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<span>Biosynthesis</span> </div> </a> <ul id="toc-Biosynthesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Pseudopeptidoglycan" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Pseudopeptidoglycan"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Pseudopeptidoglycan</span> </div> </a> <ul id="toc-Pseudopeptidoglycan-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Recognition_by_immune_system" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Recognition_by_immune_system"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Recognition by immune system</span> </div> </a> <button aria-controls="toc-Recognition_by_immune_system-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Recognition by immune system subsection</span> </button> <ul id="toc-Recognition_by_immune_system-sublist" class="vector-toc-list"> <li id="toc-Receptors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Receptors</span> </div> </a> <ul id="toc-Receptors-sublist" class="vector-toc-list"> <li id="toc-Peptidoglycan_recognition_proteins" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Peptidoglycan_recognition_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.1</span> <span>Peptidoglycan recognition proteins</span> </div> </a> <ul id="toc-Peptidoglycan_recognition_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NOD-like_receptors" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#NOD-like_receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.2</span> <span>NOD-like receptors</span> </div> </a> <ul id="toc-NOD-like_receptors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-C-type_lectin_receptors_(CLRs)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#C-type_lectin_receptors_(CLRs)"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.3</span> <span>C-type lectin receptors (CLRs)</span> </div> </a> <ul id="toc-C-type_lectin_receptors_(CLRs)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Toll-like_receptors" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Toll-like_receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1.4</span> <span>Toll-like receptors</span> </div> </a> <ul id="toc-Toll-like_receptors-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-As_vaccine_or_adjuvant" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#As_vaccine_or_adjuvant"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>As vaccine or adjuvant</span> </div> </a> <ul id="toc-As_vaccine_or_adjuvant-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inhibition_and_degradation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Inhibition_and_degradation"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Inhibition and degradation</span> </div> </a> <ul id="toc-Inhibition_and_degradation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Peptidoglycan</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 44 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-44" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">44 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A8%D8%A8%D8%AA%D9%8A%D8%AF%D9%88%D8%AC%D9%84%D9%8A%D9%83%D8%A7%D9%86" title="ببتيدوجليكان – Arabic" lang="ar" hreflang="ar" data-title="ببتيدوجليكان" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Bosnian" lang="bs" hreflang="bs" data-title="Peptidoglikan" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Peptidoglic%C3%A0" title="Peptidoglicà – Catalan" lang="ca" hreflang="ca" data-title="Peptidoglicà" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Peptidoglykan" title="Peptidoglykan – Czech" lang="cs" hreflang="cs" data-title="Peptidoglykan" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Peptidoglycan" title="Peptidoglycan – Danish" lang="da" hreflang="da" data-title="Peptidoglycan" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Peptidoglycane" title="Peptidoglycane – German" lang="de" hreflang="de" data-title="Peptidoglycane" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Peptidogl%C3%BCkaan" title="Peptidoglükaan – Estonian" lang="et" hreflang="et" data-title="Peptidoglükaan" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%A0%CE%B5%CF%80%CF%84%CE%B9%CE%B4%CE%BF%CE%B3%CE%BB%CF%85%CE%BA%CE%AC%CE%BD%CE%B7" title="Πεπτιδογλυκάνη – Greek" lang="el" hreflang="el" data-title="Πεπτιδογλυκάνη" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Peptidoglucano" title="Peptidoglucano – Spanish" lang="es" hreflang="es" data-title="Peptidoglucano" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Peptidoglikano" title="Peptidoglikano – Basque" lang="eu" hreflang="eu" data-title="Peptidoglikano" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%BE%D9%BE%D8%AA%DB%8C%D8%AF%D9%88%DA%AF%D9%84%DB%8C%DA%A9%D8%A7%D9%86" title="پپتیدوگلیکان – Persian" lang="fa" hreflang="fa" data-title="پپتیدوگلیکان" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Peptidoglycane" title="Peptidoglycane – French" lang="fr" hreflang="fr" data-title="Peptidoglycane" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Peipt%C3%ADdiglioc%C3%A1n" title="Peiptídigliocán – Irish" lang="ga" hreflang="ga" data-title="Peiptídigliocán" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Peptidoglicano" title="Peptidoglicano – Galician" lang="gl" hreflang="gl" data-title="Peptidoglicano" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%ED%8E%A9%ED%8B%B0%EB%8F%84%EA%B8%80%EB%A6%AC%EC%B9%B8" title="펩티도글리칸 – Korean" lang="ko" hreflang="ko" data-title="펩티도글리칸" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Croatian" lang="hr" hreflang="hr" data-title="Peptidoglikan" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Indonesian" lang="id" hreflang="id" data-title="Peptidoglikan" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Peptidoglicano" title="Peptidoglicano – Italian" lang="it" hreflang="it" data-title="Peptidoglicano" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A4%D7%A4%D7%98%D7%99%D7%93%D7%95%D7%92%D7%9C%D7%99%D7%A7%D7%9F" title="פפטידוגליקן – Hebrew" lang="he" hreflang="he" data-title="פפטידוגליקן" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Javanese" lang="jv" hreflang="jv" data-title="Peptidoglikan" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%9F%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D0%BE%D0%B3%D0%BB%D0%B8%D0%BA%D0%B0%D0%BD" title="Пептидогликан – Kazakh" lang="kk" hreflang="kk" data-title="Пептидогликан" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Peptidoglikanas" title="Peptidoglikanas – Lithuanian" lang="lt" hreflang="lt" data-title="Peptidoglikanas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Peptidoglik%C3%A1n" title="Peptidoglikán – Hungarian" lang="hu" hreflang="hu" data-title="Peptidoglikán" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9F%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D0%BE%D0%B3%D0%BB%D0%B8%D0%BA%D0%B0%D0%BD" title="Пептидогликан – Macedonian" lang="mk" hreflang="mk" data-title="Пептидогликан" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Malay" lang="ms" hreflang="ms" data-title="Peptidoglikan" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Peptidoglycaan" title="Peptidoglycaan – Dutch" lang="nl" hreflang="nl" data-title="Peptidoglycaan" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%9A%E3%83%97%E3%83%81%E3%83%89%E3%82%B0%E3%83%AA%E3%82%AB%E3%83%B3" title="ペプチドグリカン – Japanese" lang="ja" hreflang="ja" data-title="ペプチドグリカン" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Peptidoglykan" title="Peptidoglykan – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Peptidoglykan" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Peptidoglykan" title="Peptidoglykan – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Peptidoglykan" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Peptydoglikan" title="Peptydoglikan – Polish" lang="pl" hreflang="pl" data-title="Peptydoglikan" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Peptidoglicano" title="Peptidoglicano – Portuguese" lang="pt" hreflang="pt" data-title="Peptidoglicano" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Peptidoglican" title="Peptidoglican – Romanian" lang="ro" hreflang="ro" data-title="Peptidoglican" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D0%BE%D0%B3%D0%BB%D0%B8%D0%BA%D0%B0%D0%BD" title="Пептидогликан – Russian" lang="ru" hreflang="ru" data-title="Пептидогликан" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Peptidoglycan" title="Peptidoglycan – Simple English" lang="en-simple" hreflang="en-simple" data-title="Peptidoglycan" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Slovenian" lang="sl" hreflang="sl" data-title="Peptidoglikan" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Serbian" lang="sr" hreflang="sr" data-title="Peptidoglikan" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Peptidoglikan" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Mureiini" title="Mureiini – Finnish" lang="fi" hreflang="fi" data-title="Mureiini" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Peptidoglykan" title="Peptidoglykan – Swedish" lang="sv" hreflang="sv" data-title="Peptidoglykan" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%80%E0%B8%9B%E0%B8%9A%E0%B8%97%E0%B8%B4%E0%B9%82%E0%B8%94%E0%B9%84%E0%B8%81%E0%B8%A5%E0%B9%81%E0%B8%84%E0%B8%99" title="เปบทิโดไกลแคน – Thai" lang="th" hreflang="th" data-title="เปบทิโดไกลแคน" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Peptidoglikan" title="Peptidoglikan – Turkish" lang="tr" hreflang="tr" data-title="Peptidoglikan" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9F%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D0%BE%D0%B3%D0%BB%D1%96%D0%BA%D0%B0%D0%BD" title="Пептидоглікан – Ukrainian" lang="uk" hreflang="uk" data-title="Пептидоглікан" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Murein" title="Murein – Vietnamese" lang="vi" hreflang="vi" data-title="Murein" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%82%BD%E8%81%9A%E7%B3%96" title="肽聚糖 – Chinese" lang="zh" hreflang="zh" data-title="肽聚糖" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div 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<div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Polymer in bacterial cell walls</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">Not to be confused with <a href="/wiki/Glycopeptide" title="Glycopeptide">glycopeptide</a>, <a href="/wiki/Proteoglycan" title="Proteoglycan">proteoglycan</a>, or <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoprotein</a>.</div> <p><b>Peptidoglycan</b> or <b>murein</b> is a unique large macromolecule, a <a href="/wiki/Polysaccharide" title="Polysaccharide">polysaccharide</a>, consisting of sugars and <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> that forms a mesh-like layer (sacculus) that surrounds the <a href="/wiki/Bacterial" class="mw-redirect" title="Bacterial">bacterial</a> cytoplasmic membrane.<sup id="cite_ref-Madigan-2015_1-0" class="reference"><a href="#cite_note-Madigan-2015-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The sugar component consists of alternating residues of β-(1,4) linked <a href="/wiki/N-Acetylglucosamine" title="N-Acetylglucosamine"><i>N</i>-acetylglucosamine</a> (NAG) and <a href="/wiki/N-Acetylmuramic_acid" title="N-Acetylmuramic acid"><i>N</i>-acetylmuramic acid</a> (NAM). Attached to the <i>N</i>-acetylmuramic acid is an <a href="/wiki/Oligopeptide" title="Oligopeptide">oligopeptide</a> chain made of three to five amino acids. The peptide chain can be cross-linked to the peptide chain of another strand forming the 3D mesh-like layer.<sup id="cite_ref-Madigan-2015_1-1" class="reference"><a href="#cite_note-Madigan-2015-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Peptidoglycan serves a structural role in the bacterial cell wall, giving structural strength, as well as counteracting the <a href="/wiki/Osmotic_pressure" title="Osmotic pressure">osmotic pressure</a> of the <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>. This repetitive linking results in a dense peptidoglycan layer which is critical for maintaining cell form and withstanding high osmotic pressures, and it is regularly replaced by peptidoglycan production. Peptidoglycan hydrolysis and synthesis are two processes that must occur in order for cells to grow and multiply, a technique carried out in three stages: clipping of current material, insertion of new material, and re-crosslinking of existing material to new material.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The peptidoglycan layer is substantially thicker in <a href="/wiki/Gram-positive_bacteria" title="Gram-positive bacteria">gram-positive bacteria</a> (20 to 80 nanometers) than in <a href="/wiki/Gram-negative_bacteria" title="Gram-negative bacteria">gram-negative bacteria</a> (7 to 8 nanometers).<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Depending on pH growth conditions, the peptidoglycan forms around 40 to 90% of the <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a>'s <a href="/wiki/Dry_matter" title="Dry matter">dry weight</a> of gram-positive bacteria but only around 10% of gram-negative strains. Thus, presence of high levels of peptidoglycan is the primary determinant of the characterisation of bacteria as gram-positive.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In gram-positive strains, it is important in attachment roles and <a href="/wiki/Serotyping" class="mw-redirect" title="Serotyping">serotyping</a> purposes.<sup id="cite_ref-Salton-1996_6-0" class="reference"><a href="#cite_note-Salton-1996-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> For both gram-positive and gram-negative bacteria, particles of approximately 2 nm can pass through the peptidoglycan.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>It is difficult to tell whether an organism is gram-positive or gram-negative using a microscope; <a href="/wiki/Gram_stain" title="Gram stain">Gram staining</a>, created by <a href="/wiki/Hans_Christian_Gram" title="Hans Christian Gram">Hans Christian Gram</a> in 1884, is required. The bacteria are stained with the dyes crystal violet and <a href="/wiki/Safranin" title="Safranin">safranin</a>. Gram positive cells are purple after staining, while Gram negative cells stain pink.<sup id="cite_ref-libre_8-0" class="reference"><a href="#cite_note-libre-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size skin-invert-image" typeof="mw:File/Thumb"><a href="/wiki/File:Peptidoglycan_en.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Peptidoglycan_en.svg/220px-Peptidoglycan_en.svg.png" decoding="async" width="220" height="156" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Peptidoglycan_en.svg/330px-Peptidoglycan_en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Peptidoglycan_en.svg/440px-Peptidoglycan_en.svg.png 2x" data-file-width="620" data-file-height="439" /></a><figcaption>Peptidoglycan.</figcaption></figure> <p>The peptidoglycan layer within the bacterial cell wall is a <a href="/wiki/Crystal_lattice" class="mw-redirect" title="Crystal lattice">crystal lattice</a> structure formed from linear chains of two alternating amino <a href="/wiki/Sugar" title="Sugar">sugars</a>, namely <a href="/wiki/N-Acetylglucosamine" title="N-Acetylglucosamine"><i>N</i>-acetylglucosamine</a> (GlcNAc or NAG) and <a href="/wiki/N-Acetylmuramic_acid" title="N-Acetylmuramic acid"><i>N</i>-acetylmuramic acid</a> (MurNAc or NAM). The alternating sugars are connected by a β-(1,4)-<a href="/wiki/Glycosidic_bond" title="Glycosidic bond">glycosidic bond</a>. Each MurNAc is attached to a short (4- to 5-residue) <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> chain, containing <small>L</small>-<a href="/wiki/Alanine" title="Alanine">alanine</a>, <small>D</small>-<a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamic acid</a>, <a href="/wiki/Meso-diaminopimelic_acid" class="mw-redirect" title="Meso-diaminopimelic acid"><i>meso</i>-diaminopimelic acid</a>, and <small>D</small>-alanine in the case of <i><a href="/wiki/Escherichia_coli" title="Escherichia coli">Escherichia coli</a></i> (a gram-negative bacterium); or <small>L</small>-alanine, <small>D</small>-<a href="/wiki/Glutamine" title="Glutamine">glutamine</a>, <small>L</small>-<a href="/wiki/Lysine" title="Lysine">lysine</a>, and <small>D</small>-alanine with a 5-<a href="/wiki/Glycine" title="Glycine">glycine</a> interbridge between tetrapeptides in the case of <i><a href="/wiki/Staphylococcus_aureus" title="Staphylococcus aureus">Staphylococcus aureus</a></i> (a gram-positive bacterium). Peptidoglycan is one of the most important sources of <a href="/wiki/D-Amino_acid" title="D-Amino acid"><small>D</small>-amino acids</a> in nature.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2023)">citation needed</span></a></i>]</sup> </p><p>By enclosing the inner membrane, the peptidoglycan layer protects the cell from <a href="/wiki/Lysis" title="Lysis">lysis</a> caused by the <a href="/wiki/Turgor_pressure" title="Turgor pressure">turgor</a> pressure of the cell. When the cell wall grows, it retains its shape throughout its life, so a rod shape will remain a rod shape, and a spherical shape will remain a spherical shape for life. This happens because the freshly added septal material of synthesis transforms into a hemispherical wall for the offspring cells.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Cross-link" title="Cross-link">Cross-linking</a> between <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> in different linear amino sugar chains occurs with the help of the enzyme <a href="/wiki/DD-Transpeptidase" title="DD-Transpeptidase"><small>DD</small>-transpeptidase</a> and results in a 3-dimensional structure that is strong and rigid. The specific amino acid sequence and molecular structure vary with the bacterial <a href="/wiki/Species" title="Species">species</a>.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>The different peptidoglycan types of bacterial cell walls and their taxonomic implications have been described.<sup id="cite_ref-Schleifer-1972_11-0" class="reference"><a href="#cite_note-Schleifer-1972-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Archaea" title="Archaea">Archaea</a> (<a href="/wiki/Domain_(biology)" title="Domain (biology)">domain</a> <i>Archaea</i>)<sup id="cite_ref-Woese-1990_12-0" class="reference"><a href="#cite_note-Woese-1990-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> do not contain peptidoglycan (murein).<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Some Archaea contain <a href="/wiki/Pseudopeptidoglycan" title="Pseudopeptidoglycan">pseudopeptidoglycan</a> (pseudomurein, see below).<sup id="cite_ref-Kandler-1998_14-0" class="reference"><a href="#cite_note-Kandler-1998-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="skin-invert-image"> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 375px"> <div class="thumb" style="width: 370px; height: 270px;"><span typeof="mw:File"><a href="/wiki/File:Mureine.svg" class="mw-file-description" title="The structure of peptidoglycan. NAG = N-acetylglucosamine (also called GlcNAc or NAGA), NAM = N-acetylmuramic acid (also called MurNAc or NAMA)."><img alt="The structure of peptidoglycan. NAG = N-acetylglucosamine (also called GlcNAc or NAGA), NAM = N-acetylmuramic acid (also called MurNAc or NAMA)." src="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Mureine.svg/271px-Mureine.svg.png" decoding="async" width="271" height="240" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Mureine.svg/406px-Mureine.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/29/Mureine.svg/541px-Mureine.svg.png 2x" data-file-width="432" data-file-height="383" /></a></span></div> <div class="gallerytext">The structure of peptidoglycan. NAG = <a href="/wiki/N-Acetylglucosamine" title="N-Acetylglucosamine"><i>N</i>-acetylglucosamine</a> (also called GlcNAc or NAGA), NAM = <a href="/wiki/N-Acetylmuramic_acid" title="N-Acetylmuramic acid"><i>N</i>-acetylmuramic acid</a> (also called MurNAc or NAMA).</div> </li> <li class="gallerybox" style="width: 375px"> <div class="thumb" style="width: 370px; height: 270px;"><span typeof="mw:File"><a href="/wiki/File:Gram-positive_cellwall-schematic.png" class="mw-file-description" title="Gram-positive cell wall"><img alt="Gram-positive cell wall" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Gram-positive_cellwall-schematic.png/302px-Gram-positive_cellwall-schematic.png" decoding="async" width="302" height="240" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Gram-positive_cellwall-schematic.png/453px-Gram-positive_cellwall-schematic.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Gram-positive_cellwall-schematic.png/605px-Gram-positive_cellwall-schematic.png 2x" data-file-width="1482" data-file-height="1177" /></a></span></div> <div class="gallerytext"><a href="/wiki/Gram-positive" class="mw-redirect" title="Gram-positive">Gram-positive</a> <a href="/wiki/Cell_wall" title="Cell wall">cell wall</a></div> </li> <li class="gallerybox" style="width: 375px"> <div class="thumb" style="width: 370px; height: 270px;"><span typeof="mw:File"><a href="/wiki/File:PBP_catalysis.svg" class="mw-file-description" title="Penicillin binding protein forming cross-links in newly formed bacterial cell wall."><img alt="Penicillin binding protein forming cross-links in newly formed bacterial cell wall." src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/PBP_catalysis.svg/175px-PBP_catalysis.svg.png" decoding="async" width="175" height="240" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/PBP_catalysis.svg/262px-PBP_catalysis.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8d/PBP_catalysis.svg/350px-PBP_catalysis.svg.png 2x" data-file-width="1142" data-file-height="1567" /></a></span></div> <div class="gallerytext"><a href="/wiki/Penicillin_binding_protein" class="mw-redirect" title="Penicillin binding protein">Penicillin binding protein</a> forming cross-links in newly formed bacterial cell wall.</div> </li> </ul> </div> <p>Peptidoglycan is involved in <a href="/wiki/Binary_fission" class="mw-redirect" title="Binary fission">binary fission</a> during bacterial cell reproduction. <a href="/wiki/L-form_bacteria" title="L-form bacteria">L-form bacteria</a> and <a href="/wiki/Mycoplasma" title="Mycoplasma">mycoplasmas</a>, both lacking peptidoglycan cell walls, do not proliferate by binary fission, but by a <a href="/wiki/Budding" title="Budding">budding</a> mechanism.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>In the course of early evolution, the successive development of boundaries (membranes, walls) protecting first structures of life against their environment must have been essential for the formation of the first cells (<a href="/wiki/Cellularization" title="Cellularization">cellularisation</a>). </p><p>The invention of rigid peptidoglycan (murein) cell walls in bacteria (domain <i>Bacteria</i><sup id="cite_ref-Woese-1990_12-1" class="reference"><a href="#cite_note-Woese-1990-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>) was probably the prerequisite for their survival, extensive radiation and colonisation of virtually all habitats of the geosphere and hydrosphere.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Biosynthesis">Biosynthesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=2" title="Edit section: Biosynthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The peptidoglycan monomers are synthesized in the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> and are then attached to a membrane carrier <a href="/wiki/Bactoprenol" title="Bactoprenol">bactoprenol</a>. Bactoprenol transports peptidoglycan monomers across the cell membrane where they are inserted into the existing peptidoglycan.<sup id="cite_ref-The_Prokaryotic_Cell_19-0" class="reference"><a href="#cite_note-The_Prokaryotic_Cell-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <ol><li>In the first step of peptidoglycan synthesis, <a href="/wiki/Glutamine" title="Glutamine">glutamine</a>, which is an amino acid, donates an amino group to a sugar, <a href="/wiki/Fructose_6-phosphate" title="Fructose 6-phosphate">fructose 6-phosphate</a>.<sup id="cite_ref-White-2007_20-0" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This reaction, catalyzed by <a href="/wiki/EC_2.6.1.16" class="mw-redirect" title="EC 2.6.1.16">EC 2.6.1.16</a> (GlmS), turns fructose 6-phosphate into <a href="/w/index.php?title=Glucosamine-6-phosphate&action=edit&redlink=1" class="new" title="Glucosamine-6-phosphate (page does not exist)">glucosamine-6-phosphate</a>.<sup id="cite_ref-Otten-2018_21-0" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>In step two, an acetyl group is transferred from <a href="/wiki/Acetyl_CoA" class="mw-redirect" title="Acetyl CoA">acetyl CoA</a> to the amino group on the glucosamine-6-phosphate creating <a href="/w/index.php?title=N-acetyl-glucosamine-6-phosphate&action=edit&redlink=1" class="new" title="N-acetyl-glucosamine-6-phosphate (page does not exist)"><i>N</i>-acetyl-glucosamine-6-phosphate</a>.<sup id="cite_ref-White-2007_20-1" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This reaction is <a href="/wiki/EC_5.4.2.10" class="mw-redirect" title="EC 5.4.2.10">EC 5.4.2.10</a>, catalyzed by GlmM.<sup id="cite_ref-Otten-2018_21-1" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li><span class="anchor" id="GlmU"></span>In step three of the synthesis process, the <i>N</i>-acetyl-glucosamine-6-phosphate is isomerized, which will change <i>N</i>-acetyl-glucosamine-6-phosphate to <a href="/w/index.php?title=N-acetyl-glucosamine-1-phosphate&action=edit&redlink=1" class="new" title="N-acetyl-glucosamine-1-phosphate (page does not exist)"><i>N</i>-acetyl-glucosamine-1-phosphate</a>.<sup id="cite_ref-White-2007_20-2" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This is <a href="/wiki/EC_2.3.1.157" class="mw-redirect" title="EC 2.3.1.157">EC 2.3.1.157</a>, catalyzed by GlmU.<sup id="cite_ref-Otten-2018_21-2" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>In step 4, the <i>N</i>-acetyl-glucosamine-1-phosphate, which is now a monophosphate, attacks <a href="/wiki/Uridine_triphosphate" title="Uridine triphosphate">UTP</a>. Uridine triphosphate, which is a <a href="/wiki/Pyrimidine" title="Pyrimidine">pyrimidine</a> <a href="/wiki/Nucleotide" title="Nucleotide">nucleotide</a>, has the ability to act as an energy source. In this particular reaction, after the monophosphate has attacked the UTP, an inorganic pyrophosphate is given off and is replaced by the monophosphate, creating UDP-N-acetylglucosamine (2,4). (When <a href="/wiki/Uridine_diphosphate" title="Uridine diphosphate">UDP</a> is used as an energy source, it gives off an inorganic phosphate.) This initial stage, is used to create the precursor for the NAG in peptidoglycan.<sup id="cite_ref-White-2007_20-3" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This is <a href="/wiki/EC_2.7.7.23" class="mw-redirect" title="EC 2.7.7.23">EC 2.7.7.23</a>, also catalyzed by GlmU, which is a bifunctional enzyme.<sup id="cite_ref-Otten-2018_21-3" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>In step 5, some of the UDP-N-acetylglucosamine (UDP-GlcNAc) is converted to UDP-MurNAc (UDP-N-acetylmuramic acid) by the addition of a lactyl group to the glucosamine. Also in this reaction, the C3 hydroxyl group will remove a phosphate from the alpha carbon of <a href="/wiki/Phosphoenolpyruvic_acid" title="Phosphoenolpyruvic acid">phosphoenolpyruvate</a>. This creates what is called an enol derivative.<sup id="cite_ref-White-2007_20-4" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> <a href="/wiki/EC_2.5.1.7" class="mw-redirect" title="EC 2.5.1.7">EC 2.5.1.7</a>, catalyzed by MurA.<sup id="cite_ref-Otten-2018_21-4" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>In step 6, the enol is reduced to a "lactyl moiety" by NADPH in step six.<sup id="cite_ref-White-2007_20-5" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> <a href="/wiki/EC_1.3.1.98" class="mw-redirect" title="EC 1.3.1.98">EC 1.3.1.98</a>, catalyzed by MurB.<sup id="cite_ref-Otten-2018_21-5" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>In step 7, the UDP–MurNAc is converted to UDP-MurNAc pentapeptide by the addition of five amino acids, usually including the dipeptide <small>D</small>-alanyl-<small>D</small>-alanine.<sup id="cite_ref-White-2007_20-6" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This is a string of three reactions: <a href="/wiki/EC_6.3.2.8" class="mw-redirect" title="EC 6.3.2.8">EC 6.3.2.8</a> by MurC, <a href="/wiki/EC_6.3.2.9" class="mw-redirect" title="EC 6.3.2.9">EC 6.3.2.9</a> by MurD, and <a href="/wiki/EC_6.3.2.13" class="mw-redirect" title="EC 6.3.2.13">EC 6.3.2.13</a> by MurE.<sup id="cite_ref-Otten-2018_21-6" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li></ol> <p>Each of these reactions requires the energy source ATP.<sup id="cite_ref-White-2007_20-7" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This is all referred to as Stage one. </p><p>Stage two occurs in the cytoplasmic membrane. It is in the membrane where a lipid carrier called <a href="/wiki/Bactoprenol" title="Bactoprenol">bactoprenol</a> carries peptidoglycan precursors through the cell membrane. </p> <ol><li><a href="/wiki/Undecaprenyl_phosphate" title="Undecaprenyl phosphate">Undecaprenyl phosphate</a> will attack the UDP-MurNAc penta, creating a PP-MurNac penta, which is now a lipid (<a href="/w/index.php?title=Lipid_I&action=edit&redlink=1" class="new" title="Lipid I (page does not exist)">lipid I</a>).<sup id="cite_ref-White-2007_20-8" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> <a href="/wiki/EC_2.7.8.13" class="mw-redirect" title="EC 2.7.8.13">EC 2.7.8.13</a> by MraY.<sup id="cite_ref-Otten-2018_21-7" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>UDP-GlcNAc is then transported to MurNAc, creating Lipid-PP-MurNAc penta-GlcNAc (<a href="/wiki/Lipid_II" title="Lipid II">lipid II</a>), a disaccharide, also a precursor to peptidoglycan.<sup id="cite_ref-White-2007_20-9" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> <a href="/wiki/EC_2.4.1.227" class="mw-redirect" title="EC 2.4.1.227">EC 2.4.1.227</a> by MurG.<sup id="cite_ref-Otten-2018_21-8" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li> <li>Lipid II is transported across the membrane by <a href="/wiki/Flippase" title="Flippase">flippase</a> (MurJ), a discovery made in 2014 after decades of searching.<sup id="cite_ref-Sham-2014_22-0" class="reference"><a href="#cite_note-Sham-2014-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Once it is there, it is added to the growing glycan chain by the enzyme <a href="/wiki/Peptidoglycan_glycosyltransferase" title="Peptidoglycan glycosyltransferase">peptidoglycan glycosyltransferase</a> (GTase, EC 2.4.1.129). This reaction is known as transglycosylation. In the reaction, the hydroxyl group of the GlcNAc will attach to the MurNAc in the glycan, which will displace the lipid-PP from the glycan chain.<sup id="cite_ref-White-2007_20-10" class="reference"><a href="#cite_note-White-2007-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li> <li>In a final step, the <a href="/wiki/DD-Transpeptidase" title="DD-Transpeptidase"><small>DD</small>-transpeptidase</a> (TPase, EC 3.4.16.4) crosslinks individual glycan chains. This protein is also known as the <a href="/wiki/Penicillin-binding_protein" class="mw-redirect" title="Penicillin-binding protein">penicillin-binding protein</a>. Some versions of the enzyme also performs the glycosyltransferase function, while others leave the job to a separate enzyme.<sup id="cite_ref-Otten-2018_21-9" class="reference"><a href="#cite_note-Otten-2018-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li></ol> <div style="clear:left;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Pseudopeptidoglycan">Pseudopeptidoglycan</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=3" title="Edit section: Pseudopeptidoglycan"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Pseudopeptidoglycan" title="Pseudopeptidoglycan">Pseudopeptidoglycan</a></div> <p>In some <a href="/wiki/Archaea" title="Archaea">archaea</a>, i.e. members of the <a href="/wiki/Methanobacteriales" title="Methanobacteriales">Methanobacteriales</a> and in the genus <i><a href="/wiki/Methanopyrus" title="Methanopyrus">Methanopyrus</a></i>, <a href="/wiki/Pseudopeptidoglycan" title="Pseudopeptidoglycan">pseudopeptidoglycan</a> (pseudomurein) has been found.<sup id="cite_ref-Kandler-1998_14-1" class="reference"><a href="#cite_note-Kandler-1998-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> In pseudopeptidoglycan the sugar residues are β-(1,3) linked <i>N</i>-acetylglucosamine and <a href="/wiki/N-Acetyltalosaminuronic_acid" title="N-Acetyltalosaminuronic acid"><i>N</i>-acetyltalosaminuronic acid</a>. This makes the cell walls of such archaea insensitive to <a href="/wiki/Lysozyme" title="Lysozyme">lysozyme</a>.<sup id="cite_ref-Madigan-2009_23-0" class="reference"><a href="#cite_note-Madigan-2009-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> The biosynthesis of pseudopeptidoglycan has been described.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Recognition_by_immune_system">Recognition by immune system</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=4" title="Edit section: Recognition by immune system"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Peptidoglycan recognition is an evolutionarily conserved process.<sup id="cite_ref-Wolf-2018_25-0" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> The overall structure is similar between bacterial species, but various modifications can increase the diversity. These include modifications of the length of sugar polymers, modifications in the sugar structures, variations in cross-linking or substitutions of amino acids (primarily at the third position).<sup id="cite_ref-Wolf-2018_25-1" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bersch-2021_26-0" class="reference"><a href="#cite_note-Bersch-2021-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The aim of these modifications is to alter the properties of the cell wall, which plays a vital role in <a href="/wiki/Pathogenesis" title="Pathogenesis">pathogenesis</a>.<sup id="cite_ref-Wolf-2018_25-2" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Peptidoglycans can be degraded by several enzymes (<a href="/wiki/Lysozyme" title="Lysozyme">lysozyme</a>, glucosaminidase, <a href="/wiki/Endopeptidase" title="Endopeptidase">endopeptidase</a>...<sup id="cite_ref-Wolf-2018_25-3" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup>), producing immunostimulatory fragments (sometimes called muropeptides<sup id="cite_ref-Bastos-2021_27-0" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup>) that are critical for mediating <a href="/wiki/Host%E2%80%93pathogen_interaction" title="Host–pathogen interaction">host-pathogen interactions</a>.<sup id="cite_ref-Bersch-2021_26-1" class="reference"><a href="#cite_note-Bersch-2021-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> These include MDP (<a href="/wiki/Muramyl_dipeptide" title="Muramyl dipeptide">muramyl dipeptide</a>), NAG (<a href="/wiki/N-Acetylglucosamine" title="N-Acetylglucosamine">N-acetylglucosamine</a>) or iE-DAP (γ-d-glutamyl-meso-diaminopimelic acid).<sup id="cite_ref-Wolf-2018_25-4" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bastos-2021_27-1" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>Peptidoglycan from <a href="/wiki/Gut_microbiota" title="Gut microbiota">intestinal bacteria</a> (both pathogens and commensals) crosses the intestinal barrier even under physiological conditions.<sup id="cite_ref-Bastos-2021_27-2" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Mechanisms through which peptidoglycan or its fragments enter the host cells can be direct (carrier-independent) or indirect (carrier-dependent), and they are either bacteria-mediated (secretion systems, <a href="/wiki/Membrane_vesicle_trafficking" title="Membrane vesicle trafficking">membrane vesicles</a>) or host cell-mediated (receptor-mediated, peptide transporters).<sup id="cite_ref-Bastos-2021_27-3" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Bacterial_secretion_system" title="Bacterial secretion system">Bacterial secretion systems</a> are protein complexes used for the delivery of virulence factors across the bacterial cell envelope to the exterior environment.<sup id="cite_ref-Sun-2022_28-0" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Intracellular bacterial pathogens invade eukaryotic cells (which may lead to the formation of <a href="/wiki/Phagolysosome" title="Phagolysosome">phagolysosomes</a> and/or <a href="/wiki/Autophagy" title="Autophagy">autophagy</a> activation), or bacteria may be engulfed by <a href="/wiki/Phagocyte" title="Phagocyte">phagocytes</a> (<a href="/wiki/Macrophage" title="Macrophage">macrophages</a>, <a href="/wiki/Monocyte" title="Monocyte">monocytes</a>, <a href="/wiki/Neutrophil" title="Neutrophil">neutrophils</a>...). The bacteria-containing <a href="/wiki/Phagosome" title="Phagosome">phagosome</a> may then fuse with <a href="/wiki/Endosome" title="Endosome">endosomes</a> and <a href="/wiki/Lysosome" title="Lysosome">lysosomes</a>, leading to degradation of bacteria and generation of polymeric peptidoglycan fragments and muropeptides.<sup id="cite_ref-Bastos-2021_27-4" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Receptors">Receptors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=5" title="Edit section: Receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Innate_immune_system" title="Innate immune system">Innate immune system</a> senses intact peptidoglycan and peptidoglycan fragments using numerous PRRs (<a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern recognition receptors</a>) that are secreted, expressed intracellularly or expressed on the cell surface.<sup id="cite_ref-Wolf-2018_25-5" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Peptidoglycan_recognition_proteins">Peptidoglycan recognition proteins</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=6" title="Edit section: Peptidoglycan recognition proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Peptidoglycan_recognition_protein" title="Peptidoglycan recognition protein">PGLYRPs</a> are conserved from <a href="/wiki/Insect" title="Insect">insects</a> to <a href="/wiki/Mammal" title="Mammal">mammals</a>.<sup id="cite_ref-Bastos-2021_27-5" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Mammals produce four secreted soluble peptidoglycan recognition proteins (<a href="/wiki/Peptidoglycan_recognition_protein_1" title="Peptidoglycan recognition protein 1">PGLYRP-1</a>, <a href="/wiki/Peptidoglycan_recognition_protein_2" title="Peptidoglycan recognition protein 2">PGLYRP-2</a>, <a href="/wiki/Peptidoglycan_recognition_protein_3" title="Peptidoglycan recognition protein 3">PGLYRP-3</a> and <a href="/wiki/Peptidoglycan_recognition_protein_4" title="Peptidoglycan recognition protein 4">PGLYRP-4</a>) that recognize muramyl pentapeptide or tetrapeptide.<sup id="cite_ref-Wolf-2018_25-6" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> They can also bind to <a href="/wiki/Lipopolysaccharide" title="Lipopolysaccharide">LPS</a> and other molecules by using binding sites outside of the peptidoglycan-binding groove.<sup id="cite_ref-Sun-2022_28-1" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> After recognition of peptidoglycan, PGLYRPs activate <a href="/wiki/Polyphenol_oxidase" title="Polyphenol oxidase">polyphenol oxidase</a> (PPO) molecules, Toll, or immune deficiency (IMD) signalling pathways. That leads to production of <a href="/wiki/Antimicrobial_peptides" title="Antimicrobial peptides">antimicrobial peptides</a> (AMPs).<sup id="cite_ref-Sun-2022_28-2" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Each of the mammalian PGLYRPs display unique tissue expression patterns. PGLYRP-1 is mainly expressed in the granules of <a href="/wiki/Neutrophil" title="Neutrophil">neutrophils</a> and <a href="/wiki/Eosinophil" title="Eosinophil">eosinophils</a>.<sup id="cite_ref-Wolf-2018_25-7" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> PGLYRP-3 and 4 are expressed by several tissues such as skin, sweat glands, eyes or the intestinal tract.<sup id="cite_ref-Bastos-2021_27-6" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> PGLYRP-1, 3 and 4 form disulphide-linked <a href="/wiki/Homodimers" class="mw-redirect" title="Homodimers">homodimers</a> and <a href="/wiki/Heterodimers" class="mw-redirect" title="Heterodimers">heterodimers</a> essential for their bactericidal activity.<sup id="cite_ref-Bastos-2021_27-7" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Their binding to bacterial cell wall peptidoglycans can induce bacterial cell death by interaction with various bacterial transcriptional regulatory proteins.<sup id="cite_ref-Wolf-2018_25-8" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> PGLYRPs are likely to assist in bacterial killing by cooperating with other PRRs to enhance recognition of bacteria by phagocytes.<sup id="cite_ref-Wolf-2018_25-9" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>PGLYRP-2 is primarily expressed by the <a href="/wiki/Liver" title="Liver">liver</a> and secreted into the circulation.<sup id="cite_ref-Wolf-2018_25-10" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Also, its expression can be induced in skin <a href="/wiki/Keratinocyte" title="Keratinocyte">keratinocytes</a>, oral and intestinal <a href="/wiki/Epithelium" title="Epithelium">epithelial</a> cells.<sup id="cite_ref-Bastos-2021_27-8" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> In contrast with the other PGLYRPs, PGLYRP-2 has no direct bactericidal activity. It possesses peptidoglycan amidase activity, it hydrolyses the lactyl-amide bond between the <a href="/wiki/MurNAc" class="mw-redirect" title="MurNAc">MurNAc</a> and the first amino acid of the stem peptide of peptidoglycan.<sup id="cite_ref-Wolf-2018_25-11" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bastos-2021_27-9" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> It is proposed, that the function of PGLYRP-2 is to prevent over-activation of the immune system and <a href="/wiki/Inflammation" title="Inflammation">inflammation</a>-induced tissue damage in response to <a href="/wiki/NOD2" title="NOD2">NOD2</a> ligands (see below), as these muropeptides can no longer be recognized by NOD2 upon separation of the peptide component from MurNAc.<sup id="cite_ref-Bastos-2021_27-10" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Growing evidence suggests that peptidoglycan recognition protein family members play a dominant role in the <a href="/wiki/Tolerance_to_infections" title="Tolerance to infections">tolerance</a> of intestinal epithelial cells toward the commensal microbiota.<sup id="cite_ref-Sun-2022_28-3" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> It has been demonstrated that expression of PGLYRP-2 and 4 can influence the composition of the intestinal <a href="/wiki/Microbiota" title="Microbiota">microbiota</a>.<sup id="cite_ref-Sun-2022_28-4" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Recently, it has been discovered, that PGLYRPs (and also NOD-like receptors and peptidoglycan transporters) are highly expressed in the developing mouse <a href="/wiki/Brain" title="Brain">brain</a>.<sup id="cite_ref-Gonzalez-Santana-2020_30-0" class="reference"><a href="#cite_note-Gonzalez-Santana-2020-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> PGLYRP-2 and is highly expressed in <a href="/wiki/Neuron" title="Neuron">neurons</a> of several brain regions including the <a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">prefrontal cortex</a>, <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, and <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>, thus indicating potential direct effects of peptidoglycan on neurons. PGLYRP-2 is highly expressed also in the cerebral cortex of young children, but not in most adult cortical tissues. PGLYRP-1 is also expressed in the brain and continues to be expressed into adulthood.<sup id="cite_ref-Gonzalez-Santana-2020_30-1" class="reference"><a href="#cite_note-Gonzalez-Santana-2020-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="NOD-like_receptors">NOD-like receptors</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=7" title="Edit section: NOD-like receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Probably the most well-known receptors of peptidoglycan are the <a href="/wiki/NOD-like_receptor" title="NOD-like receptor">NOD-like receptors</a> (NLRs), mainly <a href="/wiki/NOD1" title="NOD1">NOD1</a> and <a href="/wiki/NOD2" title="NOD2">NOD2</a>. The NOD1 receptor is activated after iE-DAP (γ-d-glutamyl-meso-diaminopimelic acid) binding, while NOD2 recognizes MDP (muramyl dipeptide), by their <a href="/wiki/LRR_domain" class="mw-redirect" title="LRR domain">LRR domains</a>.<sup id="cite_ref-Sun-2022_28-5" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Activation leads to self-oligomerization, resulting in activation of two signalling cascades. One triggers activation of <a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-κB</a> (through RIP2, <a href="/wiki/MAP3K7" title="MAP3K7">TAK1</a> and <a href="/wiki/I%CE%BAB_kinase" title="IκB kinase">IKK</a><sup id="cite_ref-Murphy-2017_31-0" class="reference"><a href="#cite_note-Murphy-2017-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup>), second leads to <a href="/wiki/Mitogen-activated_protein_kinase" title="Mitogen-activated protein kinase">MAPK</a> signalling cascade. Activation of these pathways induces production of inflammatory <a href="/wiki/Cytokine" title="Cytokine">cytokines</a> and <a href="/wiki/Chemokine" title="Chemokine">chemokines</a>.<sup id="cite_ref-Wolf-2018_25-12" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>NOD1 is expressed by diverse cell types, including myeloid phagocytes, epithelial cells<sup id="cite_ref-Wolf-2018_25-13" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> and neurons.<sup id="cite_ref-Gonzalez-Santana-2020_30-2" class="reference"><a href="#cite_note-Gonzalez-Santana-2020-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> NOD2 is expressed in monocytes and macrophages, epithelial intestinal cells, <a href="/wiki/Paneth_cell" title="Paneth cell">Paneth cells</a>, <a href="/wiki/Dendritic_cell" title="Dendritic cell">dendritic cells</a>, <a href="/wiki/Osteoblast" title="Osteoblast">osteoblasts</a>, keratinocytes and other epithelial cell types.<sup id="cite_ref-Bastos-2021_27-11" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> As <a href="/wiki/Cytosol" title="Cytosol">cytosolic</a> sensors, NOD1 and NOD2 must either detect bacteria that enter the cytosol, or peptidoglycan must be degraded to generate fragments that must be transported into the cytosol for these sensors to function.<sup id="cite_ref-Wolf-2018_25-14" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Recently, it was demonstrated that <a href="/wiki/NLRP3" title="NLRP3">NLRP3</a> is activated by peptidoglycan, through a mechanism that is independent of NOD1 and NOD2.<sup id="cite_ref-Bastos-2021_27-12" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> In macrophages, N-acetylglucosamine generated by peptidoglycan degradation was found to inhibit hexokinase activity and induce its release from the <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondrial</a> <a href="/wiki/Membrane" title="Membrane">membrane</a>. It promotes NLRP3 <a href="/wiki/Inflammasome" title="Inflammasome">inflammasome</a> activation through a mechanism triggered by increased mitochondrial membrane permeability.<sup id="cite_ref-Bastos-2021_27-13" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/NLRP1" title="NLRP1">NLRP1</a> is also considered as a cytoplasmic sensor of peptidoglycan. It can sense MDP and promote <a href="/wiki/Interleukin-1_family" title="Interleukin-1 family">IL-1</a> secretion through binding NOD2.<sup id="cite_ref-Sun-2022_28-6" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bersch-2021_26-2" class="reference"><a href="#cite_note-Bersch-2021-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="C-type_lectin_receptors_(CLRs)"><span id="C-type_lectin_receptors_.28CLRs.29"></span>C-type lectin receptors (CLRs)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=8" title="Edit section: C-type lectin receptors (CLRs)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/C-type_lectin" title="C-type lectin">C-type lectins</a> are a diverse superfamily of mainly Ca<sup>2+</sup>-dependent proteins that bind a variety of <a href="/wiki/Carbohydrate" title="Carbohydrate">carbohydrates</a> (including the glycan skeleton of peptidoglycan), and function as innate immune receptors.<sup id="cite_ref-Bastos-2021_27-14" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> CLR proteins that bind to peptidoglycan include MBL (<a href="/wiki/Mannose-Binding_Lectin" class="mw-redirect" title="Mannose-Binding Lectin">mannose binding lectin</a>), <a href="/wiki/Ficolin" title="Ficolin">ficolins</a>, <a href="/wiki/REG3A" title="REG3A">Reg3A</a> (regeneration gene family protein 3A) and PTCLec1.<sup id="cite_ref-Sun-2022_28-7" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> In mammals, they initiate the <a href="/wiki/Lectin_pathway" title="Lectin pathway">lectin-pathway</a> of the <a href="/wiki/Complement_system" title="Complement system">complement</a> cascade.<sup id="cite_ref-Bastos-2021_27-15" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Toll-like_receptors">Toll-like receptors</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=9" title="Edit section: Toll-like receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The role of <a href="/wiki/Toll-like_receptor" title="Toll-like receptor">TLRs</a> in direct recognition of peptidoglycan is controversial.<sup id="cite_ref-Wolf-2018_25-15" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> In some studies, has been reported that peptidoglycan is sensed by <a href="/wiki/Toll-like_receptor_2" title="Toll-like receptor 2">TLR2</a>.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> But this TLR2-inducing activity could be due to cell wall <a href="/wiki/Lipoprotein" title="Lipoprotein">lipoproteins</a> and <a href="/wiki/Lipoteichoic_acid" title="Lipoteichoic acid">lipoteichoic acids</a> that commonly co-purify with peptidoglycan. Also variation in peptidoglycan structure in bacteria from species to species may contribute to the differing results on this topic.<sup id="cite_ref-Wolf-2018_25-16" class="reference"><a href="#cite_note-Wolf-2018-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bastos-2021_27-16" class="reference"><a href="#cite_note-Bastos-2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="As_vaccine_or_adjuvant">As vaccine or adjuvant</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=10" title="Edit section: As vaccine or adjuvant"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Peptidoglycan is immunologically active, which can stimulate immune cells to increase the expression of cytokines and enhance antibody-dependent specific response when combined with <a href="/wiki/Vaccine" title="Vaccine">vaccine</a> or as <a href="/wiki/Adjuvants,_immunologic" class="mw-redirect" title="Adjuvants, immunologic">adjuvant</a> alone.<sup id="cite_ref-Sun-2022_28-8" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> MDP, which is the basic unit of peptidoglycan, was initially used as the active component of <a href="/wiki/Freund%27s_adjuvant" title="Freund's adjuvant">Freund's adjuvant</a>.<sup id="cite_ref-Sun-2022_28-9" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Peptidoglycan from <i><a href="/wiki/Staphylococcus_aureus" title="Staphylococcus aureus">Staphylococcus aureus</a></i> was used as a vaccine to protect mice, showing that after vaccine injection for 40 weeks, the mice survived from <i>S. aureus</i> challenge at an increased <a href="/wiki/Lethal_dose" title="Lethal dose">lethal dose</a>.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Inhibition_and_degradation">Inhibition and degradation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=11" title="Edit section: Inhibition and degradation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some <a href="/wiki/Antibiotics" class="mw-redirect" title="Antibiotics">antibacterial drugs</a> such as <a href="/wiki/Penicillin" title="Penicillin">penicillin</a> interfere with the production of peptidoglycan by binding to bacterial enzymes known as <a href="/wiki/Penicillin-binding_proteins" title="Penicillin-binding proteins">penicillin-binding proteins</a> or <a href="/wiki/DD-Transpeptidase" title="DD-Transpeptidase"><small>DD</small>-transpeptidases</a>.<sup id="cite_ref-Salton-1996_6-1" class="reference"><a href="#cite_note-Salton-1996-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Penicillin-binding proteins form the bonds between oligopeptide crosslinks in peptidoglycan. For a bacterial cell to reproduce through <a href="/wiki/Binary_fission" class="mw-redirect" title="Binary fission">binary fission</a>, more than a million peptidoglycan subunits (NAM-NAG+oligopeptide) must be attached to existing subunits.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Mutations in genes coding for transpeptidases that lead to reduced interactions with an antibiotic are a significant source of emerging <a href="/wiki/Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">antibiotic resistance</a>.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Since peptidoglycan is also lacking in L-form bacteria and in mycoplasmas, both are resistant against penicillin. </p><p>Other steps of peptidoglycan synthesis can also be targeted. The topical antibiotic <a href="/wiki/Bacitracin" title="Bacitracin">bacitracin</a> targets the utilization of <a href="/wiki/C55-isoprenyl_pyrophosphate" title="C55-isoprenyl pyrophosphate">C55-isoprenyl pyrophosphate</a>. <a href="/wiki/Lantibiotics" title="Lantibiotics">Lantibiotics</a>, which includes the food preservative <a href="/wiki/Nisin" title="Nisin">nisin</a>, attack lipid II.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Lysozyme" title="Lysozyme">Lysozyme</a>, which is found in tears and constitutes part of the body's <a href="/wiki/Innate_immune_system" title="Innate immune system">innate immune system</a> exerts its antibacterial effect by breaking the β-(1,4)-glycosidic bonds in peptidoglycan (see above). Lysozyme is more effective in acting against <a href="/wiki/Gram-positive_bacteria" title="Gram-positive bacteria">gram-positive bacteria</a>, in which the peptidoglycan cell wall is exposed, than against <a href="/wiki/Gram-negative_bacteria" title="Gram-negative bacteria">gram-negative bacteria</a>, which have an outer layer of <a href="/wiki/Lipopolysaccharide" title="Lipopolysaccharide">LPS</a> covering the peptidoglycan layer.<sup id="cite_ref-Murphy-2017_31-1" class="reference"><a href="#cite_note-Murphy-2017-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Several bacterial peptidoglycan modifications can result in resistance to degradation by lysozyme. Susceptibility of bacteria to degradation is also considerably affected by exposure to <a href="/wiki/Antibiotic" title="Antibiotic">antibiotics</a>. Exposed bacteria synthesize peptidoglycan that contains shorter sugar chains that are poorly crosslinked and this peptidoglycan is then more easily degraded by lysozyme.<sup id="cite_ref-Sun-2022_28-10" class="reference"><a href="#cite_note-Sun-2022-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=12" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Undecaprenyl-diphosphatase" title="Undecaprenyl-diphosphatase">Undecaprenyl-diphosphatase</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=13" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Madigan-2015-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Madigan-2015_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Madigan-2015_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFMadiganMartinkoBenderBuckley2015" class="citation book cs1">Madigan, Michael T.; Martinko, John M.; Bender, Kelly S.; Buckley, Daniel H.; Stahl, David A. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=MedChemComm&rft.atitle=A+review+on+cell+wall+synthesis+inhibitors+with+an+emphasis+on+glycopeptide+antibiotics&rft.volume=8&rft.issue=3&rft.pages=516-533&rft.date=2017-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6072328%23id-name%3DPMC&rft_id=info%3Apmid%2F30108769&rft_id=info%3Adoi%2F10.1039%2Fc6md00585c&rft.aulast=Sarkar&rft.aufirst=P&rft.au=Yarlagadda%2C+V&rft.au=Ghosh%2C+C&rft.au=Haldar%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6072328&rfr_id=info%3Asid%2Fen.wikipedia.org%3APeptidoglycan" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Peptidoglycan&action=edit&section=14" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mmed.figgrp.298">Diagrammatic representation of peptidoglycan structures.</a></li> <li><a rel="nofollow" class="external text" href="http://pubs.acs.org/doi/abs/10.1021/bi4010446">Structure of MurNAc 6-Phosphate Hydrolase (MurQ) from Haemophilus influenzae with a Bound Inhibitor.</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol 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.navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Protein,_glycoconjugate:_glycoproteins_and_glycopeptides" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:#e7dcc3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar 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template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Glycoproteins" title="Special:EditPage/Template:Glycoproteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Protein,_glycoconjugate:_glycoproteins_and_glycopeptides" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein" title="Protein">Protein</a>, <a href="/wiki/Glycoconjugate" title="Glycoconjugate">glycoconjugate</a>: <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoproteins</a> and <a href="/wiki/Glycopeptide" title="Glycopeptide">glycopeptides</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Mucoprotein" title="Mucoprotein">Mucoproteins</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Mucin" title="Mucin">Mucin</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CD43" title="CD43">CD43</a></li> <li><a href="/wiki/CD164" title="CD164">CD164</a></li> <li><a href="/wiki/MUC1" class="mw-redirect" title="MUC1">MUC1</a></li> <li><a href="/wiki/MUC2" class="mw-redirect" title="MUC2">MUC2</a></li> <li><a href="/wiki/MUC3A" class="mw-redirect" title="MUC3A">MUC3A</a></li> <li><a href="/wiki/MUC3B" class="mw-redirect" title="MUC3B">MUC3B</a></li> <li><a href="/wiki/MUC4" class="mw-redirect" title="MUC4">MUC4</a></li> <li><a href="/wiki/MUC5AC" class="mw-redirect" title="MUC5AC">MUC5AC</a></li> <li><a href="/wiki/MUC5B" class="mw-redirect" title="MUC5B">MUC5B</a></li> <li><a href="/wiki/MUC6" class="mw-redirect" title="MUC6">MUC6</a></li> <li><a href="/wiki/MUC7" class="mw-redirect" title="MUC7">MUC7</a></li> <li><a href="/wiki/MUC8" class="mw-redirect" title="MUC8">MUC8</a></li> <li><a href="/wiki/Mucin" title="Mucin">MUC12</a></li> <li><a href="/wiki/MUC13" class="mw-redirect" title="MUC13">MUC13</a></li> <li><a href="/wiki/MUC15" class="mw-redirect" title="MUC15">MUC15</a></li> <li><a href="/wiki/CA-125" class="mw-redirect" title="CA-125">MUC16</a></li> <li><a href="/wiki/MUC17" class="mw-redirect" title="MUC17">MUC17</a></li> <li><a href="/wiki/MUC19" class="mw-redirect" title="MUC19">MUC19</a></li> <li><a href="/wiki/MUC20" class="mw-redirect" title="MUC20">MUC20</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Haptoglobin" title="Haptoglobin">Haptoglobin</a></li> <li><a href="/wiki/Intrinsic_factor" title="Intrinsic factor">Intrinsic factor</a></li> <li><a href="/wiki/Orosomucoid" title="Orosomucoid">Orosomucoid</a></li> <li><a class="mw-selflink selflink">Peptidoglycan</a></li> <li><a href="/wiki/Phytohaemagglutinin" title="Phytohaemagglutinin">Phytohaemagglutinin</a></li> <li><a href="/wiki/Ovomucin" title="Ovomucin">Ovomucin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Proteoglycan" title="Proteoglycan">Proteoglycans</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Chondroitin_sulfate" title="Chondroitin sulfate">CS</a>/<a href="/wiki/Dermatan_sulfate" title="Dermatan sulfate">DS</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Decorin" title="Decorin">Decorin</a></li> <li><a href="/wiki/Biglycan" title="Biglycan">Biglycan</a></li> <li><a href="/wiki/Versican" title="Versican">Versican</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Heparan_sulfate" title="Heparan sulfate">HS</a>/<a href="/wiki/Chondroitin_sulfate" title="Chondroitin sulfate">CS</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Testican" title="Testican">Testican</a></li> <li><a href="/wiki/Perlecan" title="Perlecan">Perlecan</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Chondroitin_sulfate" title="Chondroitin sulfate">CS</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chondroitin_sulfate_proteoglycan" title="Chondroitin sulfate proteoglycan">Chondroitin sulfate proteoglycans</a>: <a href="/wiki/Aggrecan" title="Aggrecan">Aggrecan</a></li> <li><a href="/wiki/Neurocan" title="Neurocan">Neurocan</a></li> <li><a href="/wiki/Brevican" title="Brevican">Brevican</a></li> <li><a href="/wiki/CD44" title="CD44">CD44</a></li> <li><a href="/wiki/CSPG4" title="CSPG4">CSPG4</a></li> <li><a href="/wiki/CSPG5" title="CSPG5">CSPG5</a></li> <li><a href="/wiki/Platelet_factor_4" title="Platelet factor 4">Platelet factor 4</a></li> <li><a href="/wiki/SMC3_(gene)" class="mw-redirect" title="SMC3 (gene)">Structural maintenance of chromosomes 3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Keratan_sulfate" title="Keratan sulfate">KS</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Fibromodulin" title="Fibromodulin">Fibromodulin</a></li> <li><a href="/wiki/Lumican" title="Lumican">Lumican</a></li> <li><a href="/wiki/Keratocan" title="Keratocan">Keratocan</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite"><a href="/wiki/Heparan_sulfate" title="Heparan sulfate">HS</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Syndecan_1" title="Syndecan 1">Syndecan 1</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background-color: AntiqueWhite">Other</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/Activin_and_inhibin" title="Activin and inhibin">Activin and inhibin</a></li> <li><a href="/wiki/A_disintegrin_and_metalloproteinase" class="mw-redirect" title="A disintegrin and metalloproteinase">ADAM</a></li> <li><a href="/wiki/Alpha_1-antichymotrypsin" title="Alpha 1-antichymotrypsin">Alpha 1-antichymotrypsin</a></li> <li><a href="/wiki/Apolipoprotein_H" title="Apolipoprotein H">Apolipoprotein H</a></li> <li><a href="/wiki/CD70" title="CD70">CD70</a></li> <li><a href="/wiki/Asialoglycoprotein" title="Asialoglycoprotein">Asialoglycoprotein</a></li> <li><a href="/wiki/Avidin" title="Avidin">Avidin</a></li> <li><a href="/wiki/B-cell_activating_factor" title="B-cell activating factor">B-cell activating factor</a></li> <li><a href="/wiki/4-1BB_ligand" class="mw-redirect" title="4-1BB ligand">4-1BB ligand</a></li> <li><a href="/wiki/Cholesterylester_transfer_protein" class="mw-redirect" title="Cholesterylester transfer protein">Cholesterylester transfer protein</a></li> <li><a href="/wiki/Clusterin" title="Clusterin">Clusterin</a></li> <li><a href="/wiki/Colony-stimulating_factor" title="Colony-stimulating factor">Colony-stimulating factor</a></li> <li><a href="/wiki/Hemopexin" title="Hemopexin">Hemopexin</a></li> <li><a href="/wiki/Lactoferrin" title="Lactoferrin">Lactoferrin</a></li> <li><a href="/wiki/Membrane_glycoproteins" title="Membrane glycoproteins">Membrane glycoproteins</a></li> <li><a href="/wiki/Myelin_protein_zero" title="Myelin protein zero">Myelin protein zero</a></li> <li><a href="/wiki/Osteonectin" title="Osteonectin">Osteonectin</a></li> <li><a href="/wiki/Protein_C" title="Protein C">Protein C</a></li> <li><a href="/wiki/Protein_S" title="Protein S">Protein S</a></li> <li><a href="/wiki/Serum_amyloid_P_component" title="Serum amyloid P component">Serum amyloid P component</a></li> <li><a href="/wiki/Sialoglycoprotein" title="Sialoglycoprotein">Sialoglycoprotein</a> <ul><li><a href="/wiki/CD43" title="CD43">CD43</a></li> <li><a href="/wiki/Glycophorin" title="Glycophorin">Glycophorin</a></li> <li><a href="/wiki/Glycophorin_C" title="Glycophorin C">Glycophorin C</a></li></ul></li> <li><a href="/wiki/Thrombopoietin" title="Thrombopoietin">Thrombopoietin</a></li> <li><a href="/wiki/Thyroglobulin" title="Thyroglobulin">Thyroglobulin</a></li> <li><a href="/wiki/Thyroxine-binding_proteins" title="Thyroxine-binding proteins">Thyroxine-binding proteins</a></li> <li><a href="/wiki/Transcortin" title="Transcortin">Transcortin</a></li> <li><a href="/wiki/Tumor_necrosis_factor_alpha" class="mw-redirect" title="Tumor necrosis factor alpha">Tumor necrosis factor alpha</a></li> <li><a href="/wiki/Uteroglobin" title="Uteroglobin">Uteroglobin</a></li> <li><a href="/wiki/Vitronectin" title="Vitronectin">Vitronectin</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Microbiology:_Bacteria" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align: center;"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Bacteria" title="Template:Bacteria"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Bacteria" title="Template talk:Bacteria"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Bacteria" title="Special:EditPage/Template:Bacteria"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Microbiology:_Bacteria" style="font-size:114%;margin:0 4em"><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a>: <a href="/wiki/Bacteria" title="Bacteria">Bacteria</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Medical_microbiology" title="Medical microbiology">Medical<br />microbiology</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/Pathogenic_bacteria" title="Pathogenic bacteria">Infection</a></li> <li><a href="/wiki/Exotoxin" title="Exotoxin">Exotoxin</a></li> <li><a href="/wiki/Lysogenic_cycle" title="Lysogenic cycle">Lysogenic cycle</a></li> <li><a href="/wiki/Pathogenic_bacteria" title="Pathogenic bacteria">Pathogenic bacteria</a></li> <li><a href="/wiki/Antimicrobial_resistance" title="Antimicrobial resistance">Resistance</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Biochemistry" title="Biochemistry">Biochemistry</a><br />and <a href="/wiki/Ecology" title="Ecology">ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Oxygen<br />preference</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aerobic_organism" title="Aerobic organism">Aerobic</a> <ul><li><a href="/wiki/Obligate_aerobe" title="Obligate aerobe">Obligate</a></li> <li><a href="/wiki/Facultative_aerobe" class="mw-redirect" title="Facultative aerobe">Facultative</a></li></ul></li> <li><a href="/wiki/Anaerobic_organism" title="Anaerobic organism">Anaerobic</a> <ul><li><a href="/wiki/Facultative_anaerobic_organism" title="Facultative anaerobic organism">Facultative</a></li> <li><a href="/wiki/Obligate_anaerobe" title="Obligate anaerobe">Obligate</a></li></ul></li> <li><a href="/wiki/Microaerophile" title="Microaerophile">Microaerophile</a></li> <li><a href="/wiki/Nanaerobe" title="Nanaerobe">Nanaerobe</a></li> <li><a href="/wiki/Aerotolerant_anaerobe" title="Aerotolerant anaerobe">Aerotolerant</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Extremophile" title="Extremophile">Extremophile</a></li> <li><a href="/wiki/Human_microbiome" title="Human microbiome">Human microbiome</a> <ul><li><a href="/wiki/Gut_microbiota" title="Gut microbiota">Gut</a></li> <li><a href="/wiki/Lung_microbiota" title="Lung microbiota">Lung</a></li> <li><a href="/wiki/Oral_microbiology" title="Oral microbiology">Mouth</a></li> <li><a href="/wiki/Skin_flora" title="Skin flora">Skin</a></li> <li><a href="/wiki/Vaginal_flora" title="Vaginal flora">Vaginal</a> (<a href="/wiki/Vaginal_microbiota_in_pregnancy" class="mw-redirect" title="Vaginal microbiota in pregnancy">in pregnancy</a>)</li> <li><a href="/wiki/Placental_microbiome" title="Placental microbiome">Placental</a></li> <li><a href="/wiki/Uterine_microbiome" title="Uterine microbiome">Uterine</a></li> <li><a href="/wiki/Salivary_microbiome" title="Salivary microbiome">Salivary</a></li></ul></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Nitrogen_fixation" title="Nitrogen fixation">Nitrogen fixation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li>Substrate preference <ul><li><a href="/wiki/Lipophilic_bacteria" title="Lipophilic bacteria">Lipophilic</a></li> <li><a href="/wiki/Osmophile" title="Osmophile">Saccharophilic</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">Shape</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/Bacterial_cellular_morphologies" title="Bacterial cellular morphologies">Bacterial cellular morphologies</a> <ul><li><a href="/wiki/Bacterial_cell_structure" title="Bacterial cell structure">cell structure</a></li> <li><a href="/wiki/Bacterial_morphological_plasticity" title="Bacterial morphological plasticity">plasticity</a></li></ul></li> <li><a href="/wiki/Coccus" class="mw-redirect" title="Coccus">Cocci</a></li> <li><a href="/wiki/Bacillus_(shape)" class="mw-redirect" title="Bacillus (shape)">Bacilli</a></li> <li><a href="/wiki/Coccobacillus" class="mw-redirect" title="Coccobacillus">Coccobacilli</a></li> <li><a href="/wiki/Spiral_bacteria" class="mw-redirect" title="Spiral bacteria">Spiral</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Bacterial_cell_structure" title="Bacterial cell structure">Structure</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;"><a href="/wiki/Cell_envelope" title="Cell envelope">Cell<br />envelope</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_membrane" title="Cell membrane">Cell membrane</a></li> <li><a href="/wiki/Cell_wall" title="Cell wall">Cell wall</a>: <a class="mw-selflink selflink">Peptidoglycan</a> <ul><li><a href="/wiki/N-Acetylmuramic_acid" title="N-Acetylmuramic acid">NAM</a></li> <li><a href="/wiki/N-Acetylglucosamine" title="N-Acetylglucosamine">NAG</a></li> <li><a href="/wiki/Diaminopimelic_acid" title="Diaminopimelic acid">DAP</a></li></ul></li> <li><i><a href="/wiki/Gram-positive_bacteria" title="Gram-positive bacteria">Gram-positive bacteria</a> only:</i> <a href="/wiki/Teichoic_acid" title="Teichoic acid">Teichoic acid</a></li> <li><a href="/wiki/Lipoteichoic_acid" title="Lipoteichoic acid">Lipoteichoic acid</a></li> <li><a href="/wiki/Endospore" title="Endospore">Endospore</a></li> <li><i><a href="/wiki/Gram-negative_bacteria" title="Gram-negative bacteria">Gram-negative bacteria</a> only:</i> <a href="/wiki/Bacterial_outer_membrane" title="Bacterial outer membrane">Bacterial outer membrane</a> <ul><li><a href="/wiki/General_bacterial_porin_family" title="General bacterial porin family">Porin</a></li> <li><a href="/wiki/Lipopolysaccharide" title="Lipopolysaccharide">Lipopolysaccharide</a></li></ul></li> <li><a href="/wiki/Periplasmic_space" class="mw-redirect" title="Periplasmic space">Periplasmic space</a></li> <li><i><a href="/wiki/Mycobacterium" title="Mycobacterium">Mycobacteria</a> only:</i> <a href="/wiki/Arabinogalactan" title="Arabinogalactan">Arabinogalactan</a></li> <li><a href="/wiki/Mycolic_acid" title="Mycolic acid">Mycolic acid</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Outside<br />envelope</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_capsule" title="Bacterial capsule">Bacterial capsule</a></li> <li><a href="/wiki/Slime_layer" title="Slime layer">Slime layer</a></li> <li><a href="/wiki/S-layer" title="S-layer">S-layer</a></li> <li><a href="/wiki/Glycocalyx" title="Glycocalyx">Glycocalyx</a></li> <li><a href="/wiki/Pilus" title="Pilus">Pilus</a></li> <li><a href="/wiki/Fimbria_(bacteriology)" class="mw-redirect" title="Fimbria (bacteriology)">Fimbria</a></li> <li><a href="/wiki/Non-motile_bacteria" title="Non-motile bacteria">Non-motile bacteria</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;">Composite</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biofilm" title="Biofilm">Biofilm</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="/wiki/Bacterial_taxonomy" title="Bacterial taxonomy">Taxonomy</a><br />and <a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">evolution</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="https://species.wikimedia.org/wiki/Bacteria_(classifications)" class="extiw" title="wikispecies:Bacteria (classifications)">Bacteria (classifications)</a></li> <li><a href="/wiki/Bacterial_phyla" title="Bacterial phyla">Bacterial phyla</a></li> <li><i>Former groupings</i>: <a href="/wiki/Schizomycetes" class="mw-redirect" title="Schizomycetes">Schizomycetes</a></li> <li><a href="/wiki/Monera" title="Monera">Monera</a></li> <li><a href="/wiki/Prokaryota" class="mw-redirect" title="Prokaryota">Prokaryota</a> <ul><li><a href="/wiki/Gracilicutes" title="Gracilicutes">Gracilicutes</a></li> <li><a href="/wiki/Bacillota" title="Bacillota">Firmicutes</a></li> <li><a href="/wiki/Mollicutes" title="Mollicutes">Mollicutes</a></li> <li><a href="/wiki/Mendosicutes" class="mw-redirect" title="Mendosicutes">Mendosicutes</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="2" style="text-align: center;"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Bacteria" title="Category:Bacteria">Category</a></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, 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