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class="vector-toc-numb">2</span> <span>Activation</span> </div> </a> <button aria-controls="toc-Activation-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 Activation subsection</span> </button> <ul id="toc-Activation-sublist" class="vector-toc-list"> <li id="toc-T_cell-dependent_activation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#T_cell-dependent_activation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>T cell-dependent activation</span> </div> </a> <ul id="toc-T_cell-dependent_activation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-T_cell-independent_activation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#T_cell-independent_activation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>T cell-independent activation</span> </div> </a> <ul id="toc-T_cell-independent_activation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Memory_B_cell_activation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Memory_B_cell_activation"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Memory B cell activation</span> </div> </a> <ul id="toc-Memory_B_cell_activation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-B_cell_types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#B_cell_types"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>B cell types</span> </div> </a> <ul id="toc-B_cell_types-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-B_cell-related_pathology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#B_cell-related_pathology"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>B cell-related pathology</span> </div> </a> <ul id="toc-B_cell-related_pathology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Epigenetics" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Epigenetics"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Epigenetics</span> </div> </a> <ul id="toc-Epigenetics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><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">B cell</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 55 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-55" 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">55 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%AE%D9%84%D9%8A%D8%A9_%D8%A8%D8%A7%D8%A6%D9%8A%D8%A9" title="خلية بائية – Arabic" lang="ar" hreflang="ar" data-title="خلية بائية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AC%E0%A6%BF_%E0%A6%95%E0%A7%8B%E0%A6%B7" title="বি কোষ – Bangla" lang="bn" hreflang="bn" data-title="বি কোষ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/B-%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="B-клетка – Bulgarian" lang="bg" hreflang="bg" data-title="B-клетка" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/B-%C4%87elija" title="B-ćelija – Bosnian" lang="bs" hreflang="bs" data-title="B-ćelija" 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/Limf%C3%B2cit_B" title="Limfòcit B – Catalan" lang="ca" hreflang="ca" data-title="Limfòcit B" 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/B-lymfocyt" title="B-lymfocyt – Czech" lang="cs" hreflang="cs" data-title="B-lymfocyt" 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/B-celle" title="B-celle – Danish" lang="da" hreflang="da" data-title="B-celle" 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/B-Lymphozyt" title="B-Lymphozyt – German" lang="de" hreflang="de" data-title="B-Lymphozyt" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-dv mw-list-item"><a href="https://dv.wikipedia.org/wiki/%DE%84%DE%A9_%DE%8D%DE%A8%DE%89%DE%B0%DE%8A%DE%A6%DE%90%DE%A6%DE%87%DE%A8%DE%93%DE%B0" title="ބީ ލިމްފަސައިޓް – Divehi" lang="dv" hreflang="dv" data-title="ބީ ލިމްފަސައިޓް" data-language-autonym="ދިވެހިބަސް" data-language-local-name="Divehi" class="interlanguage-link-target"><span>ދިވެހިބަސް</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/B-l%C3%BCmfots%C3%BC%C3%BCdid" title="B-lümfotsüüdid – Estonian" lang="et" hreflang="et" data-title="B-lümfotsüüdid" 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%92_%CE%BA%CF%8D%CF%84%CF%84%CE%B1%CF%81%CE%B1" title="Β κύτταρα – Greek" lang="el" hreflang="el" data-title="Β κύτταρα" 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/Linfocito_B" title="Linfocito B – Spanish" lang="es" hreflang="es" data-title="Linfocito B" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/B-%C4%89elo" title="B-ĉelo – Esperanto" lang="eo" hreflang="eo" data-title="B-ĉelo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/B_linfozito" title="B linfozito – Basque" lang="eu" hreflang="eu" data-title="B linfozito" 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%84%D9%86%D9%81%D9%88%D8%B3%DB%8C%D8%AA_%D8%A8%DB%8C" title="لنفوسیت بی – Persian" lang="fa" hreflang="fa" data-title="لنفوسیت بی" 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/Lymphocyte_B" title="Lymphocyte B – French" lang="fr" hreflang="fr" data-title="Lymphocyte B" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Linfocito_B" title="Linfocito B – Galician" lang="gl" hreflang="gl" data-title="Linfocito B" 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/B%EC%84%B8%ED%8F%AC" title="B세포 – Korean" lang="ko" hreflang="ko" data-title="B세포" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/B_%D5%AC%D5%AB%D5%B4%D6%86%D5%B8%D6%81%D5%AB%D5%BF%D5%B6%D5%A5%D6%80" title="B լիմֆոցիտներ – Armenian" lang="hy" hreflang="hy" data-title="B լիմֆոցիտներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Sel_B" title="Sel B – Indonesian" lang="id" hreflang="id" data-title="Sel B" 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-is mw-list-item"><a href="https://is.wikipedia.org/wiki/B-eitilfruma" title="B-eitilfruma – Icelandic" lang="is" hreflang="is" data-title="B-eitilfruma" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Linfocita_B" title="Linfocita B – Italian" lang="it" hreflang="it" data-title="Linfocita B" 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%9C%D7%99%D7%9E%D7%A4%D7%95%D7%A6%D7%99%D7%98_B" title="לימפוציט B – Hebrew" lang="he" hreflang="he" data-title="לימפוציט B" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/B_%E1%83%9A%E1%83%98%E1%83%9B%E1%83%A4%E1%83%9D%E1%83%AA%E1%83%98%E1%83%A2%E1%83%98" title="B ლიმფოციტი – Georgian" lang="ka" hreflang="ka" data-title="B ლიმფოციტი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%92-%D0%BB%D0%B8%D0%BC%D1%84%D0%BE%D1%86%D0%B8%D1%82%D1%82%D0%B5%D1%80" title="В-лимфоциттер – Kazakh" lang="kk" hreflang="kk" data-title="В-лимфоциттер" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Xaneya_B" title="Xaneya B – Kurdish" lang="ku" hreflang="ku" data-title="Xaneya B" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Lymphocytus_B" title="Lymphocytus B – Latin" lang="la" hreflang="la" data-title="Lymphocytus B" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/B_limfocitai" title="B limfocitai – Lithuanian" lang="lt" hreflang="lt" data-title="B limfocitai" 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/B-limfocita" title="B-limfocita – Hungarian" lang="hu" hreflang="hu" data-title="B-limfocita" 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/B-%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="B-клетка – Macedonian" lang="mk" hreflang="mk" data-title="B-клетка" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%AC%E0%B4%BF-%E0%B4%B2%E0%B4%B8%E0%B4%BF%E0%B4%95%E0%B4%BE%E0%B4%A3%E0%B5%81" title="ബി-ലസികാണു – Malayalam" lang="ml" hreflang="ml" data-title="ബി-ലസികാണു" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Sel_B" title="Sel B – Malay" lang="ms" hreflang="ms" data-title="Sel B" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%92_%D1%8D%D1%81" title="В эс – Mongolian" lang="mn" hreflang="mn" data-title="В эс" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/B-cel" title="B-cel – Dutch" lang="nl" hreflang="nl" data-title="B-cel" 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/B%E7%B4%B0%E8%83%9E" title="B細胞 – Japanese" lang="ja" hreflang="ja" data-title="B細胞" 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/B-celler" title="B-celler – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="B-celler" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Linfocit_B" title="Linfocit B – Occitan" lang="oc" hreflang="oc" data-title="Linfocit B" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Limfocyty_B" title="Limfocyty B – Polish" lang="pl" hreflang="pl" data-title="Limfocyty B" 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/Linf%C3%B3cito_B" title="Linfócito B – Portuguese" lang="pt" hreflang="pt" data-title="Linfócito B" 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/Celule_B" title="Celule B – Romanian" lang="ro" hreflang="ro" data-title="Celule B" 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/B-%D0%BB%D0%B8%D0%BC%D1%84%D0%BE%D1%86%D0%B8%D1%82%D1%8B" title="B-лимфоциты – Russian" lang="ru" hreflang="ru" data-title="B-лимфоциты" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Limfociti-B" title="Limfociti-B – Albanian" lang="sq" hreflang="sq" data-title="Limfociti-B" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/B_cell" title="B cell – Simple English" lang="en-simple" hreflang="en-simple" data-title="B cell" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/B-lymfocyt" title="B-lymfocyt – Slovak" lang="sk" hreflang="sk" data-title="B-lymfocyt" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Limfocit_B" title="Limfocit B – Slovenian" lang="sl" hreflang="sl" data-title="Limfocit B" 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/%D0%91-%D1%9B%D0%B5%D0%BB%D0%B8%D1%98%D0%B0" title="Б-ћелија – Serbian" lang="sr" hreflang="sr" data-title="Б-ћелија" 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-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/B-solu" title="B-solu – Finnish" lang="fi" hreflang="fi" data-title="B-solu" 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/B-cell" title="B-cell – Swedish" lang="sv" hreflang="sv" data-title="B-cell" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AA%E0%AE%BF_%E0%AE%89%E0%AE%AF%E0%AE%BF%E0%AE%B0%E0%AE%A3%E0%AF%81" title="பி உயிரணு – Tamil" lang="ta" hreflang="ta" data-title="பி உயிரணு" 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class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Type of white blood cell</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">This article is about the immune system cell. For the electrical cell, see <a href="/wiki/Battery_(vacuum_tube)" class="mw-redirect" title="Battery (vacuum tube)">Battery (vacuum tube)</a>. For β-cell of pancreas, see <a href="/wiki/Beta_cell" title="Beta cell">beta cell</a>.</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:beige">B lymphocyte cell</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Blausen_0624_Lymphocyte_B_cell_(crop).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Blausen_0624_Lymphocyte_B_cell_%28crop%29.png/250px-Blausen_0624_Lymphocyte_B_cell_%28crop%29.png" decoding="async" width="250" height="242" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Blausen_0624_Lymphocyte_B_cell_%28crop%29.png/375px-Blausen_0624_Lymphocyte_B_cell_%28crop%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Blausen_0624_Lymphocyte_B_cell_%28crop%29.png/500px-Blausen_0624_Lymphocyte_B_cell_%28crop%29.png 2x" data-file-width="1165" data-file-height="1130" /></a></span><div class="infobox-caption">Animation of B cell</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Embryology" title="Embryology">Precursor</a></th><td class="infobox-data"><a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">Hematopoietic stem cell</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Organ_system" title="Organ system">System</a></th><td class="infobox-data"><a href="/wiki/Immune_system" title="Immune system">Immune system</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>lymphocytus B</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D001402">D001402</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma62869">62869</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_microanatomy" title="Anatomical terms of microanatomy">Anatomical terms of microanatomy</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q188930" class="extiw" title="d:Q188930">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p><b>B cells</b>, also known as <b>B lymphocytes</b>, are a type of <a href="/wiki/White_blood_cell" title="White blood cell">white blood cell</a> of the <a href="/wiki/Lymphocyte" title="Lymphocyte">lymphocyte</a> subtype.<sup id="cite_ref-:0_1-0" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> They function in the <a href="/wiki/Humoral_immunity" title="Humoral immunity">humoral immunity</a> component of the <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a>.<sup id="cite_ref-:0_1-1" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> B cells produce <a href="/wiki/Antibody" title="Antibody">antibody</a> molecules which may be either secreted or inserted into the plasma membrane where they serve as a part of <a href="/wiki/B-cell_receptor" title="B-cell receptor">B-cell receptors</a>.<sup id="cite_ref-Alberts_2002_2-0" class="reference"><a href="#cite_note-Alberts_2002-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> When a naïve or <a href="/wiki/Memory_B_cell" title="Memory B cell">memory B cell</a> is activated by an antigen, it proliferates and differentiates into an antibody-secreting effector cell, known as a plasmablast or plasma cell.<sup id="cite_ref-Alberts_2002_2-1" class="reference"><a href="#cite_note-Alberts_2002-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In addition, B cells <a href="/wiki/Antigen_presentation" title="Antigen presentation">present antigens</a> (they are also classified as professional <a href="/wiki/Antigen-presenting_cell" title="Antigen-presenting cell">antigen-presenting cells, APCs</a>) and secrete <a href="/wiki/Cytokine" title="Cytokine">cytokines</a>.<sup id="cite_ref-:0_1-2" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Mammal" title="Mammal">mammals</a> B cells <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">mature</a> in the <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a>, which is at the core of most <a href="/wiki/Bone" title="Bone">bones</a>.<sup id="cite_ref-:1_3-0" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Bird" title="Bird">birds</a>, B cells mature in the <a href="/wiki/Bursa_of_Fabricius" title="Bursa of Fabricius">bursa of Fabricius</a>, a lymphoid organ where they were first discovered by Chang and Glick,<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> which is why the <i>B</i> stands for <i>bursa</i> and not <i>bone marrow</i>, as commonly believed. </p><p>B cells, unlike the other two classes of lymphocytes, <a href="/wiki/T_cell" title="T cell">T cells</a> and <a href="/wiki/Natural_killer_cell" title="Natural killer cell">natural killer cells</a>, express <a href="/wiki/B-cell_receptor" title="B-cell receptor">B cell receptors (BCRs)</a> on their <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a>.<sup id="cite_ref-:0_1-3" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> BCRs allow the B cell to <a href="/wiki/Molecular_binding" title="Molecular binding">bind</a> to a foreign <a href="/wiki/Antigen" title="Antigen">antigen</a>, against which it will initiate an antibody response.<sup id="cite_ref-:0_1-4" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> B cell receptors are extremely specific, with all BCRs on a B cell recognizing the same <a href="/wiki/Epitope" title="Epitope">epitope</a>.<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> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Development">Development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=1" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Early_B_cell_development.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Early_B_cell_development.jpg/250px-Early_B_cell_development.jpg" decoding="async" width="220" height="81" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/56/Early_B_cell_development.jpg/330px-Early_B_cell_development.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/56/Early_B_cell_development.jpg/500px-Early_B_cell_development.jpg 2x" data-file-width="2295" data-file-height="846" /></a><figcaption>Early B cell development: from stem cell to immature B cell</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Transitional_B_cell_development.PNG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Transitional_B_cell_development.PNG/250px-Transitional_B_cell_development.PNG" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/d/dd/Transitional_B_cell_development.PNG 1.5x" data-file-width="303" data-file-height="217" /></a><figcaption><a href="/wiki/Transitional_B_cell" title="Transitional B cell">Transitional B cell</a> development: from immature B cell to MZ B cell or mature (FO) B cell</figcaption></figure> <p>B cells develop from <a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">hematopoietic stem cells (HSCs)</a> that originate from <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a>.<sup id="cite_ref-BCDTfBcells_6-0" class="reference"><a href="#cite_note-BCDTfBcells-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_7-0" class="reference"><a href="#cite_note-:2-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> HSCs first differentiate into <a href="/wiki/Multipotent_progenitor" class="mw-redirect" title="Multipotent progenitor">multipotent progenitor</a> (MPP) cells, then <a href="/wiki/Common_lymphoid_progenitor" class="mw-redirect" title="Common lymphoid progenitor">common lymphoid progenitor</a> (CLP) cells.<sup id="cite_ref-:2_7-1" class="reference"><a href="#cite_note-:2-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> From here, their development into B cells occurs in several stages (shown in image to the right), each marked by various <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> patterns and <a href="/wiki/Antibody" title="Antibody">immunoglobulin</a> <a href="/wiki/Immunoglobulin_heavy_chain" title="Immunoglobulin heavy chain">H chain</a> and <a href="/wiki/Immunoglobulin_light_chain" title="Immunoglobulin light chain">L chain</a> <a href="/wiki/Locus_(genetics)" title="Locus (genetics)">gene loci</a> arrangements, the latter due to B cells undergoing <a href="/wiki/V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a> as they develop.<sup id="cite_ref-:3_8-0" class="reference"><a href="#cite_note-:3-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>B cells undergo two types of selection while developing in the bone marrow to ensure proper development, both involving B cell receptors (BCR) on the surface of the cell. Positive selection occurs through antigen-independent signalling involving both the pre-BCR and the BCR.<sup id="cite_ref-:4_9-0" class="reference"><a href="#cite_note-:4-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_10-0" class="reference"><a href="#cite_note-:5-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> If these receptors do not bind to their <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligand</a>, B cells do not receive the proper signals and cease to develop.<sup id="cite_ref-:4_9-1" class="reference"><a href="#cite_note-:4-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_10-1" class="reference"><a href="#cite_note-:5-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Negative selection occurs through the binding of self-antigen with the BCR; if the BCR can bind strongly to self-antigen, then the B cell undergoes one of four fates: <a href="/wiki/Clonal_deletion" title="Clonal deletion">clonal deletion</a>, <a href="/wiki/Receptor_editing" title="Receptor editing">receptor editing</a>, <a href="/wiki/Clonal_anergy" title="Clonal anergy">anergy</a>, or ignorance (B cell ignores signal and continues development).<sup id="cite_ref-:5_10-2" class="reference"><a href="#cite_note-:5-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> This negative selection process leads to a state of <a href="/wiki/Central_tolerance" title="Central tolerance">central tolerance</a>, in which the mature B cells do not bind self antigens present in the bone marrow.<sup id="cite_ref-:3_8-1" class="reference"><a href="#cite_note-:3-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>To complete development, immature B cells migrate from the bone marrow into the spleen as <a href="/wiki/Transitional_B_cells" class="mw-redirect" title="Transitional B cells">transitional B cells</a>, passing through two transitional stages: T1 and T2.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Throughout their migration to the spleen and after spleen entry, they are considered T1 B cells.<sup id="cite_ref-:6_12-0" class="reference"><a href="#cite_note-:6-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Within the spleen, T1 B cells transition to T2 B cells.<sup id="cite_ref-:6_12-1" class="reference"><a href="#cite_note-:6-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> T2 B cells differentiate into either follicular (FO) B cells or marginal zone (MZ) B cells depending on signals received through the BCR and other receptors.<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> Once differentiated, they are now considered mature B cells, or naïve B cells.<sup id="cite_ref-:6_12-2" class="reference"><a href="#cite_note-:6-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Activation">Activation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=2" title="Edit section: Activation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:B_cell_activation_naive_to_plasma_cell.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/B_cell_activation_naive_to_plasma_cell.png/220px-B_cell_activation_naive_to_plasma_cell.png" decoding="async" width="220" height="124" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/B_cell_activation_naive_to_plasma_cell.png/330px-B_cell_activation_naive_to_plasma_cell.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e9/B_cell_activation_naive_to_plasma_cell.png/440px-B_cell_activation_naive_to_plasma_cell.png 2x" data-file-width="463" data-file-height="261" /></a><figcaption>B cell activation: from immature B cell to plasma cell or memory B cell</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:B_cell_function.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/B_cell_function.png/220px-B_cell_function.png" decoding="async" width="220" height="76" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c5/B_cell_function.png/330px-B_cell_function.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c5/B_cell_function.png/440px-B_cell_function.png 2x" data-file-width="813" data-file-height="280" /></a><figcaption>Basic B cell function: bind to an antigen, receive help from a cognate helper T cell, and differentiate into a <a href="/wiki/Plasma_cell" title="Plasma cell">plasma cell</a> that secretes large numbers of antibodies</figcaption></figure> <p>B cell activation occurs in the <a href="/wiki/Secondary_lymphoid_organs" class="mw-redirect" title="Secondary lymphoid organs">secondary lymphoid organs</a> (SLOs), such as the <a href="/wiki/Spleen" title="Spleen">spleen</a> and <a href="/wiki/Lymph_nodes" class="mw-redirect" title="Lymph nodes">lymph nodes</a>.<sup id="cite_ref-:0_1-5" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> After B cells mature in the bone marrow, they migrate through the blood to SLOs, which receive a constant supply of antigen through circulating <a href="/wiki/Lymph" title="Lymph">lymph</a>.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> At the SLO, B cell activation begins when the B cell binds to an antigen via its BCR.<sup id="cite_ref-:7_15-0" class="reference"><a href="#cite_note-:7-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> Although the events taking place immediately after activation have yet to be completely determined, it is believed that B cells are activated in accordance with the kinetic segregation model <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="the mechanism of activation is actually quite controversial (April 2019)">citation needed</span></a></i>]</sup>, initially determined in T lymphocytes. This model denotes that before antigen stimulation, receptors diffuse through the membrane coming into contact with <a href="/wiki/Lck" class="mw-redirect" title="Lck">Lck</a> and <a href="/wiki/CD45" class="mw-redirect" title="CD45">CD45</a> in equal frequency, rendering a net equilibrium of phosphorylation and non-phosphorylation. It is only when the cell comes in contact with an antigen presenting cell that the larger CD45 is displaced due to the close distance between the two membranes. This allows for net phosphorylation of the BCR and the initiation of the signal transduction pathway<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. (April 2019)">citation needed</span></a></i>]</sup>. Of the three B cell subsets, FO B cells preferentially undergo T cell-dependent activation while MZ B cells and B1 B cells preferentially undergo T cell-independent activation.<sup id="cite_ref-:8_16-0" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>B cell activation is enhanced through the activity of <a href="/wiki/Complement_receptor_2" title="Complement receptor 2">CD21</a>, a surface receptor in complex with surface proteins <a href="/wiki/CD19" title="CD19">CD19</a> and <a href="/wiki/CD81" title="CD81">CD81</a> (all three are collectively known as the B cell coreceptor complex).<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> When a BCR binds an antigen tagged with a fragment of the C3 complement protein, CD21 binds the C3 fragment, co-ligates with the bound BCR, and signals are transduced through CD19 and CD81 to lower the activation threshold of the cell.<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-heading3"><h3 id="T_cell-dependent_activation">T cell-dependent activation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=3" title="Edit section: T cell-dependent activation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Antigens that activate B cells with the help of T-cell are known as T cell-dependent (TD) antigens and include foreign proteins.<sup id="cite_ref-:0_1-6" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> They are named as such because they are unable to induce a humoral response in organisms that lack T cells.<sup id="cite_ref-:0_1-7" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> B cell responses to these antigens takes multiple days, though antibodies generated have a higher affinity and are more functionally versatile than those generated from T cell-independent activation.<sup id="cite_ref-:0_1-8" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p> Once a BCR binds a TD antigen, the antigen is taken up into the B cell through <a href="/wiki/Receptor-mediated_endocytosis" title="Receptor-mediated endocytosis">receptor-mediated endocytosis</a>, <a href="/wiki/Proteolysis" title="Proteolysis">degraded</a>, and presented to T cells as peptide pieces in complex with <a href="/wiki/MHC_class_II" title="MHC class II">MHC-II molecules</a> on the cell membrane.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> <a href="/wiki/T_helper_cell" title="T helper cell">T helper (T<sub>H</sub>) cells</a>, typically <a href="/wiki/Follicular_B_helper_T_cells" title="Follicular B helper T cells">follicular T helper (T<sub>FH</sub>) cells</a> recognize and bind these MHC-II-peptide complexes through their <a href="/wiki/T_cell_receptor" class="mw-redirect" title="T cell receptor">T cell receptor (TCR)</a>.<sup id="cite_ref-:9_20-0" class="reference"><a href="#cite_note-:9-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Following TCR-MHC-II-peptide binding, T cells express the surface protein <a href="/wiki/CD154" title="CD154">CD40L</a> as well as cytokines such as <a href="/wiki/Interleukin_4" title="Interleukin 4">IL-4</a> and <a href="/wiki/Interleukin_21" title="Interleukin 21">IL-21</a>.<sup id="cite_ref-:9_20-1" class="reference"><a href="#cite_note-:9-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> CD40L serves as a necessary co-stimulatory factor for B cell activation by binding the B cell surface receptor <a href="/wiki/CD40_(protein)" title="CD40 (protein)">CD40</a>, which promotes B cell <a href="/wiki/Cell_growth" title="Cell growth">proliferation</a>, <a href="/wiki/Immunoglobulin_class_switching" title="Immunoglobulin class switching">immunoglobulin class switching</a>, and <a href="/wiki/Somatic_hypermutation" title="Somatic hypermutation">somatic hypermutation</a> as well as sustains T cell growth and differentiation.<sup id="cite_ref-:0_1-9" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> T cell-derived cytokines bound by B cell <a href="/wiki/Cytokine_receptor" title="Cytokine receptor">cytokine receptors</a> also promote B cell proliferation, immunoglobulin class switching, and somatic hypermutation as well as guide differentiation.<sup id="cite_ref-:9_20-2" class="reference"><a href="#cite_note-:9-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> After B cells receive these signals, they are considered activated.<sup id="cite_ref-:9_20-3" class="reference"><a href="#cite_note-:9-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:T-dependent_B_cell_activation.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/T-dependent_B_cell_activation.png/250px-T-dependent_B_cell_activation.png" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/T-dependent_B_cell_activation.png/330px-T-dependent_B_cell_activation.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/T-dependent_B_cell_activation.png/500px-T-dependent_B_cell_activation.png 2x" data-file-width="960" data-file-height="720" /></a><figcaption>T-dependent B cell activation</figcaption></figure> <p>Once activated, B cells participate in a two-step differentiation process that yields both short-lived plasmablasts for immediate protection and <a href="/wiki/Long-lived_plasma_cell" title="Long-lived plasma cell">long-lived plasma cells</a> and memory B cells for persistent protection.<sup id="cite_ref-:8_16-1" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The first step, known as the extrafollicular response, occurs outside lymphoid follicles but still in the SLO.<sup id="cite_ref-:8_16-2" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> During this step activated B cells proliferate, may undergo immunoglobulin class switching, and differentiate into plasmablasts that produce early, weak antibodies mostly of class IgM.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Dark,_light,_mantle_and_marginal_zones_of_a_secondary_follicle.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png/220px-Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png" decoding="async" width="220" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png/330px-Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8b/Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png/440px-Dark%2C_light%2C_mantle_and_marginal_zones_of_a_secondary_follicle.png 2x" data-file-width="1313" data-file-height="979" /></a><figcaption>Histology of a normal <a href="/wiki/Lymphoid_follicle" class="mw-redirect" title="Lymphoid follicle">lymphoid follicle</a>, with germinal center in the middle.</figcaption></figure> <p>The second step consists of activated B cells entering a lymphoid follicle and forming a <a href="/wiki/Germinal_center" title="Germinal center">germinal center (GC)</a>, which is a specialized microenvironment where B cells undergo extensive proliferation, immunoglobulin class switching, and <a href="/wiki/Affinity_maturation" title="Affinity maturation">affinity maturation</a> directed by somatic hypermutation.<sup id="cite_ref-:10_22-0" class="reference"><a href="#cite_note-:10-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> These processes are facilitated by T<sub>FH</sub> and follicular dendritic cells within the GC and generate both high-affinity memory B cells and long-lived plasma cells.<sup id="cite_ref-:8_16-3" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Resultant plasma cells secrete large numbers of antibodies and either stay within the SLO or, more preferentially, migrate to bone marrow.<sup id="cite_ref-:10_22-1" class="reference"><a href="#cite_note-:10-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="T_cell-independent_activation">T cell-independent activation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=4" title="Edit section: T cell-independent activation"><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/T_independent_antigen_(TI)" title="T independent antigen (TI)">T independent antigen (TI)</a></div> <p>Antigens that activate B cells without T cell help are known as <a href="/wiki/T_independent_antigen_(TI)" title="T independent antigen (TI)">T cell-independent (TI) antigens</a><sup id="cite_ref-:0_1-10" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> and include foreign polysaccharides and unmethylated CpG DNA.<sup id="cite_ref-:8_16-4" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> They are named as such because they are able to induce a humoral response in organisms that lack T cells.<sup id="cite_ref-:0_1-11" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> B cell response to these antigens is rapid, though antibodies generated tend to have lower affinity and are less functionally versatile than those generated from T cell-dependent activation.<sup id="cite_ref-:0_1-12" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>As with TD antigens, B cells activated by TI antigens need additional signals to complete activation, but instead of receiving them from T cells, they are provided either by recognition and binding of a common microbial constituent to <a href="/wiki/Toll-like_receptor" title="Toll-like receptor">toll-like receptors (TLRs)</a> or by extensive crosslinking of BCRs to repeated epitopes on a bacterial cell.<sup id="cite_ref-:0_1-13" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> B cells activated by TI antigens go on to proliferate outside lymphoid follicles but still in SLOs (GCs do not form), possibly undergo immunoglobulin class switching, and differentiate into short-lived plasmablasts that produce early, weak antibodies mostly of class IgM, but also some populations of long-lived plasma cells.<sup id="cite_ref-:11_24-0" class="reference"><a href="#cite_note-:11-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Memory_B_cell_activation">Memory B cell activation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=5" title="Edit section: Memory B cell activation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Memory B cell activation begins with the detection and binding of their target antigen, which is shared by their parent B cell.<sup id="cite_ref-:13_25-0" class="reference"><a href="#cite_note-:13-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Some memory B cells can be activated without T cell help, such as certain virus-specific memory B cells, but others need T cell help.<sup id="cite_ref-:12_26-0" class="reference"><a href="#cite_note-:12-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Upon antigen binding, the memory B cell takes up the antigen through receptor-mediated endocytosis, degrades it, and presents it to T cells as peptide pieces in complex with MHC-II molecules on the cell membrane.<sup id="cite_ref-:13_25-1" class="reference"><a href="#cite_note-:13-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Memory T helper (T<sub>H</sub>) cells, typically memory follicular T helper (T<sub>FH</sub>) cells, that were derived from T cells activated with the same antigen recognize and bind these MHC-II-peptide complexes through their TCR.<sup id="cite_ref-:13_25-2" class="reference"><a href="#cite_note-:13-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Following TCR-MHC-II-peptide binding and the relay of other signals from the memory T<sub>FH</sub> cell, the memory B cell is activated and differentiates either into plasmablasts and plasma cells via an extrafollicular response or enter a germinal center reaction where they generate plasma cells and more memory B cells.<sup id="cite_ref-:13_25-3" class="reference"><a href="#cite_note-:13-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:12_26-1" class="reference"><a href="#cite_note-:12-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> It is unclear whether the memory B cells undergo further affinity maturation within these secondary GCs.<sup id="cite_ref-:13_25-4" class="reference"><a href="#cite_note-:13-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/In_vitro" title="In vitro">In vitro</a></i> activation of memory B cells can be achieved through stimulation with various activators, such as pokeweed mitogen or anti-<a href="/wiki/CD40_(protein)" title="CD40 (protein)">CD40</a> <a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">monoclonal antibodies</a>, however, a study found a combination of <a href="/wiki/Resiquimod" title="Resiquimod">R-848</a> and <a href="/wiki/Interleukin_2" title="Interleukin 2">recombinant human IL-2</a> to be the most efficient activator.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="B_cell_types">B cell types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=6" title="Edit section: B cell types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Plasmablast,_Wright_stain.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Plasmablast%2C_Wright_stain.png/250px-Plasmablast%2C_Wright_stain.png" decoding="async" width="220" height="144" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Plasmablast%2C_Wright_stain.png/330px-Plasmablast%2C_Wright_stain.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Plasmablast%2C_Wright_stain.png/500px-Plasmablast%2C_Wright_stain.png 2x" data-file-width="2000" data-file-height="1309" /></a><figcaption>Plasmablast, <a href="/wiki/Wright_stain" class="mw-redirect" title="Wright stain">Wright stain</a>.</figcaption></figure> <dl><dt>Plasmablast</dt> <dd>A short-lived, proliferating antibody-secreting cell arising from B cell differentiation.<sup id="cite_ref-:0_1-14" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Plasmablasts are generated early in an infection and their antibodies tend to have a weaker affinity towards their target antigen compared to plasma cell.<sup id="cite_ref-:8_16-5" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Plasmablasts can result from T cell-independent activation of B cells or the extrafollicular response from T cell-dependent activation of B cells.<sup id="cite_ref-:0_1-15" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></dd> <dt><a href="/wiki/Plasma_cell" title="Plasma cell">Plasma cell</a></dt> <dd>A long-lived, non-proliferating antibody-secreting cell arising from B cell differentiation.<sup id="cite_ref-:0_1-16" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> There is evidence that B cells first differentiate into a plasmablast-like cell, then differentiate into a plasma cell.<sup id="cite_ref-:8_16-6" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Plasma cells are generated later in an infection and, compared to plasmablasts, have antibodies with a higher affinity towards their target antigen due to affinity maturation in the germinal center (GC) and produce more antibodies.<sup id="cite_ref-:8_16-7" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Plasma cells typically result from the germinal center reaction from T cell-dependent activation of B cells, though they can also result from T cell-independent activation of B cells.<sup id="cite_ref-:11_24-1" class="reference"><a href="#cite_note-:11-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup></dd> <dt>Lymphoplasmacytoid cell</dt> <dd>A cell with a mixture of B lymphocyte and plasma cell morphological features that is thought to be closely related to or a subtype of plasma cells. This cell type is found in pre-malignant and malignant <a href="/wiki/Plasma_cell_dyscrasia" class="mw-redirect" title="Plasma cell dyscrasia">plasma cell dyscrasias</a> that are associated with the secretion of <a href="/wiki/IgM" class="mw-redirect" title="IgM">IgM</a> monoclonal proteins; these dyscrasias include <a href="/wiki/Plasma_cell_dyscrasia#IgM_MGUS" class="mw-redirect" title="Plasma cell dyscrasia">IgM monoclonal gammopathy of undetermined significance</a> and <a href="/wiki/Waldenstr%C3%B6m%27s_macroglobulinemia" class="mw-redirect" title="Waldenström's macroglobulinemia">Waldenström's macroglobulinemia</a>.<sup id="cite_ref-pmid25899140_28-0" class="reference"><a href="#cite_note-pmid25899140-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup></dd> <dt><a href="/wiki/Memory_B_cell" title="Memory B cell">Memory B cell</a></dt> <dd>Dormant B cell arising from B cell differentiation.<sup id="cite_ref-:0_1-17" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Their function is to circulate through the body and initiate a stronger, more rapid antibody response (known as the anamnestic secondary antibody response) if they detect the antigen that had activated their parent B cell (memory B cells and their parent B cells share the same BCR, thus they detect the same antigen).<sup id="cite_ref-:12_26-2" class="reference"><a href="#cite_note-:12-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Memory B cells can be generated from T cell-dependent activation through both the extrafollicular response and the germinal center reaction as well as from T cell-independent activation of B1 cells.<sup id="cite_ref-:12_26-3" class="reference"><a href="#cite_note-:12-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup></dd> <dt>B-2 cell</dt> <dd>FO B cells and MZ B cells.<sup id="cite_ref-:14_29-0" class="reference"><a href="#cite_note-:14-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> <dl><dt><a href="/wiki/Follicular_B_cells" class="mw-redirect" title="Follicular B cells">Follicular (FO) B cell</a> (also known as a B-2 cell)</dt> <dd>Most common type of B cell and, when not circulating through the blood, is found mainly in the lymphoid follicles of secondary lymphoid organs (SLOs).<sup id="cite_ref-:8_16-8" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> They are responsible for generating the majority of high-affinity antibodies during an infection.<sup id="cite_ref-:0_1-18" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></dd> <dt><a href="/wiki/Marginal-zone_B_cell" title="Marginal-zone B cell">Marginal-zone (MZ) B cell</a></dt> <dd>Found mainly in the marginal zone of the spleen and serves as a first line of defense against blood-borne pathogens, as the marginal zone receives large amounts of blood from the general circulation.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> They can undergo both T cell-independent and T cell-dependent activation, but preferentially undergo T cell-independent activation.<sup id="cite_ref-:8_16-9" class="reference"><a href="#cite_note-:8-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></dd></dl></dd></dl> <dl><dt><a href="/wiki/B-1_cell" class="mw-redirect" title="B-1 cell">B-1 cell</a></dt> <dd>Arises from a developmental pathway different from FO B cells and MZ B cells.<sup id="cite_ref-:14_29-1" class="reference"><a href="#cite_note-:14-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In mice, they predominantly populate the <a href="/wiki/Peritoneal_cavity" title="Peritoneal cavity">peritoneal cavity</a> and <a href="/wiki/Pleural_cavity" title="Pleural cavity">pleural cavity</a>, generate <a href="/wiki/Antibody#Natural_antibodies" title="Antibody">natural antibodies</a> (antibodies produced without infection), defend against mucosal pathogens, and primarily exhibit T cell-independent activation.<sup id="cite_ref-:14_29-2" class="reference"><a href="#cite_note-:14-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> A true homologue of mouse B-1 cells has not been discovered in humans, though various cell populations similar to B-1 cells have been described.<sup id="cite_ref-:14_29-3" class="reference"><a href="#cite_note-:14-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup></dd> <dt><a href="/wiki/Regulatory_B_cells" class="mw-redirect" title="Regulatory B cells">Regulatory B (Breg) cell</a></dt> <dd>An <a href="/wiki/Immunosuppression" title="Immunosuppression">immunosuppressive</a> B cell type that stops the expansion of pathogenic, pro-inflammatory lymphocytes through the secretion of IL-10, IL-35, and TGF-β.<sup id="cite_ref-:15_31-0" class="reference"><a href="#cite_note-:15-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> Also, it promotes the generation of <a href="/wiki/Regulatory_T_cell" title="Regulatory T cell">regulatory T (Treg) cells</a> by directly interacting with T cells to skew their differentiation towards Tregs.<sup id="cite_ref-:15_31-1" class="reference"><a href="#cite_note-:15-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> No common Breg cell identity has been described and many Breg cell subsets sharing regulatory functions have been found in both mice and humans.<sup id="cite_ref-:15_31-2" class="reference"><a href="#cite_note-:15-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> It is currently unknown if Breg cell subsets are developmentally linked and how exactly differentiation into a Breg cell occurs.<sup id="cite_ref-:15_31-3" class="reference"><a href="#cite_note-:15-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> There is evidence showing that nearly all B cell types can differentiate into a Breg cell through mechanisms involving inflammatory signals and BCR recognition.<sup id="cite_ref-:15_31-4" class="reference"><a href="#cite_note-:15-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup></dd></dl> <div class="mw-heading mw-heading2"><h2 id="B_cell-related_pathology">B cell-related pathology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=7" title="Edit section: B cell-related pathology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Autoimmune disease can result from abnormal B cell recognition of self-antigens followed by the production of autoantibodies.<sup id="cite_ref-:16_32-0" class="reference"><a href="#cite_note-:16-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Autoimmune diseases where disease activity is correlated with B cell activity include <a href="/wiki/Scleroderma" title="Scleroderma">scleroderma</a>, <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a>, <a href="/wiki/Systemic_lupus_erythematosus" class="mw-redirect" title="Systemic lupus erythematosus">systemic lupus erythematosus</a>, <a href="/wiki/Diabetes_mellitus_type_1" class="mw-redirect" title="Diabetes mellitus type 1">type 1 diabetes</a>, <a href="/wiki/Irritable_bowel_syndrome" title="Irritable bowel syndrome">post-infectious IBS</a>, and <a href="/wiki/Rheumatoid_arthritis" title="Rheumatoid arthritis">rheumatoid arthritis</a>.<sup id="cite_ref-:16_32-1" class="reference"><a href="#cite_note-:16-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Malignant_transformation" title="Malignant transformation">Malignant transformation</a> of B cells and their precursors can cause a host of <a href="/wiki/Cancer" title="Cancer">cancers</a>, including <a href="/wiki/B-cell_chronic_lymphocytic_leukemia" class="mw-redirect" title="B-cell chronic lymphocytic leukemia">chronic lymphocytic leukemia (CLL)</a>, <a href="/wiki/Acute_lymphoblastic_leukemia" title="Acute lymphoblastic leukemia">acute lymphoblastic leukemia (ALL)</a>, <a href="/wiki/Hairy_cell_leukemia" title="Hairy cell leukemia">hairy cell leukemia</a>, <a href="/wiki/Follicular_lymphoma" title="Follicular lymphoma">follicular lymphoma</a>, <a href="/wiki/Non-Hodgkin_lymphoma" title="Non-Hodgkin lymphoma">non-Hodgkin's lymphoma</a>, <a href="/wiki/Hodgkin%27s_lymphoma" class="mw-redirect" title="Hodgkin's lymphoma">Hodgkin's lymphoma</a>, and <a href="/wiki/Plasma_cell_dyscrasia" class="mw-redirect" title="Plasma cell dyscrasia">plasma cell malignancies</a> such as <a href="/wiki/Multiple_myeloma" title="Multiple myeloma">multiple myeloma</a>, <a href="/wiki/Waldenstr%C3%B6m%27s_macroglobulinemia" class="mw-redirect" title="Waldenström's macroglobulinemia">Waldenström's macroglobulinemia</a>, and certain forms of <a href="/wiki/Amyloidosis" title="Amyloidosis">amyloidosis</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><sup id="cite_ref-pmid27866585_34-0" class="reference"><a href="#cite_note-pmid27866585-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>Abnormal B cells may be relatively large and some diseases include this in their names, such as <a href="/wiki/Diffuse_large_B-cell_lymphoma" title="Diffuse large B-cell lymphoma">diffuse large B-cell lymphomas</a> (DLBCLs) and <a href="/wiki/Intravascular_large_B-cell_lymphoma" class="mw-redirect" title="Intravascular large B-cell lymphoma">intravascular large B-cell lymphoma</a>. </p><p>Patients with B cell alymphocytosis are predisposed to infections.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Epigenetics">Epigenetics</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=B_cell&action=edit&section=8" title="Edit section: Epigenetics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A study that investigated the <a href="/wiki/Methylome" class="mw-redirect" title="Methylome">methylome</a> of B cells along their differentiation cycle, using <a href="/wiki/Whole_genome_bisulfite_sequencing" title="Whole genome bisulfite sequencing">whole-genome bisulfite sequencing</a> (WGBS), showed that there is a hypomethylation from the earliest stages to the most differentiated stages. The largest methylation difference is between the stages of germinal center B cells and memory B cells. Furthermore, this study showed that there is a similarity between B cell tumors and long-lived B cells in their <a href="/wiki/DNA_methylation" title="DNA methylation">DNA methylation</a> signatures.<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> <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=B_cell&action=edit&section=9" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/A20_cells" title="A20 cells">A20 cells</a></li> <li><a href="/wiki/List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">List of distinct cell types in the adult human body</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=B_cell&action=edit&section=10" 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 reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-:0-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:0_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:0_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-:0_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-:0_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-:0_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-:0_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-:0_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-:0_1-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-:0_1-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-:0_1-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-:0_1-12"><sup><i><b>m</b></i></sup></a> <a href="#cite_ref-:0_1-13"><sup><i><b>n</b></i></sup></a> <a href="#cite_ref-:0_1-14"><sup><i><b>o</b></i></sup></a> <a href="#cite_ref-:0_1-15"><sup><i><b>p</b></i></sup></a> <a href="#cite_ref-:0_1-16"><sup><i><b>q</b></i></sup></a> <a href="#cite_ref-:0_1-17"><sup><i><b>r</b></i></sup></a> <a href="#cite_ref-:0_1-18"><sup><i><b>s</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 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href="/wiki/Transitional_B_cell" title="Transitional B cell">Transitional B cell</a></li> <li><a href="/w/index.php?title=Lymphoplasmacytoid_cell&action=edit&redlink=1" class="new" title="Lymphoplasmacytoid cell (page does not exist)">Lymphoplasmacytoid cell</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/T_cell" title="T cell">T cells</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><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Thymocyte" title="Thymocyte">Thymocyte</a></li> <li>αβ (<a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">Cytotoxic CD8<sup>+</sup></a></li> <li><a href="/wiki/T_helper_cell" title="T helper cell">Helper CD4<sup>+</sup></a> / <a href="/wiki/Follicular_B_helper_T_cells" title="Follicular B helper T cells">T<sub>FH</sub></a> / <a href="/wiki/T_helper_3_cell" title="T helper 3 cell">T<sub>h</sub>3</a> / <a href="/wiki/T_helper_17_cell" title="T helper 17 cell">T<sub>h</sub>17</a> / <a href="/wiki/Regulatory_T_cell" title="Regulatory T cell">Regulatory</a>)</li> <li><a href="/wiki/Naive_T_cell" title="Naive T cell">Naïve</a></li> <li><a href="/wiki/Memory_T_cell" title="Memory T cell">Memory T cell</a> <ul><li><a href="/wiki/Central_memory_T_cell" class="mw-redirect" title="Central memory T cell">T<sub>CM</sub></a></li> <li><a href="/wiki/Effector_memory_T_cell" class="mw-redirect" title="Effector memory T cell">T<sub>EM</sub></a></li> <li><a href="/wiki/Tissue-resident_memory_T_cell" title="Tissue-resident memory T cell">T<sub>RM</sub></a></li> <li><a href="/wiki/Virtual_memory_T_cell" title="Virtual memory T cell">T<sub>VM</sub></a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Innate-like_T_cell" class="mw-redirect" title="Innate-like T cell">Innate-like T cells</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/Natural_killer_T_cell" title="Natural killer T cell">Natural killer T cell</a></li> <li><a href="/wiki/Gamma_delta_T_cell" title="Gamma delta T cell">γδ</a></li> <li><a href="/wiki/Mucosal_associated_invariant_T_cell" class="mw-redirect" title="Mucosal associated invariant T cell">Mucosal associated invariant T cell</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Innate_lymphoid_cell" title="Innate lymphoid cell">Innate lymphoid cells</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 id="NK_cells32" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Natural_killer_cell" title="Natural killer cell">NK cells</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/Cytokine-induced_killer_cell" title="Cytokine-induced killer cell">Cytokine-induced killer cell</a></li> <li><a href="/wiki/Lymphokine-activated_killer_cell" title="Lymphokine-activated killer cell">Lymphokine-activated killer cell</a></li> <li><a href="/wiki/Null_cell" title="Null cell">Null cell</a></li> <li><a href="/wiki/Adaptive_NK_cells" class="mw-redirect" title="Adaptive NK cells">Adaptive NK cell</a></li> <li><a href="/wiki/Uterine_natural_killer_cells" title="Uterine natural killer cells">Uterine natural killer cells</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Type_1_innate_lymphoid_cell" class="mw-redirect" title="Type 1 innate lymphoid cell">Type 1 innate lymphoid cells</a></li> <li><a href="/wiki/Type_2_innate_lymphoid_cell" class="mw-redirect" title="Type 2 innate lymphoid cell">Type 2 innate lymphoid cells</a> <ul><li><a href="/wiki/Nuocyte" title="Nuocyte">Nuocytes</a></li></ul></li> <li><a href="/wiki/Type_3_innate_lymphoid_cells" title="Type 3 innate lymphoid cells">Type 3 innate lymphoid cells</a></li> <li><a href="/wiki/LTi_cells" class="mw-redirect" title="LTi cells">LTi cells</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lymphopoiesis" title="Lymphopoiesis">Lymphopoiesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">Hematopoietic stem cell</a></li> <li><a href="/wiki/Lymphoblast" title="Lymphoblast">Lymphoblast</a></li> <li><a href="/wiki/Prolymphocyte" title="Prolymphocyte">Prolymphocyte</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="Lymphocytic_adaptive_immune_system_and_complement123" 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"><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:Lymphocytic_immune_system" title="Template:Lymphocytic immune system"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Lymphocytic_immune_system" title="Template talk:Lymphocytic immune system"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Lymphocytic_immune_system" title="Special:EditPage/Template:Lymphocytic immune system"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Lymphocytic_adaptive_immune_system_and_complement123" style="font-size:114%;margin:0 4em"><a href="/wiki/Lymphocyte" title="Lymphocyte">Lymphocytic</a> <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a> and <a href="/wiki/Complement_system" title="Complement system">complement</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lymphoid</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%">Antigens</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/Antigen" title="Antigen">Antigen</a> <ul><li><a href="/wiki/Superantigen" title="Superantigen">Superantigen</a></li> <li><a href="/wiki/Allergen" title="Allergen">Allergen</a></li> <li><a href="/wiki/Antigenic_variation" title="Antigenic variation">Antigenic variation</a></li></ul></li> <li><a href="/wiki/Hapten" title="Hapten">Hapten</a></li></ul> <ul><li><a href="/wiki/Epitope" title="Epitope">Epitope</a> <ul><li><a href="/wiki/Linear_epitope" title="Linear epitope">Linear</a></li> <li><a href="/wiki/Conformational_epitope" title="Conformational epitope">Conformational</a></li></ul></li> <li><a href="/wiki/Mimotope" title="Mimotope">Mimotope</a></li></ul> <ul><li><a href="/wiki/Antigen_presentation" title="Antigen presentation">Antigen presentation</a>/<a href="/wiki/Antigen-presenting_cell" title="Antigen-presenting cell">professional APCs</a>: <a href="/wiki/Dendritic_cell" title="Dendritic cell">Dendritic cell</a></li> <li><a href="/wiki/Macrophage" title="Macrophage">Macrophage</a></li> <li><a class="mw-selflink selflink">B cell</a></li> <li><a href="/wiki/Immunogen" title="Immunogen">Immunogen</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Antibodies</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/Antibody" title="Antibody">Antibody</a> <ul><li><a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">Monoclonal antibodies</a></li> <li><a href="/wiki/Polyclonal_antibodies" title="Polyclonal antibodies">Polyclonal antibodies</a></li> <li><a href="/wiki/Autoantibody" title="Autoantibody">Autoantibody</a></li> <li><a href="/wiki/Microantibody" title="Microantibody">Microantibody</a></li></ul></li> <li><a href="/wiki/Polyclonal_B_cell_response" title="Polyclonal B cell response">Polyclonal B cell response</a></li> <li><a href="/wiki/Allotype_(immunology)" title="Allotype (immunology)">Allotype</a></li> <li><a href="/wiki/Isotype_(immunology)" title="Isotype (immunology)">Isotype</a></li> <li><a href="/wiki/Idiotype" title="Idiotype">Idiotype</a></li></ul> <ul><li><a href="/wiki/Immune_complex" title="Immune complex">Immune complex</a></li> <li><a href="/wiki/Paratope" title="Paratope">Paratope</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Immunity vs.<br /> tolerance</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Action: <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">Immunity</a></li> <li><a href="/wiki/Autoimmunity" title="Autoimmunity">Autoimmunity</a></li> <li><a href="/wiki/Alloimmunity" title="Alloimmunity">Alloimmunity</a></li> <li><a href="/wiki/Allergy" title="Allergy">Allergy</a></li> <li><a href="/wiki/Hypersensitivity" title="Hypersensitivity">Hypersensitivity</a></li> <li><a href="/wiki/Inflammation" title="Inflammation">Inflammation</a></li> <li><a href="/wiki/Cross-reactivity" title="Cross-reactivity">Cross-reactivity</a></li> <li><a href="/wiki/Co-stimulation" title="Co-stimulation">Co-stimulation</a></li></ul> <ul><li>Inaction: <a href="/wiki/Immune_tolerance" title="Immune tolerance">Tolerance</a> <ul><li><a href="/wiki/Central_tolerance" title="Central tolerance">Central</a></li> <li><a href="/wiki/Peripheral_tolerance" title="Peripheral tolerance">Peripheral</a></li> <li><a href="/wiki/Clonal_anergy" title="Clonal anergy">Clonal anergy</a></li> <li><a href="/wiki/Clonal_deletion" title="Clonal deletion">Clonal deletion</a></li> <li><a href="/wiki/Immune_tolerance_in_pregnancy" title="Immune tolerance in pregnancy">Tolerance in pregnancy</a></li></ul></li> <li><a href="/wiki/Immunodeficiency" title="Immunodeficiency">Immunodeficiency</a></li> <li><a href="/wiki/Immune_privilege" title="Immune privilege">Immune privilege</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</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/Affinity_maturation" title="Affinity maturation">Affinity maturation</a> <ul><li><a href="/wiki/Somatic_hypermutation" title="Somatic hypermutation">Somatic hypermutation</a></li> <li><a href="/wiki/Clonal_selection" title="Clonal selection">Clonal selection</a></li></ul></li> <li><a href="/wiki/V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a></li> <li><a href="/wiki/Junctional_diversity" title="Junctional diversity">Junctional diversity</a></li> <li><a href="/wiki/Immunoglobulin_class_switching" title="Immunoglobulin class switching">Immunoglobulin class switching</a></li> <li><a href="/wiki/Major_histocompatibility_complex" title="Major histocompatibility complex">MHC</a>/<a href="/wiki/Human_leukocyte_antigen" title="Human leukocyte antigen">HLA</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lymphocyte" title="Lymphocyte">Lymphocytes</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/Cell-mediated_immunity" title="Cell-mediated immunity">Cellular</a> <ul><li><a href="/wiki/T_cell" title="T cell">T cell</a></li></ul></li> <li><a href="/wiki/Humoral_immunity" title="Humoral immunity">Humoral</a> <ul><li><a class="mw-selflink selflink">B cell</a></li></ul></li> <li><a href="/wiki/Natural_killer_cell" title="Natural killer cell">NK cell</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Substances</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/Cytokine" title="Cytokine">Cytokines</a></li> <li><a href="/wiki/Opsonin" title="Opsonin">Opsonin</a></li> <li><a href="/wiki/Cytolysin" title="Cytolysin">Cytolysin</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" /><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319" 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