_epidemics" class="mw-redirect" title="List of epidemics">earlier epidemics</a> can also be determined by PCR analysis.</li> <li>Viral DNA can be detected by PCR. The primers used must be specific to the targeted sequences in the DNA of a virus, and PCR can be used for diagnostic analyses or DNA sequencing of the viral genome. The high sensitivity of PCR permits virus detection soon after infection and even before the onset of disease.<a href="#cite_note-Cai2014-36">[36]</a> Such early detection may give physicians a significant lead time in treatment. The amount of virus ("<a href="/wiki/Viral_load" title="Viral load">viral load</a>") in a patient can also be quantified by PCR-based DNA quantitation techniques (see below). A variant of PCR (<a href="/wiki/Reverse_transcription_polymerase_chain_reaction" title="Reverse transcription polymerase chain reaction">RT-PCR</a>) is used for detecting viral RNA rather than DNA: in this test the enzyme reverse transcriptase is used to generate a DNA sequence which matches the viral RNA; this DNA is then amplified as per the usual PCR method. RT-PCR is widely used to detect the SARS-CoV-2 viral genome.<a href="#cite_note-39">[39]</a></li> <li>Diseases such as pertussis (or <a href="/wiki/Whooping_cough" title="Whooping cough">whooping cough</a>) are caused by the bacteria <a href="/wiki/Bordetella_pertussis" title="Bordetella pertussis">Bordetella pertussis</a>. This bacteria is marked by a serious acute respiratory infection that affects various animals and humans and has led to the deaths of many young children. The pertussis toxin is a protein exotoxin that binds to cell receptors by two <a href="/wiki/Dimer_(chemistry)" class="mw-redirect" title="Dimer (chemistry)">dimers</a> and reacts with different cell types such as T lymphocytes which play a role in cell immunity.<a href="#cite_note-40">[40]</a> PCR is an important testing tool that can detect sequences within the gene for the pertussis toxin. Because PCR has a high sensitivity for the toxin and a rapid turnaround time, it is very efficient for diagnosing pertussis when compared to culture.<a href="#cite_note-41">[41]</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Forensic_applications">Forensic applications</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=10" title="Edit section: Forensic applications">edit</a>]</div> The development of PCR-based <a href="/wiki/Genetic_fingerprinting" class="mw-redirect" title="Genetic fingerprinting">genetic</a> (or <a href="/wiki/DNA_fingerprinting" class="mw-redirect" title="DNA fingerprinting">DNA</a>) fingerprinting protocols has seen widespread application in <a href="/wiki/Forensics" class="mw-redirect" title="Forensics">forensics</a>: <ul><li><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:US_Army_CID_agents_at_crime_scene.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/US_Army_CID_agents_at_crime_scene.jpg/220px-US_Army_CID_agents_at_crime_scene.jpg" decoding="async" width="220" height="206" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f1/US_Army_CID_agents_at_crime_scene.jpg/330px-US_Army_CID_agents_at_crime_scene.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f1/US_Army_CID_agents_at_crime_scene.jpg/440px-US_Army_CID_agents_at_crime_scene.jpg 2x" data-file-width="453" data-file-height="425" /></a><figcaption>DNA samples are often taken at crime scenes and analyzed by PCR.</figcaption></figure>In its most discriminating form, <a href="/wiki/Genetic_fingerprinting" class="mw-redirect" title="Genetic fingerprinting">genetic fingerprinting</a> can uniquely discriminate any one person from the entire population of the <a href="/wiki/Earth" title="Earth">world</a>. Minute samples of DNA can be isolated from a <a href="/wiki/O._J._Simpson_murder_case" class="mw-redirect" title="O. J. Simpson murder case">crime scene</a>, and <a href="/wiki/Combined_DNA_Index_System" title="Combined DNA Index System">compared</a> to that from suspects, or from a <a href="/wiki/National_DNA_database" class="mw-redirect" title="National DNA database">DNA database</a> of earlier evidence or convicts. Simpler versions of these tests are often used to rapidly rule out suspects during a criminal investigation. Evidence from decades-old crimes can be tested, confirming or <a href="/wiki/Innocence_Project" title="Innocence Project">exonerating</a> the people originally convicted.</li> <li>Forensic DNA typing has been an effective way of identifying or exonerating criminal suspects due to analysis of evidence discovered at a crime scene. The human genome has many repetitive regions that can be found within gene sequences or in non-coding regions of the genome. Specifically, up to 40% of human DNA is repetitive.<a href="#cite_note-Ninfa-2009-5">[5]</a> There are two distinct categories for these repetitive, non-coding regions in the genome. The first category is called variable number tandem repeats (VNTR), which are 10–100 base pairs long and the second category is called short tandem repeats (STR) and these consist of repeated 2–10 base pair sections. PCR is used to amplify several well-known VNTRs and STRs using primers that flank each of the repetitive regions. The sizes of the fragments obtained from any individual for each of the STRs will indicate which alleles are present. By analyzing several STRs for an individual, a set of alleles for each person will be found that statistically is likely to be unique.<a href="#cite_note-Ninfa-2009-5">[5]</a> Researchers have identified the complete sequence of the human genome. This sequence can be easily accessed through the NCBI website and is used in many real-life applications. For example, the FBI has compiled a set of DNA marker sites used for identification, and these are called the Combined DNA Index System (CODIS) DNA database.<a href="#cite_note-Ninfa-2009-5">[5]</a> Using this database enables statistical analysis to be used to determine the probability that a DNA sample will match. PCR is a very powerful and significant analytical tool to use for forensic DNA typing because researchers only need a very small amount of the target DNA to be used for analysis. For example, a single human hair with attached hair follicle has enough DNA to conduct the analysis. Similarly, a few sperm, skin samples from under the fingernails, or a small amount of blood can provide enough DNA for conclusive analysis.<a href="#cite_note-Ninfa-2009-5">[5]</a></li> <li>Less discriminating forms of <a href="/wiki/DNA_fingerprinting" class="mw-redirect" title="DNA fingerprinting">DNA fingerprinting</a> can help in <a href="/wiki/DNA_paternity_testing" title="DNA paternity testing">DNA paternity testing</a>, where an individual is matched with their close relatives. DNA from unidentified human remains can be tested, and compared with that from possible parents, siblings, or children. Similar testing can be used to confirm the biological parents of an adopted (or kidnapped) child. The actual biological father of a newborn can also be <a href="/wiki/DNA_paternity_testing" title="DNA paternity testing">confirmed</a> (or ruled out).</li> <li>The PCR AMGX/AMGY design has been shown to not only[<a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify">clarification needed</a>] facilitate in amplifying DNA sequences from a very minuscule amount of genome. However it can also be used for real-time sex determination from forensic bone samples. This provides a powerful and effective way to determine gender in forensic cases and ancient specimens.<a href="#cite_note-42">[42]</a></li></ul> <div class="mw-heading mw-heading3"><h3 id="Research_applications">Research applications</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=11" title="Edit section: Research applications">edit</a>]</div> PCR has been applied to many areas of research in molecular genetics: <ul><li>PCR allows rapid production of short pieces of DNA, even when not more than the sequence of the two primers is known. This ability of PCR augments many methods, such as generating <a href="/wiki/Nucleic_acid_hybridization" title="Nucleic acid hybridization">hybridization</a> <a href="/wiki/Hybridization_probe" title="Hybridization probe">probes</a> for <a href="/wiki/Southern_blot" title="Southern blot">Southern</a> or <a href="/wiki/Northern_blot" title="Northern blot">northern blot</a> hybridization. PCR supplies these techniques with large amounts of pure DNA, sometimes as a single strand, enabling analysis even from very small amounts of starting material.</li> <li>The task of <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> can also be assisted by PCR. Known segments of DNA can easily be produced from a patient with a genetic disease mutation. Modifications to the amplification technique can extract segments from a completely unknown genome, or can generate just a single strand of an area of interest.</li> <li>PCR has numerous applications to the more traditional process of <a href="/wiki/DNA_cloning" class="mw-redirect" title="DNA cloning">DNA cloning</a>. It can extract segments for insertion into a vector from a larger genome, which may be only available in small quantities. Using a single set of 'vector primers', it can also analyze or extract fragments that have already been inserted into vectors. Some alterations to the PCR protocol can generate mutations (general or site-directed) of an inserted fragment.</li> <li><a href="/wiki/Sequence-tagged_site" title="Sequence-tagged site">Sequence-tagged sites</a> is a process where PCR is used as an indicator that a particular segment of a genome is present in a particular clone. The <a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a> found this application vital to mapping the cosmid clones they were sequencing, and to coordinating the results from different laboratories.</li> <li>An application of PCR is the <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogenic</a> analysis of DNA from <a href="/wiki/Ancient_DNA" title="Ancient DNA">ancient sources</a>, such as that found in the recovered bones of <a href="/wiki/Neanderthal" title="Neanderthal">Neanderthals</a>, from frozen tissues of <a href="/wiki/Mammoth" title="Mammoth">mammoths</a>, or from the brain of Egyptian mummies.<a href="#cite_note-Schochetman_1988_1154–1157-16">[16]</a> In some cases the highly degraded DNA from these sources might be reassembled during the early stages of amplification.</li> <li>A common application of PCR is the study of patterns of <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>. Tissues (or even individual cells) can be analyzed at different stages to see which genes have become active, or which have been switched off. This application can also use <a href="/wiki/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">quantitative PCR</a> to quantitate the actual levels of expression</li> <li>The ability of PCR to simultaneously amplify several loci from individual sperm<a href="#cite_note-43">[43]</a> has greatly enhanced the more traditional task of <a href="/wiki/Genetic_linkage" title="Genetic linkage">genetic mapping</a> by studying <a href="/wiki/Chromosomal_crossover" title="Chromosomal crossover">chromosomal crossovers</a> after <a href="/wiki/Meiosis" title="Meiosis">meiosis</a>. Rare crossover events between very close loci have been directly observed by analyzing thousands of individual sperms. Similarly, unusual deletions, insertions, translocations, or inversions can be analyzed, all without having to wait (or pay) for the long and laborious processes of fertilization, embryogenesis, etc.</li> <li><a href="/wiki/Site-directed_mutagenesis" title="Site-directed mutagenesis">Site-directed mutagenesis</a>: PCR can be used to create mutant genes with mutations chosen by scientists at will. These mutations can be chosen in order to understand how proteins accomplish their functions, and to change or improve protein function.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Advantages">Advantages</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=12" title="Edit section: Advantages">edit</a>]</div> PCR has a number of advantages. It is fairly simple to understand and to use, and produces results rapidly. The technique is highly sensitive with the potential to produce millions to billions of copies of a specific product for sequencing, cloning, and analysis. qRT-PCR shares the same advantages as the PCR, with an added advantage of quantification of the synthesized product. Therefore, it has its uses to analyze alterations of gene expression levels in tumors, microbes, or other disease states.<a href="#cite_note-Garibyan,_Avashia_1–4-27">[27]</a> PCR is a very powerful and practical research tool. The sequencing of unknown etiologies of many diseases are being figured out by the PCR. The technique can help identify the sequence of previously unknown viruses related to those already known and thus give us a better understanding of the disease itself. If the procedure can be further simplified and sensitive non-radiometric detection systems can be developed, the PCR will assume a prominent place in the clinical laboratory for years to come.<a href="#cite_note-Schochetman_1988_1154–1157-16">[16]</a> <div class="mw-heading mw-heading2"><h2 id="Limitations">Limitations</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=13" title="Edit section: Limitations">edit</a>]</div> One major limitation of PCR is that prior information about the target sequence is necessary in order to generate the primers that will allow its selective amplification.<a href="#cite_note-Garibyan,_Avashia_1–4-27">[27]</a> This means that, typically, PCR users must know the precise sequence(s) upstream of the target region on each of the two single-stranded templates in order to ensure that the DNA polymerase properly binds to the primer-template hybrids and subsequently generates the entire target region during DNA synthesis.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] Like all enzymes, DNA polymerases are also prone to error, which in turn causes mutations in the PCR fragments that are generated.<a href="#cite_note-44">[44]</a> Another limitation of PCR is that even the smallest amount of contaminating DNA can be amplified, resulting in misleading or ambiguous results. To minimize the chance of contamination, investigators should reserve separate rooms for reagent preparation, the PCR, and analysis of product. Reagents should be dispensed into single-use <a href="/wiki/Sample_(material)" title="Sample (material)">aliquots</a>. Pipettors with disposable plungers and extra-long pipette tips should be routinely used.<a href="#cite_note-Schochetman_1988_1154–1157-16">[16]</a> It is moreover recommended to ensure that the lab set-up follows a unidirectional workflow. No materials or reagents used in the PCR and analysis rooms should ever be taken into the PCR preparation room without thorough decontamination.<a href="#cite_note-45">[45]</a> Environmental samples that contain humic acids may inhibit PCR amplification and lead to inaccurate results.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <div class="mw-heading mw-heading2"><h2 id="Variations">Variations</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=14" title="Edit section: Variations">edit</a>]</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/Variants_of_PCR" title="Variants of PCR">Variants of PCR</a></div> <ul><li>Allele-specific PCR or The amplification refractory mutation system (ARMS): a diagnostic or cloning technique based on single-nucleotide variations (SNVs not to be confused with <a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">SNPs</a>) (single-base differences in a patient). Any mutation involving single base change can be detected by this system. It requires prior knowledge of a DNA sequence, including differences between <a href="/wiki/Allele" title="Allele">alleles</a>, and uses primers whose 3' ends encompass the SNV (base pair buffer around SNV usually incorporated).<a href="#cite_note-46">[46]</a> PCR amplification under stringent conditions is much less efficient in the presence of a mismatch between template and primer, so successful amplification with an SNP-specific primer signals presence of the specific SNP or small deletions in a sequence.<a href="#cite_note-47">[47]</a> See <a href="/wiki/SNP_genotyping" title="SNP genotyping">SNP genotyping</a> for more information.</li> <li><a href="/wiki/Polymerase_cycling_assembly" title="Polymerase cycling assembly">Assembly PCR</a> or Polymerase Cycling Assembly (PCA): artificial synthesis of long DNA sequences by performing PCR on a pool of long oligonucleotides with short overlapping segments. The oligonucleotides alternate between sense and antisense directions, and the overlapping segments determine the order of the PCR fragments, thereby selectively producing the final long DNA product.<a href="#cite_note-Stemmer_et_al.-48">[48]</a></li> <li><a href="/wiki/Asymmetric_PCR" title="Asymmetric PCR">Asymmetric PCR</a>: preferentially amplifies one DNA strand in a double-stranded DNA template. It is used in <a href="/wiki/Sequencing" title="Sequencing">sequencing</a> and hybridization probing where amplification of only one of the two complementary strands is required. PCR is carried out as usual, but with a great excess of the primer for the strand targeted for amplification. Because of the slow (<a href="/wiki/Arithmetic" title="Arithmetic">arithmetic</a>) amplification later in the reaction after the limiting primer has been used up, extra cycles of PCR are required.<a href="#cite_note-Innis_et_al.-49">[49]</a> A recent modification on this process, known as Linear-After-The-Exponential-PCR (LATE-PCR), uses a limiting primer with a higher melting temperature (<a href="/wiki/DNA_melting" class="mw-redirect" title="DNA melting">Tm</a>) than the excess primer to maintain reaction efficiency as the limiting primer concentration decreases mid-reaction.<a href="#cite_note-Pierce_and_Wangh-50">[50]</a></li> <li>Convective PCR: a pseudo-isothermal way of performing PCR. Instead of repeatedly heating and cooling the PCR mixture, the solution is subjected to a thermal gradient. The resulting thermal instability driven convective flow automatically shuffles the PCR reagents from the hot and cold regions repeatedly enabling PCR.<a href="#cite_note-51">[51]</a> Parameters such as thermal boundary conditions and geometry of the PCR enclosure can be optimized to yield robust and rapid PCR by harnessing the emergence of chaotic flow fields.<a href="#cite_note-52">[52]</a> Such convective flow PCR setup significantly reduces device power requirement and operation time.</li> <li>Dial-out PCR: a highly parallel method for retrieving accurate DNA molecules for gene synthesis. A complex library of DNA molecules is modified with unique flanking tags before massively parallel sequencing. Tag-directed primers then enable the retrieval of molecules with desired sequences by PCR.<a href="#cite_note-Schwartz_et_al.-53">[53]</a></li> <li><a href="/wiki/Digital_PCR" class="mw-redirect" title="Digital PCR">Digital PCR</a> (dPCR): used to measure the quantity of a target DNA sequence in a DNA sample. The DNA sample is highly diluted so that after running many PCRs in parallel, some of them do not receive a single molecule of the target DNA. The target DNA concentration is calculated using the proportion of negative outcomes. Hence the name 'digital PCR'.</li> <li><a href="/wiki/Helicase-dependent_amplification" title="Helicase-dependent amplification">Helicase-dependent amplification</a>: similar to traditional PCR, but uses a constant temperature rather than cycling through denaturation and annealing/extension cycles. <a href="/wiki/DNA_helicase" class="mw-redirect" title="DNA helicase">DNA helicase</a>, an enzyme that unwinds DNA, is used in place of thermal denaturation.<a href="#cite_note-54">[54]</a></li> <li><a href="/wiki/Hot_start_PCR" title="Hot start PCR">Hot start PCR</a>: a technique that reduces non-specific amplification during the initial set up stages of the PCR. It may be performed manually by heating the reaction components to the denaturation temperature (e.g., 95 °C) before adding the polymerase.<a href="#cite_note-general_hot_start-55">[55]</a> Specialized enzyme systems have been developed that inhibit the polymerase's activity at ambient temperature, either by the binding of an <a href="/wiki/Antibody" title="Antibody">antibody</a><a href="#cite_note-antibody_hot_start-13">[13]</a><a href="#cite_note-antibody_hot_start_2-56">[56]</a> or by the presence of covalently bound inhibitors that dissociate only after a high-temperature activation step. Hot-start/cold-finish PCR is achieved with new hybrid polymerases that are inactive at ambient temperature and are instantly activated at elongation temperature.</li> <li><a href="/wiki/In_silico_PCR" title="In silico PCR">In silico PCR</a> (digital PCR, virtual PCR, electronic PCR, e-PCR) refers to computational tools used to calculate theoretical polymerase chain reaction results using a given set of <a href="/wiki/Primer_(molecular_biology)" title="Primer (molecular biology)">primers</a> (<a href="/wiki/Hybridization_probe" title="Hybridization probe">probes</a>) to amplify <a href="/wiki/DNA" title="DNA">DNA</a> sequences from a sequenced <a href="/wiki/Genome" title="Genome">genome</a> or <a href="/wiki/Transcriptome" title="Transcriptome">transcriptome</a>. In silico PCR was proposed as an educational tool for molecular biology.<a href="#cite_note-57">[57]</a></li> <li>Intersequence-specific PCR (ISSR): a PCR method for DNA fingerprinting that amplifies regions between simple sequence repeats to produce a unique fingerprint of amplified fragment lengths.<a href="#cite_note-58">[58]</a></li> <li><a href="/wiki/Inverse_polymerase_chain_reaction" title="Inverse polymerase chain reaction">Inverse PCR</a>: is commonly used to identify the flanking sequences around <a href="/wiki/Genomic" class="mw-redirect" title="Genomic">genomic</a> inserts. It involves a series of <a href="/wiki/Restriction_digest" title="Restriction digest">DNA digestions</a> and <a href="/wiki/Self_ligation" class="mw-redirect" title="Self ligation">self ligation</a>, resulting in known sequences at either end of the unknown sequence.<a href="#cite_note-59">[59]</a></li> <li>Ligation-mediated PCR: uses small DNA linkers ligated to the DNA of interest and multiple primers annealing to the DNA linkers; it has been used for <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a>, <a href="/wiki/Genome_walking" class="mw-redirect" title="Genome walking">genome walking</a>, and <a href="/wiki/DNA_footprinting" title="DNA footprinting">DNA footprinting</a>.<a href="#cite_note-Mueller_and_Wold-60">[60]</a></li> <li><a href="/wiki/Methylation-specific_PCR" class="mw-redirect" title="Methylation-specific PCR">Methylation-specific PCR</a> (MSP): developed by <a href="/wiki/Stephen_Baylin" class="mw-redirect" title="Stephen Baylin">Stephen Baylin</a> and <a href="/wiki/James_G._Herman" title="James G. Herman">James G. Herman</a> at the Johns Hopkins School of Medicine,<a href="#cite_note-61">[61]</a> and is used to detect methylation of CpG islands in genomic DNA. DNA is first treated with sodium bisulfite, which converts unmethylated cytosine bases to uracil, which is recognized by PCR primers as thymine. Two PCRs are then carried out on the modified DNA, using primer sets identical except at any CpG islands within the primer sequences. At these points, one primer set recognizes DNA with cytosines to amplify methylated DNA, and one set recognizes DNA with uracil or thymine to amplify unmethylated DNA. MSP using qPCR can also be performed to obtain quantitative rather than qualitative information about methylation.</li> <li>Miniprimer PCR: uses a thermostable polymerase (S-Tbr) that can extend from short primers ("smalligos") as short as 9 or 10 nucleotides. This method permits PCR targeting to smaller primer binding regions, and is used to amplify conserved DNA sequences, such as the 16S (or eukaryotic 18S) rRNA gene.<a href="#cite_note-62">[62]</a></li> <li><a href="/wiki/Multiplex_ligation-dependent_probe_amplification" title="Multiplex ligation-dependent probe amplification">Multiplex ligation-dependent probe amplification</a> (MLPA): permits amplifying multiple targets with a single primer pair, thus avoiding the resolution limitations of multiplex PCR (see below).</li> <li><a href="/wiki/Multiplex-PCR" class="mw-redirect" title="Multiplex-PCR">Multiplex-PCR</a>: consists of multiple primer sets within a single PCR mixture to produce <a href="/wiki/Amplicon" title="Amplicon">amplicons</a> of varying sizes that are specific to different DNA sequences. By targeting multiple genes at once, additional information may be gained from a single test-run that otherwise would require several times the reagents and more time to perform. Annealing temperatures for each of the primer sets must be optimized to work correctly within a single reaction, and amplicon sizes. That is, their base pair length should be different enough to form distinct bands when visualized by <a href="/wiki/Gel_electrophoresis" title="Gel electrophoresis">gel electrophoresis</a>.</li> <li>Nanoparticle-assisted PCR (nanoPCR): some nanoparticles (NPs) can enhance the efficiency of PCR (thus being called nanoPCR), and some can even outperform the original PCR enhancers. It was reported that quantum dots (QDs) can improve PCR specificity and efficiency. Single-walled carbon nanotubes (SWCNTs) and multi-walled carbon nanotubes (MWCNTs) are efficient in enhancing the amplification of long PCR. Carbon nanopowder (CNP) can improve the efficiency of repeated PCR and long PCR, while <a href="/wiki/Zinc_oxide" title="Zinc oxide">zinc oxide</a>, <a href="/wiki/Titanium_dioxide" title="Titanium dioxide">titanium dioxide</a> and Ag NPs were found to increase the PCR yield. Previous data indicated that non-metallic NPs retained acceptable amplification fidelity. Given that many NPs are capable of enhancing PCR efficiency, it is clear that there is likely to be great potential for nanoPCR technology improvements and product development.<a href="#cite_note-63">[63]</a><a href="#cite_note-64">[64]</a></li> <li><a href="/wiki/Nested_PCR" class="mw-redirect" title="Nested PCR">Nested PCR</a>: increases the specificity of DNA amplification, by reducing background due to non-specific amplification of DNA. Two sets of primers are used in two successive PCRs. In the first reaction, one pair of primers is used to generate DNA products, which besides the intended target, may still consist of non-specifically amplified DNA fragments. The product(s) are then used in a second PCR with a set of primers whose binding sites are completely or partially different from and located 3' of each of the primers used in the first reaction. Nested PCR is often more successful in specifically amplifying long DNA fragments than conventional PCR, but it requires more detailed knowledge of the target sequences.</li> <li><a href="/wiki/Overlap-extension_PCR" class="mw-redirect" title="Overlap-extension PCR">Overlap-extension PCR</a> or Splicing by overlap extension (SOEing) : a <a href="/wiki/Genetic_engineering" title="Genetic engineering">genetic engineering</a> technique that is used to splice together two or more DNA fragments that contain complementary sequences. It is used to join DNA pieces containing genes, regulatory sequences, or mutations; the technique enables creation of specific and long DNA constructs. It can also introduce deletions, insertions or point mutations into a DNA sequence.<a href="#cite_note-65">[65]</a><a href="#cite_note-66">[66]</a></li> <li>PAN-AC: uses isothermal conditions for amplification, and may be used in living cells.<a href="#cite_note-DavidTurlotte1998-67">[67]</a><a href="#cite_note-68">[68]</a></li> <li>PAN-PCR: A computational method for designing bacterium typing assays based on whole genome sequence data.<a href="#cite_note-69">[69]</a></li> <li><a href="/wiki/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">Quantitative PCR</a> (qPCR): used to measure the quantity of a target sequence (commonly in real-time). It quantitatively measures starting amounts of DNA, cDNA, or RNA. Quantitative PCR is commonly used to determine whether a DNA sequence is present in a sample and the number of its copies in the sample. Quantitative PCR has a very high degree of precision. Quantitative PCR methods use fluorescent dyes, such as Sybr Green, EvaGreen or <a href="/wiki/Fluorophore" title="Fluorophore">fluorophore</a>-containing DNA probes, such as <a href="/wiki/TaqMan" title="TaqMan">TaqMan</a>, to measure the amount of amplified product in real time. It is also sometimes abbreviated to <a href="/wiki/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">RT-PCR</a> (real-time PCR) but this abbreviation should be used only for <a href="/wiki/Reverse_transcription_polymerase_chain_reaction" title="Reverse transcription polymerase chain reaction">reverse transcription PCR</a>. qPCR is the appropriate contractions for <a href="/wiki/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">quantitative PCR</a> (real-time PCR).</li> <li><a href="/wiki/Reverse_Complement_Polymerase_Chain_Reaction" class="mw-redirect" title="Reverse Complement Polymerase Chain Reaction">Reverse Complement PCR</a> (RC-PCR): Allows the addition of functional domains or sequences of choice to be appended independently to either end of the generated amplicon in a single closed tube reaction. This method generates target specific primers within the reaction by the interaction of universal primers (which contain the desired sequences or domains to be appended) and RC probes.</li> <li>Reverse Transcription PCR (<a href="/wiki/RT-PCR" class="mw-redirect" title="RT-PCR">RT-PCR</a>): for amplifying DNA from RNA. <a href="/wiki/Reverse_transcriptase" title="Reverse transcriptase">Reverse transcriptase</a> reverse transcribes <a href="/wiki/RNA" title="RNA">RNA</a> into <a href="/wiki/CDNA" class="mw-redirect" title="CDNA">cDNA</a>, which is then amplified by PCR. RT-PCR is widely used in <a href="/wiki/Expression_profiling" class="mw-redirect" title="Expression profiling">expression profiling</a>, to determine the expression of a gene or to identify the sequence of an RNA transcript, including transcription start and termination sites. If the genomic DNA sequence of a gene is known, RT-PCR can be used to map the location of <a href="/wiki/Exons" class="mw-redirect" title="Exons">exons</a> and <a href="/wiki/Introns" class="mw-redirect" title="Introns">introns</a> in the gene. The 5' end of a gene (corresponding to the transcription start site) is typically identified by <a href="/wiki/RACE_(biology)" class="mw-redirect" title="RACE (biology)">RACE-PCR</a> (Rapid Amplification of cDNA Ends).</li> <li><a href="/wiki/RNase_H-dependent_PCR" title="RNase H-dependent PCR">RNase H-dependent PCR</a> (rhPCR): a modification of PCR that utilizes primers with a 3' extension block that can be removed by a thermostable RNase HII enzyme. This system reduces primer-dimers and allows for multiplexed reactions to be performed with higher numbers of primers.<a href="#cite_note-dobosy_2011-70">[70]</a></li> <li>Single specific primer-PCR (SSP-PCR): allows the amplification of double-stranded DNA even when the sequence information is available at one end only. This method permits amplification of genes for which only a partial sequence information is available, and allows unidirectional genome walking from known into unknown regions of the chromosome.<a href="#cite_note-71">[71]</a></li> <li>Solid Phase PCR: encompasses multiple meanings, including <a href="/wiki/Polony_(biology)" title="Polony (biology)">Polony Amplification</a> (where PCR colonies are derived in a gel matrix, for example), Bridge PCR<a href="#cite_note-72">[72]</a> (primers are covalently linked to a solid-support surface), conventional Solid Phase PCR (where Asymmetric PCR is applied in the presence of solid support bearing primer with sequence matching one of the aqueous primers) and Enhanced Solid Phase PCR<a href="#cite_note-73">[73]</a> (where conventional Solid Phase PCR can be improved by employing high Tm and nested solid support primer with optional application of a thermal 'step' to favour solid support priming).</li> <li>Suicide PCR: typically used in <a href="/wiki/Paleogenetics" title="Paleogenetics">paleogenetics</a> or other studies where avoiding false positives and ensuring the specificity of the amplified fragment is the highest priority. It was originally described in a study to verify the presence of the microbe <a href="/wiki/Yersinia_pestis" title="Yersinia pestis">Yersinia pestis</a> in dental samples obtained from 14th Century graves of people supposedly killed by the plague during the medieval <a href="/wiki/Black_Death" title="Black Death">Black Death</a> epidemic.<a href="#cite_note-74">[74]</a> The method prescribes the use of any primer combination only once in a PCR (hence the term "suicide"), which should never have been used in any positive control PCR reaction, and the primers should always target a genomic region never amplified before in the lab using this or any other set of primers. This ensures that no contaminating DNA from previous PCR reactions is present in the lab, which could otherwise generate false positives.</li> <li>Thermal asymmetric interlaced PCR (TAIL-PCR): for isolation of an unknown sequence flanking a known sequence. Within the known sequence, TAIL-PCR uses a nested pair of primers with differing annealing temperatures; a degenerate primer is used to amplify in the other direction from the unknown sequence.<a href="#cite_note-75">[75]</a></li> <li><a href="/wiki/Touchdown_PCR" class="mw-redirect" title="Touchdown PCR">Touchdown PCR</a> (Step-down PCR): a variant of PCR that aims to reduce nonspecific background by gradually lowering the annealing temperature as PCR cycling progresses. The annealing temperature at the initial cycles is usually a few degrees (3–5 °C) above the Tm of the primers used, while at the later cycles, it is a few degrees (3–5 °C) below the primer Tm. The higher temperatures give greater specificity for primer binding, and the lower temperatures permit more efficient amplification from the specific products formed during the initial cycles.<a href="#cite_note-76">[76]</a></li> <li>Universal Fast Walking: for genome walking and genetic fingerprinting using a more specific 'two-sided' PCR than conventional 'one-sided' approaches (using only one gene-specific primer and one general primer—which can lead to artefactual 'noise')<a href="#cite_note-Myrick_and_Gelbart-77">[77]</a> by virtue of a mechanism involving lariat structure formation. Streamlined derivatives of UFW are LaNe RAGE (lariat-dependent nested PCR for rapid amplification of genomic DNA ends),<a href="#cite_note-lane_rage-78">[78]</a> 5'RACE LaNe<a href="#cite_note-Parkb-79">[79]</a> and 3'RACE LaNe.<a href="#cite_note-Park-80">[80]</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="History">History</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=15" title="Edit section: History">edit</a>]</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/History_of_polymerase_chain_reaction" title="History of polymerase chain reaction">History of polymerase chain reaction</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Primers_RevComp.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Primers_RevComp.svg/220px-Primers_RevComp.svg.png" decoding="async" width="220" height="115" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Primers_RevComp.svg/330px-Primers_RevComp.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/91/Primers_RevComp.svg/440px-Primers_RevComp.svg.png 2x" data-file-width="1028" data-file-height="536" /></a><figcaption>Diagrammatic representation of an example primer pair. The use of primers in an in vitro assay to allow DNA synthesis was a major innovation that allowed the development of PCR.</figcaption></figure> The heat-resistant enzymes that are a key component in polymerase chain reaction were discovered in the 1960s as a product of a microbial life form that lived in the superheated waters of <a href="/wiki/Yellowstone_National_Park" title="Yellowstone National Park">Yellowstone</a>'s Mushroom Spring.<a href="#cite_note-81">[81]</a> A 1971 paper in the <a href="/wiki/Journal_of_Molecular_Biology" title="Journal of Molecular Biology">Journal of Molecular Biology</a> by <a href="/wiki/Kjell_Kleppe" title="Kjell Kleppe">Kjell Kleppe</a> and co-workers in the laboratory of <a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">H. Gobind Khorana</a> first described a method of using an enzymatic assay to replicate a short DNA template with primers in vitro.<a href="#cite_note-82">[82]</a> However, this early manifestation of the basic PCR principle did not receive much attention at the time and the invention of the polymerase chain reaction in 1983 is generally credited to <a href="/wiki/Kary_Mullis" title="Kary Mullis">Kary Mullis</a>.<a href="#cite_note-83">[83]</a>[<a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">page needed</a>] <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Baby_Blue_-_a_prototype_polymerase_chain_reaction_(PCR),_c_1986._(9663810586).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg/220px-Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg" decoding="async" width="220" height="142" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/db/Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg/330px-Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/db/Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg/440px-Baby_Blue_-_a_prototype_polymerase_chain_reaction_%28PCR%29%2C_c_1986._%289663810586%29.jpg 2x" data-file-width="1250" data-file-height="806" /></a><figcaption>"Baby Blue", a 1986 prototype machine for doing PCR</figcaption></figure> When Mullis developed the PCR in 1983, he was working in <a href="/wiki/Emeryville,_California" title="Emeryville, California">Emeryville</a>, California for <a href="/wiki/Cetus_Corporation" title="Cetus Corporation">Cetus Corporation</a>, one of the first <a href="/wiki/Biotechnology" title="Biotechnology">biotechnology</a> companies, where he was responsible for synthesizing short chains of DNA. Mullis has written that he conceived the idea for PCR while cruising along the <a href="/wiki/California_State_Route_1" title="California State Route 1">Pacific Coast Highway</a> one night in his car.<a href="#cite_note-Mullis-84">[84]</a> He was playing in his mind with a new way of analyzing changes (mutations) in DNA when he realized that he had instead invented a method of amplifying any DNA region through repeated cycles of duplication driven by DNA polymerase. In <a href="/wiki/Scientific_American" title="Scientific American">Scientific American</a>, Mullis summarized the procedure: "Beginning with a single molecule of the genetic material DNA, the PCR can generate 100 billion similar molecules in an afternoon. The reaction is easy to execute. It requires no more than a test tube, a few simple reagents, and a source of heat."<a href="#cite_note-85">[85]</a> DNA fingerprinting was first used for <a href="/wiki/Paternity_testing" class="mw-redirect" title="Paternity testing">paternity testing</a> in 1988.<a href="#cite_note-86">[86]</a> Mullis has credited his use of <a href="/wiki/LSD" title="LSD">LSD</a> as integral to his development of PCR: "Would I have invented PCR if I hadn't taken LSD? I seriously doubt it. I could sit on a DNA molecule and watch the polymers go by. I learnt that partly on psychedelic drugs."<a href="#cite_note-87">[87]</a> Mullis and biochemist <a href="/wiki/Michael_Smith_(chemist)" title="Michael Smith (chemist)">Michael Smith</a>, who had developed other essential ways of manipulating DNA,<a href="#cite_note-NobelPrize-1">[1]</a> were jointly awarded the <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a> in 1993, seven years after Mullis and his colleagues at Cetus first put his proposal to practice.<a href="#cite_note-Kary_Mullis_Nobel_Lecture-88">[88]</a> Mullis's 1985 paper with R. K. Saiki and H. A. Erlich, "Enzymatic Amplification of β-globin Genomic Sequences and Restriction Site Analysis for Diagnosis of Sickle Cell Anemia"—the polymerase chain reaction invention (PCR)—was honored by a Citation for Chemical Breakthrough Award from the Division of History of Chemistry of the American Chemical Society in 2017.<a href="#cite_note-breakthrough-89">[89]</a><a href="#cite_note-Saiki1-2">[2]</a> At the core of the PCR method is the use of a suitable <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerase</a> able to withstand the high temperatures of >90 °C (194 °F) required for separation of the two DNA strands in the <a href="/wiki/DNA_double_helix" class="mw-redirect" title="DNA double helix">DNA double helix</a> after each <a href="/wiki/DNA_replication" title="DNA replication">replication</a> cycle. The DNA polymerases initially employed for <a href="/wiki/In_vitro" title="In vitro">in vitro</a> experiments presaging PCR were unable to withstand these high temperatures.<a href="#cite_note-Saiki1-2">[2]</a> So the early procedures for DNA replication were very inefficient and time-consuming, and required large amounts of DNA polymerase and continuous handling throughout the process. The discovery in 1976 of <a href="/wiki/Taq_polymerase" title="Taq polymerase">Taq polymerase</a>—a DNA polymerase purified from the <a href="/wiki/Thermophile" title="Thermophile">thermophilic bacterium</a>, <a href="/wiki/Thermus_aquaticus" title="Thermus aquaticus">Thermus aquaticus</a>, which naturally lives in hot (50 to 80 °C (122 to 176 °F)) environments<a href="#cite_note-Chien_et_al.-14">[14]</a> such as hot springs—paved the way for dramatic improvements of the PCR method. The DNA polymerase isolated from T. aquaticus is stable at high temperatures remaining active even after DNA denaturation,<a href="#cite_note-Lawyer_et_al.-15">[15]</a> thus obviating the need to add new DNA polymerase after each cycle.<a href="#cite_note-Saiki2-3">[3]</a> This allowed an automated thermocycler-based process for DNA amplification. <div class="mw-heading mw-heading3"><h3 id="Patent_disputes">Patent disputes</h3>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=16" title="Edit section: Patent disputes">edit</a>]</div> The PCR technique was patented by <a href="/wiki/Kary_Mullis" title="Kary Mullis">Kary Mullis</a> and assigned to <a href="/wiki/Cetus_Corporation" title="Cetus Corporation">Cetus Corporation</a>, where Mullis worked when he invented the technique in 1983. The Taq polymerase enzyme was also covered by patents. There have been several high-profile lawsuits related to the technique, including an unsuccessful lawsuit<a href="#cite_note-90">[90]</a> brought by <a href="/wiki/DuPont" title="DuPont">DuPont</a>. The Swiss pharmaceutical company <a href="/wiki/Hoffmann-La_Roche" class="mw-redirect" title="Hoffmann-La Roche">Hoffmann-La Roche</a> purchased the rights to the patents in 1992. The last of the commercial PCR patents expired in 2017.<a href="#cite_note-91">[91]</a> A related patent battle over the Taq polymerase enzyme is still ongoing[<a href="/wiki/Wikipedia:Manual_of_Style/Dates_and_numbers#Chronological_items" title="Wikipedia:Manual of Style/Dates and numbers">as of?</a>] in several jurisdictions around the world between Roche and <a href="/wiki/Promega" title="Promega">Promega</a>. The legal arguments have extended beyond the lives of the original PCR and Taq polymerase patents, which expired on 28 March 2005.<a href="#cite_note-92">[92]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=17" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1266661725">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 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testing">COVID-19 testing</a></li> <li><a href="/wiki/DNA_spiking" title="DNA spiking">DNA spiking</a></li> <li><a href="/wiki/Loop-mediated_isothermal_amplification" title="Loop-mediated isothermal amplification">Loop-mediated isothermal amplification</a></li> <li><a href="/wiki/Selector_technique" title="Selector technique">Selector technique</a></li> <li><a href="/wiki/Thermus_thermophilus#Applications" title="Thermus thermophilus">Thermus thermophilus</a></li> <li><a href="/wiki/Pfu_DNA_polymerase" title="Pfu DNA polymerase">Pfu DNA polymerase</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Polymerase_chain_reaction&action=edit&section=18" title="Edit section: References">edit</a>]</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-NobelPrize-1">^ <a href="#cite_ref-NobelPrize_1-0">a</a> <a href="#cite_ref-NobelPrize_1-1">b</a> <style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output 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Proceedings of the National Academy of Sciences of the United States of America. 91 (12): 5695–99. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1994PNAS...91.5695C">1994PNAS...91.5695C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.91.12.5695">10.1073/pnas.91.12.5695</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC44063">44063</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/8202550">8202550</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Effective+amplification+of+long+targets+from+cloned+inserts+and+human+genomic+DNA&rft.volume=91&rft.issue=12&rft.pages=%3Cspan+class%3D%22nowrap%22%3E5695-%3C%2Fspan%3E99&rft.date=1994-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC44063%23id-name%3DPMC&rft_id=info%3Apmid%2F8202550&rft_id=info%3Adoi%2F10.1073%2Fpnas.91.12.5695&rft_id=info%3Abibcode%2F1994PNAS...91.5695C&rft.aulast=Cheng&rft.aufirst=S&rft.au=Fockler%2C+C&rft.au=Barnes%2C+WM&rft.au=Higuchi%2C+R&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC44063&rfr_id=info%3Asid%2Fen.wikipedia.org%3APolymerase+chain+reaction" class="Z3988"> </li> <li id="cite_note-7"><a href="#cite_ref-7">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarrMoore2012" class="citation journal cs1">Carr AC, Moore SD (2012). 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Cold Spring Harbor, N.Y.: Cold Spring Harbor Laboratory Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87969-576-7" title="Special:BookSources/978-0-87969-576-7"><bdi>978-0-87969-576-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Molecular+Cloning%3A+A+Laboratory+Manual&rft.place=Cold+Spring+Harbor%2C+N.Y.&rft.edition=third&rft.pub=Cold+Spring+Harbor+Laboratory+Press&rft.date=2001&rft.isbn=978-0-87969-576-7&rft.au=Joseph+Sambrook&rft.au=David+W.+Russel&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmolecularcloning0000samb_p7p5&rfr_id=info%3Asid%2Fen.wikipedia.org%3APolymerase+chain+reaction" class="Z3988"> Chapter 8: In vitro Amplification of DNA by the Polymerase Chain Reaction </li> <li id="cite_note-9"><a href="#cite_ref-9">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/probe/docs/techpcr/">"Polymerase Chain Reaction (PCR)"</a>. 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reaction"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Polymerase_chain_reaction" title="Special:EditPage/Template:Polymerase chain reaction"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Polymerase_chain_reaction_techniques218" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Polymerase chain reaction</a> techniques</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Procedure</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>Initialization</li> <li>Cycle <ul><li><a href="/wiki/Denaturation_(biochemistry)" title="Denaturation (biochemistry)">Denaturation</a></li> <li><a href="/wiki/Annealing_(biology)" class="mw-redirect" title="Annealing (biology)">Annealing</a></li> <li><a href="/wiki/Transcription_(biology)#Elongation" title="Transcription (biology)">Elongation</a></li></ul></li> <li>Final elongation</li> <li>Final hold</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Polymerase" title="Polymerase">Polymerase</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/Klenow_fragment" title="Klenow fragment">Klenow</a></li> <li><a href="/w/index.php?title=T4_DNA_polymerase&action=edit&redlink=1" class="new" title="T4 DNA polymerase (page does not exist)">T4</a> and <a href="/wiki/T7_DNA_polymerase" title="T7 DNA polymerase">T7</a></li> <li><a href="/wiki/Taq_polymerase" title="Taq polymerase">Taq</a></li> <li><a href="/wiki/Tth_DNA_polymerase" class="mw-redirect" title="Tth DNA polymerase">Tth</a></li> <li><a href="/wiki/Pfu_DNA_polymerase" title="Pfu DNA polymerase">Pfu</a></li> <li><a href="/wiki/Vent_DNA_polymerase" title="Vent DNA polymerase">Vent</a></li> <li><a href="/wiki/Pwo_DNA_polymerase" title="Pwo DNA polymerase">Pwo</a></li> <li><a href="/wiki/Tli_polymerase" class="mw-redirect" title="Tli polymerase">Tli</a></li> <li><a href="/w/index.php?title=Tfu_DNA_polymerase&action=edit&redlink=1" class="new" title="Tfu DNA polymerase (page does not exist)">Tfu</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Polymerase_chain_reaction_optimization" title="Polymerase chain reaction optimization">Optimization</a> and <a href="/wiki/Variants_of_PCR" title="Variants of PCR">variants</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/Quantitative_PCR" class="mw-redirect" title="Quantitative PCR">Quantitative polymerase chain reaction</a> (qPCR)</li> <li><a href="/wiki/Reverse_transcription_polymerase_chain_reaction" title="Reverse transcription polymerase chain reaction">Reverse transcription polymerase chain reaction</a> (RT-PCR)</li> <li><a href="/wiki/Inverse_polymerase_chain_reaction" title="Inverse polymerase chain reaction">Inverse polymerase chain reaction</a> (inverse PCR)</li> <li><a href="/wiki/Nested_polymerase_chain_reaction" title="Nested polymerase chain reaction">Nested polymerase chain reaction</a> (nested PCR)</li> <li><a href="/wiki/Touchdown_polymerase_chain_reaction" title="Touchdown polymerase chain reaction">Touchdown polymerase chain reaction</a></li> <li><a href="/wiki/Hot_start_PCR" title="Hot start PCR">Hot start PCR</a></li> <li><a href="/wiki/Overlap_extension_polymerase_chain_reaction" title="Overlap extension polymerase chain reaction">Overlap extension polymerase chain reaction</a> (OE-PCR)</li> <li><a href="/wiki/Multiplex_polymerase_chain_reaction" title="Multiplex polymerase chain reaction">Multiplex polymerase chain reaction</a> (multiplex PCR)</li> <li><a href="/wiki/Multiplex_ligation-dependent_probe_amplification" title="Multiplex ligation-dependent probe amplification">Multiplex ligation-dependent probe amplification</a> (MLPA)</li> <li><a href="/wiki/COLD-PCR" title="COLD-PCR">Co-amplification at lower denaturation temperature PCR</a> (COLD-PCR)</li> <li><a href="/wiki/Digital_polymerase_chain_reaction" title="Digital polymerase chain reaction">Digital polymerase chain reaction</a> (dePCR)</li> <li><a href="/wiki/Random_amplification_of_polymorphic_DNA" title="Random amplification of polymorphic DNA">Random amplification of polymorphic DNA</a> (RAPD)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_polymerase_chain_reaction" title="History of polymerase chain reaction">History</a> and people</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/Kary_Mullis" title="Kary Mullis">Kary Mullis</a></li> <li><a href="/wiki/Kjell_Kleppe" title="Kjell Kleppe">Kjell Kleppe</a></li> <li><a href="/w/index.php?title=Alice_Chien&action=edit&redlink=1" class="new" title="Alice Chien (page does not exist)">Alice Chien</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 .portal-bar-content{display:flex;flex-flow:row wrap;flex:0 1 auto;padding:0.15em 0;column-gap:1em;align-items:baseline;margin:0;list-style:none}.mw-parser-output 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