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id="toc-Invention_of_TNF_inhibitors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Discovery_of_the_LPS_receptor,_and_the_role_of_TLRs_in_innate_immune_sensing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Discovery_of_the_LPS_receptor,_and_the_role_of_TLRs_in_innate_immune_sensing"> <div class="vector-toc-text"> 2.3 Discovery of the LPS receptor, and the role of TLRs in innate immune sensing </div> </a> <ul id="toc-Discovery_of_the_LPS_receptor,_and_the_role_of_TLRs_in_innate_immune_sensing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Random_Germline_Mutagenesis/Forward_Genetics_in_the_mouse" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Random_Germline_Mutagenesis/Forward_Genetics_in_the_mouse"> <div class="vector-toc-text"> 2.4 Random Germline Mutagenesis/Forward Genetics in the mouse </div> </a> <ul id="toc-Random_Germline_Mutagenesis/Forward_Genetics_in_the_mouse-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Invention_of_Automated_Meiotic_Mapping" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Invention_of_Automated_Meiotic_Mapping"> <div class="vector-toc-text"> 2.5 Invention of Automated Meiotic Mapping </div> </a> <ul id="toc-Invention_of_Automated_Meiotic_Mapping-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Developing_drugs_that_activate_TLRs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Developing_drugs_that_activate_TLRs"> <div class="vector-toc-text"> 2.6 Developing drugs that activate TLRs </div> </a> <ul id="toc-Developing_drugs_that_activate_TLRs-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Awards_and_recognition" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Awards_and_recognition"> <div class="vector-toc-text"> 3 Awards and recognition </div> </a> <button aria-controls="toc-Awards_and_recognition-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Awards and recognition subsection </button> <ul id="toc-Awards_and_recognition-sublist" class="vector-toc-list"> <li id="toc-Awards" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Awards"> <div class="vector-toc-text"> 3.1 Awards </div> </a> <ul id="toc-Awards-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Honorary_Doctoral_Degrees" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Honorary_Doctoral_Degrees"> <div class="vector-toc-text"> 3.2 Honorary Doctoral Degrees </div> </a> <ul id="toc-Honorary_Doctoral_Degrees-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Family" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Family"> <div class="vector-toc-text"> 4 Family </div> </a> <ul id="toc-Family-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"> 5 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 6 References </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 7 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links_2" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links_2"> <div class="vector-toc-text"> 8 External links </div> </a> <ul id="toc-External_links_2-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" > Toggle the table of contents </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Bruce Beutler</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" > 55 languages </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%B1%D9%88%D8%B3_%D8%A8%D9%88%D8%AA%D9%84%D8%B1" title="بروس بوتلر – Arabic" lang="ar" hreflang="ar" data-title="بروس بوتلر" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Br%C3%BCs_B%C3%B6tler" title="Brüs Bötler – Azerbaijani" lang="az" hreflang="az" data-title="Brüs Bötler" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D8%B3_%D8%A8%D9%88%DB%8C%D8%AA%D9%84%D8%B1" title="بروس بویتلر – South Azerbaijani" lang="azb" hreflang="azb" data-title="بروس بویتلر" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target">تۆرکجه</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AC%E0%A7%8D%E0%A6%B0%E0%A7%81%E0%A6%B8_%E0%A6%AC%E0%A6%AF%E0%A6%BC%E0%A7%87%E0%A6%9F%E0%A6%B2%E0%A6%BE%E0%A6%B0" title="ব্রুস বয়েটলার – Bangla" lang="bn" hreflang="bn" data-title="ব্রুস বয়েটলার" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%91%D1%80%D1%83%D1%81_%D0%91%D1%91%D1%82%D0%BB%D0%B5%D1%80" title="Брус Бётлер – Belarusian" lang="be" hreflang="be" data-title="Брус Бётлер" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%91%D1%80%D1%83%D1%81_%D0%91%D0%BE%D0%B9%D1%82%D0%BB%D0%B5%D1%80" title="Брус Бойтлер – Bulgarian" lang="bg" hreflang="bg" data-title="Брус Бойтлер" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Catalan" lang="ca" hreflang="ca" data-title="Bruce Beutler" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Czech" lang="cs" hreflang="cs" data-title="Bruce Beutler" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Bruce_A._Beutler" title="Bruce A. Beutler – Danish" lang="da" hreflang="da" data-title="Bruce A. Beutler" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – German" lang="de" hreflang="de" data-title="Bruce Beutler" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Estonian" lang="et" hreflang="et" data-title="Bruce Beutler" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Spanish" lang="es" hreflang="es" data-title="Bruce Beutler" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Esperanto" lang="eo" hreflang="eo" data-title="Bruce Beutler" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Basque" lang="eu" hreflang="eu" data-title="Bruce Beutler" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D8%B3_%D8%A8%D9%88%DB%8C%D8%AA%D9%84%D8%B1" title="بروس بویتلر – Persian" lang="fa" hreflang="fa" data-title="بروس بویتلر" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – French" lang="fr" hreflang="fr" data-title="Bruce Beutler" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%B8%8C%EB%A3%A8%EC%8A%A4_%EB%B3%B4%EC%9D%B4%ED%8B%80%EB%9F%AC" title="브루스 보이틀러 – Korean" lang="ko" hreflang="ko" data-title="브루스 보이틀러" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D6%80%D5%B5%D5%B8%D6%82%D5%BD_%D4%B2%D5%B8%D5%B5%D5%A9%D5%AC%D5%A5%D6%80" title="Բրյուս Բոյթլեր – Armenian" lang="hy" hreflang="hy" data-title="Բրյուս Բոյթլեր" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Croatian" lang="hr" hreflang="hr" data-title="Bruce Beutler" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Bruce_Alan_Beutler" title="Bruce Alan Beutler – Ido" lang="io" hreflang="io" data-title="Bruce Alan Beutler" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target">Ido</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Indonesian" lang="id" hreflang="id" data-title="Bruce Beutler" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Italian" lang="it" hreflang="it" data-title="Bruce Beutler" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%91%D7%A8%D7%95%D7%A1_%D7%91%D7%95%D7%99%D7%98%D7%9C%D7%A8" title="ברוס בויטלר – Hebrew" lang="he" hreflang="he" data-title="ברוס בויטלר" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%91%D1%80%D1%8E%D1%81_%D0%91%D0%B5%D1%82%D0%BB%D0%B5%D1%80" title="Брюс Бетлер – Kazakh" lang="kk" hreflang="kk" data-title="Брюс Бетлер" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Swahili" lang="sw" hreflang="sw" data-title="Bruce Beutler" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target">Kiswahili</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Brussius_Beutler" title="Brussius Beutler – Latin" lang="la" hreflang="la" data-title="Brussius Beutler" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Br%C5%ABss_Betlers" title="Brūss Betlers – Latvian" lang="lv" hreflang="lv" data-title="Brūss Betlers" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Hungarian" lang="hu" hreflang="hu" data-title="Bruce Beutler" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D8%B3_%D8%A8%D9%88%D8%AA%D9%84%D8%B1" title="بروس بوتلر – Egyptian Arabic" lang="arz" hreflang="arz" data-title="بروس بوتلر" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target">مصرى</a></li><li class="interlanguage-link interwiki-mzn mw-list-item"><a href="https://mzn.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D8%B3_%D8%A8%D9%88%DB%8C%D8%AA%D9%84%D8%B1" title="بروس بویتلر – Mazanderani" lang="mzn" hreflang="mzn" data-title="بروس بویتلر" data-language-autonym="مازِرونی" data-language-local-name="Mazanderani" class="interlanguage-link-target">مازِرونی</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Malay" lang="ms" hreflang="ms" data-title="Bruce Beutler" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Dutch" lang="nl" hreflang="nl" data-title="Bruce Beutler" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%96%E3%83%AB%E3%83%BC%E3%82%B9%E3%83%BB%E3%83%9C%E3%82%A4%E3%83%88%E3%83%A9%E3%83%BC" title="ブルース・ボイトラー – Japanese" lang="ja" hreflang="ja" data-title="ブルース・ボイトラー" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Bruce Beutler" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Bruce_A._Beutler" title="Bruce A. Beutler – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Bruce A. Beutler" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D8%B3_%D8%A8%D9%B9%D9%84%D8%B1" title="بروس بٹلر – Western Punjabi" lang="pnb" hreflang="pnb" data-title="بروس بٹلر" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target">پنجابی</a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Bruce_Alan_Beutler" title="Bruce Alan Beutler – Low German" lang="nds" hreflang="nds" data-title="Bruce Alan Beutler" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target">Plattdüütsch</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Polish" lang="pl" hreflang="pl" data-title="Bruce Beutler" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Portuguese" lang="pt" hreflang="pt" data-title="Bruce Beutler" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Romanian" lang="ro" hreflang="ro" data-title="Bruce Beutler" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%91%D1%91%D1%82%D0%BB%D0%B5%D1%80,_%D0%91%D1%80%D1%8E%D1%81" title="Бётлер, Брюс – Russian" lang="ru" hreflang="ru" data-title="Бётлер, Брюс" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Bruce_Beutler" title="Bruce Beutler – Simple English" lang="en-simple" hreflang="en-simple" data-title="Bruce Beutler" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%A8%D8%B1%D9%88%D9%88%D8%B3_%D8%A8%DB%86%DB%8C%D8%AA%D9%84%DB%95%D8%B1" title="برووس بۆیتلەر – Central Kurdish" lang="ckb" hreflang="ckb" data-title="برووس بۆیتلەر" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a 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divorced; 3 children)</td></tr><tr><th scope="row" class="infobox-label">Awards</th><td class="infobox-data">2011 <a href="/wiki/Nobel_Prize" title="Nobel Prize">Nobel Prize</a> in <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Physiology or Medicine</a></td></tr><tr><td colspan="2" class="infobox-full-data"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546">Scientific career</td></tr><tr><th scope="row" class="infobox-label">Fields</th><td class="infobox-data category"><a href="/wiki/Immunology" title="Immunology">Immunology</a></td></tr><tr><th scope="row" class="infobox-label">Institutions</th><td class="infobox-data"><a href="/wiki/University_of_Texas_Southwestern_Medical_Center" title="University of Texas Southwestern Medical Center">University of Texas Southwestern Medical Center</a></td></tr><tr style="display:none"><td colspan="2"> </td></tr></tbody></table> Bruce Alan Beutler (<a href="/wiki/Help:IPA/English" title="Help:IPA/English">/ˈbɔɪtlər/</a> <a href="/wiki/Help:Pronunciation_respelling_key" title="Help:Pronunciation respelling key">BOYT-lər</a>; born December 29, 1957) is an American immunologist and geneticist. Together with <a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules A. Hoffmann</a>, he received one-half of the 2011 <a href="/wiki/Nobel_Prize" title="Nobel Prize">Nobel Prize</a> in <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Physiology or Medicine</a>, for "discoveries concerning the activation of innate immunity."<a href="#cite_note-Nobel-1">[1]</a> Beutler discovered the long-elusive receptor for <a href="/wiki/Lipopolysaccharide" title="Lipopolysaccharide">lipopolysaccharide</a> (LPS; also known as endotoxin). He did so by identifying spontaneous mutations in the gene coding for mouse <a href="/wiki/Toll-like_receptor_4" title="Toll-like receptor 4">Toll-like receptor 4</a> (Tlr4) in two unrelated strains of LPS-refractory mice and proving they were responsible for that phenotype.<a href="#cite_note-:1-2">[2]</a> Subsequently, and chiefly through the work of <a href="/wiki/Shizuo_Akira" title="Shizuo Akira">Shizuo Akira</a>, other TLRs were shown to detect signature molecules of most infectious microbes, in each case triggering an <a href="/wiki/Innate_immune_system" title="Innate immune system">innate immune response</a>.<a href="#cite_note-3">[3]</a><a href="#cite_note-4">[4]</a><a href="#cite_note-5">[5]</a><a href="#cite_note-6">[6]</a><a href="#cite_note-7">[7]</a> The other half of the Nobel Prize went to <a href="/wiki/Ralph_M._Steinman" title="Ralph M. Steinman">Ralph M. Steinman</a> for "his discovery of the dendritic cell and its role in <a href="/wiki/Adaptive_immunity" class="mw-redirect" title="Adaptive immunity">adaptive immunity</a>."<a href="#cite_note-Nobel-1">[1]</a> Beutler is currently a Regental Professor and Director of the Center for the Genetics of Host Defense at the <a href="/wiki/University_of_Texas_Southwestern_Medical_Center" title="University of Texas Southwestern Medical Center">University of Texas Southwestern Medical Center</a> in <a href="/wiki/Dallas" title="Dallas">Dallas</a>, <a href="/wiki/Texas" title="Texas">Texas</a>.<a href="#cite_note-profile-8">[8]</a><a href="#cite_note-9">[9]</a> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Early_life_and_education">Early life and education</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=1" title="Edit section: Early life and education">edit</a>]</div> Born in <a href="/wiki/Chicago" title="Chicago">Chicago</a>, Illinois, to a <a href="/wiki/Jews" title="Jews">Jewish</a><a href="#cite_note-10">[10]</a> family, Beutler lived in Southern California between the ages of 2 and 18 (1959 to 1977). For most of this time, he lived in city of <a href="/wiki/Arcadia,_California" title="Arcadia, California">Arcadia</a>, a northeastern suburb of <a href="/wiki/Los_Angeles" title="Los Angeles">Los Angeles</a> in the <a href="/wiki/San_Gabriel_Valley" title="San Gabriel Valley">San Gabriel Valley</a>. During these years, he spent much time hiking in the San Gabriel Mountains, and in regional national parks (<a href="/wiki/Sequoia_National_Park" title="Sequoia National Park">Sequoia</a>, <a href="/wiki/Yosemite_National_Park" title="Yosemite National Park">Yosemite</a>, <a href="/wiki/Joshua_Tree_National_Park" title="Joshua Tree National Park">Joshua Tree</a>, and <a href="/wiki/Grand_Canyon_National_Park" title="Grand Canyon National Park">Grand Canyon</a>), and was particularly fascinated by living things.<a href="#cite_note-:0-11">[11]</a> These experiences impelled an intense interest in biological science. His introduction to experimental biology, acquired between the ages of 14 and 18, included work in the laboratory of his father, <a href="/wiki/Ernest_Beutler" title="Ernest Beutler">Ernest Beutler</a>, then at the <a href="/wiki/City_of_Hope_National_Medical_Center" title="City of Hope National Medical Center">City of Hope Medical Center</a> in <a href="/wiki/Duarte,_California" title="Duarte, California">Duarte</a>, CA. There he learned to assay enzymes of red blood cells and became familiar with methods for protein isolation. He published his studies of an electrophoretic variant of glutathione peroxidase,<a href="#cite_note-:32-12">[12]</a> as well as the inherent catalytic activity of inorganic selenite,<a href="#cite_note-:33-13">[13]</a> at the age of 17. Beutler also worked in the City of Hope laboratory of <a href="/wiki/Susumu_Ohno" title="Susumu Ohno">Susumu Ohno</a>, a geneticist known for his studies of evolution, genome structure, and sex differentiation in mammals. Ohno hypothesized that the major histocompatibility complex proteins served as anchorage sites for organogenesis-directing proteins.<a href="#cite_note-14">[14]</a> He further suggested that <a href="/wiki/H-Y_antigen" title="H-Y antigen">H-Y antigen</a>, a minor histocompatibility protein encoded by a gene on the Y chromosome and absent in female mammals, was responsible for directing organogenesis of the indifferent gonad to form a testis. In studying H-Y antigen,<a href="#cite_note-15">[15]</a> Beutler became conversant with immunology and mouse genetics during the 1970s. While a college student at the <a href="/wiki/University_of_California,_San_Diego" title="University of California, San Diego">University of California at San Diego</a>, Beutler worked in the laboratory of <a href="/wiki/Dan_Lindsley" title="Dan Lindsley">Dan Lindsley</a>, a <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila</a> geneticist interested in spermatogenesis and spermiogenesis in the fruit fly. There, he learned to map phenotypes to chromosomal regions using visible phenotypic markers.<a href="#cite_note-:0-11">[11]</a> He also worked in the laboratory of Abraham Braude, an expert in the biology of LPS. Beutler received his secondary school education at <a href="/wiki/Polytechnic_School_(California)" title="Polytechnic School (California)">Polytechnic School</a> in <a href="/wiki/Pasadena,_California" title="Pasadena, California">Pasadena</a>, California. A precocious student, he graduated from high school at the age of 16, enrolled in college at the University of California, San Diego, and graduated with a BA degree at the age of 18 in 1976. He then enrolled in medical school at the <a href="/wiki/University_of_Chicago" title="University of Chicago">University of Chicago</a> in 1977 and received his M.D. degree in 1981 at the age of 23.<a href="#cite_note-16">[16]</a> From 1981 to 1983 Beutler continued his medical training at the University of Texas Southwestern Medical Center in Dallas, Texas, as an intern in the Department of Internal Medicine, and as a resident in the Department of Neurology. However, he found clinical medicine less interesting than laboratory science, and decided to return to the laboratory. <div class="mw-heading mw-heading2"><h2 id="Scientific_contributions">Scientific contributions</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=2" title="Edit section: Scientific contributions">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Isolation_of_tumor_necrosis_factor_and_discovery_of_its_inflammation-promoting_effect">Isolation of tumor necrosis factor and discovery of its inflammation-promoting effect</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=3" title="Edit section: Isolation of tumor necrosis factor and discovery of its inflammation-promoting effect">edit</a>]</div> Beutler’s focus on innate immunity began when he was a postdoctoral associate and later an assistant professor in the lab of <a href="/wiki/Anthony_Cerami" title="Anthony Cerami">Anthony Cerami</a> at <a href="/wiki/Rockefeller_University" title="Rockefeller University">Rockefeller University</a> (1983-1986). Drawing upon skills he had acquired earlier, he isolated mouse “cachectin” from the conditioned medium of LPS-activated mouse macrophages.<a href="#cite_note-17">[17]</a> Cachectin was hypothesized by Cerami to be a mediator of wasting in chronic disease. Its biological activity, the suppression of lipoprotein lipase synthesis in adipocytes, was thought to contribute to wasting, since lipoprotein lipase cleaves fatty acids from circulating triglycerides, allowing their uptake and re-esterification within fat cells.<a href="#cite_note-:2-18">[18]</a> By sequential fractionation of LPS-activated macrophage medium, measuring cachectin activity at each step, Beutler purified cachectin to homogeneity.<a href="#cite_note-:3-19">[19]</a> Determining its N-terminal sequence, he recognized it as mouse <a href="/wiki/Tumor_necrosis_factor" title="Tumor necrosis factor">tumor necrosis factor</a> (TNF), and showed that it had strong TNF activity; moreover that human TNF, isolated by a very different assay, had strong cachectin activity.<a href="#cite_note-:2-18">[18]</a> Human TNF, isolated contemporaneously by other workers,<a href="#cite_note-20">[20]</a> had to that time been defined only by its ability to kill cancer cells. The discovery of a separate role for TNF as a catabolic switch was of considerable interest. Of still greater importance, Beutler demonstrated that TNF acted as a key mediator of  endotoxin-induced shock.<a href="#cite_note-:4-21">[21]</a> This he accomplished by raising an antibody against mouse TNF, which he used to neutralize TNF in living mice challenged with lipopolysaccharide (LPS).<a href="#cite_note-:4-21">[21]</a> The often-lethal systemic inflammatory response to LPS was significantly mitigated by passive <a href="/wiki/Immunization" title="Immunization">immunization</a> against TNF. The discovery that TNF caused an acute systemic inflammatory disease (LPS-induced shock) presaged its causative role in numerous chronic inflammatory diseases. With J.-M. Dayer, Beutler demonstrated that purified TNF could cause inflammation-associated responses in cultured human synoviocytes: secretion of <a href="/wiki/Collagenase" title="Collagenase">collagenase</a> and <a href="/wiki/Prostaglandin_E2" title="Prostaglandin E2">prostaglandin E2</a>.<a href="#cite_note-22">[22]</a>  This was an early hint that TNF might be causally important in <a href="/wiki/Rheumatoid_arthritis" title="Rheumatoid arthritis">rheumatoid arthritis</a> (as later shown by Feldmann, Brennan, and Maini<a href="#cite_note-23">[23]</a>). Beutler also demonstrated the existence of TNF receptors on most cell types,<a href="#cite_note-:3-19">[19]</a> and correctly inferred the presence of two types of TNF receptor distinguished by their affinities, later cloned and designated <a href="/wiki/Tumor_necrosis_factor_receptor_1" title="Tumor necrosis factor receptor 1">p55</a> and <a href="/wiki/Tumor_necrosis_factor_receptor_2" title="Tumor necrosis factor receptor 2">p75</a> TNF receptors to denote their approximate molecular weights.<a href="#cite_note-24">[24]</a><a href="#cite_note-25">[25]</a><a href="#cite_note-26">[26]</a><a href="#cite_note-27">[27]</a><a href="#cite_note-28">[28]</a> Before a sensitive immunoassay for TNF was feasible, Beutler used these receptors in a binding competition assay using radio-iodinated TNF as a tracer, which allowed him to precisely measure TNF in biological fluids.<a href="#cite_note-29">[29]</a> <div class="mw-heading mw-heading3"><h3 id="Invention_of_TNF_inhibitors">Invention of TNF inhibitors</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=4" title="Edit section: Invention of TNF inhibitors">edit</a>]</div> Beutler was recruited to a faculty position at UT Southwestern Medical Center and the Howard Hughes Medical Institute in 1986. Aware that TNF blockade might have clinical applications, he (along with a graduate student, David Crawford, and a postdoctoral associate, Karsten Peppel) invented and patented recombinant molecules expressly designed to neutralize TNF in vivo (Patent No. <a rel="nofollow" class="external text" href="https://patents.google.com/patent/US5447851B1/en">US5447851B1</a>).<a href="#cite_note-:5-30">[30]</a> Fusing the binding portion of TNF receptor proteins to the <a href="/wiki/Immunoglobulin_heavy_chain" title="Immunoglobulin heavy chain">heavy chain</a> of an immunoglobulin molecule to force receptor dimerization,<a href="#cite_note-:5-30">[30]</a> they produced chimeric reagents with surprisingly high affinity and specificity for both TNF and a closely related cytokine called lymphotoxin, low antigenicity, and excellent stability in vivo. The human p75 receptor chimeric protein was later used extensively as the drug <a href="/wiki/Etanercept" title="Etanercept">Etanercept</a> in the treatment of rheumatoid arthritis, <a href="/wiki/Crohn%27s_disease" title="Crohn's disease">Crohn's disease</a>, <a href="/wiki/Psoriasis" title="Psoriasis">psoriasis</a>, and other forms of inflammation. Marketed by <a href="/wiki/Amgen" title="Amgen">Amgen</a>, Etanercept achieved more than $74B in sales.<a href="#cite_note-31">[31]</a> <div class="mw-heading mw-heading3"><h3 id="Discovery_of_the_LPS_receptor,_and_the_role_of_TLRs_in_innate_immune_sensing">Discovery of the LPS receptor, and the role of TLRs in innate immune sensing</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=5" title="Edit section: Discovery of the LPS receptor, and the role of TLRs in innate immune sensing">edit</a>]</div> From the mid-1980s onward Beutler was interested in the mechanism by which LPS activates mammalian immune cells (chiefly <a href="/wiki/Macrophage" title="Macrophage">macrophages</a>,<a href="#cite_note-:2-18">[18]</a><a href="#cite_note-:4-21">[21]</a> but <a href="/wiki/Dendritic_cell" title="Dendritic cell">dendritic cells</a> and <a href="/wiki/B_cell" title="B cell">B cells</a> as well), sometimes leading to uncontrollable Gram negative septic shock,<a href="#cite_note-32">[32]</a><a href="#cite_note-33">[33]</a><a href="#cite_note-34">[34]</a> but also promoting the well-known adjuvant effect of LPS,<a href="#cite_note-35">[35]</a> and B cell mitogenesis<a href="#cite_note-:6-36">[36]</a><a href="#cite_note-37">[37]</a> and antibody production. A single, highly specific LPS receptor was presumed to exist as early as the 1960s, based on the fact that allelic mutations in two separate strains of mice, affecting a discrete genetic locus on chromosome 4 termed Lps, abolished LPS sensing.<a href="#cite_note-:6-36">[36]</a><a href="#cite_note-38">[38]</a> Although this receptor had been widely pursued, it remained elusive. Beutler reasoned that in finding the LPS receptor, insight might be gained into the first molecular events that transpire upon an encounter between the host and microbial invaders.<a href="#cite_note-39">[39]</a> Utilizing <a href="/wiki/Positional_cloning" class="mw-redirect" title="Positional cloning">positional cloning</a> in an effort that began in 1993 and lasted five years, Beutler, together with several postdoctoral associates including Alexander Poltorak, measured TNF production as a qualitative phenotypic endpoint of the LPS response. Analyzing more than 2,000 <a href="/wiki/Meiosis" title="Meiosis">meioses</a>, they confined the LPS receptor-encoding gene to a region of the genome encompassing approximately 5.8 million base pairs of <a href="/wiki/DNA" title="DNA">DNA</a>.<a href="#cite_note-:1-2">[2]</a> Sequencing most of the interval, they identified a gene within which each of two LPS-refractory strains of mice (C3H/HeJ and C57BL/10ScCr) had deleterious mutations. The gene, Tlr4, encoded a cell surface protein with cytoplasmic domain homology to the <a href="/wiki/Interleukin-1_receptor" title="Interleukin-1 receptor">interleukin-1 receptor</a>, and several other homologous genes that were scattered across the mouse genome. Beutler and his team thus proved that one of the mammalian Toll-like receptors, TLR4, acts as the membrane-spanning component of the mammalian LPS receptor complex.<a href="#cite_note-:1-2">[2]</a><a href="#cite_note-40">[40]</a><a href="#cite_note-:7-41">[41]</a> They also showed that while mouse TLR4 is activated by a tetra-acylated LPS-like molecule (lipid IVa), human TLR4 is not, recapitulating the species specificity for LPS partial structures.<a href="#cite_note-:7-41">[41]</a> It was deduced that direct contact between TLR4 and LPS is a prerequisite for cell activation.<a href="#cite_note-:7-41">[41]</a>  Later, an extracellular component of the LPS receptor complex, MD-2 (also known as lymphocyte antigen 96), was identified by R. Shimazu and colleagues.<a href="#cite_note-42">[42]</a> The structure of the complex, with and without LPS bound, was solved by Jie-Oh Lee and colleagues in 2009.<a href="#cite_note-43">[43]</a> Jules Hoffmann and colleagues had earlier shown that the Drosophila Toll protein, originally known for its role in embryogenesis, was essential for the <a href="/wiki/Antimicrobial_peptides" title="Antimicrobial peptides">antimicrobial peptide</a> response to fungal infection.<a href="#cite_note-44">[44]</a>  However, no molecule derived from <a href="/wiki/Fungus" title="Fungus">fungi</a> actually became bound to Toll; rather, a proteolytic cascade led to the activation of an endogenous ligand, the protein <a href="/wiki/Sp%C3%A4tzle_(gene)" title="Spätzle (gene)">Spätzle</a>. This activated <a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-kB</a> within cells of the fat body, leading to antimicrobial peptide secretion. Aware of this work, <a href="/wiki/Charles_Janeway" title="Charles Janeway">Charles Janeway</a> and <a href="/wiki/Ruslan_Medzhitov" title="Ruslan Medzhitov">Ruslan Medzhitov</a> overexpressed a modified version of human TLR4 (which they called ‘h-Toll’) and found it capable of activating the transcription factor NF-κB in mammalian cells.<a href="#cite_note-45">[45]</a> They speculated that TLR4 was a “<a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern recognition receptor</a>.”  However, they provided no evidence that TLR4 recognized any molecule of microbial origin. If a ligand did exist, it might have been endogenous (as in the fruit fly, where Toll recognizes the endogenous protein Spätzle, or as in the case of the IL-1 receptor, which recognizes the endogenous cytokine <a href="/wiki/Interleukin-1_family" title="Interleukin-1 family">IL-1</a>). Indeed, numerous cell surface receptors, including the <a href="/wiki/TGF_beta_receptor" title="TGF beta receptor">TGFβ receptor</a>, <a href="/wiki/B-cell_receptor" title="B-cell receptor">B cell receptor</a>, and <a href="/wiki/T-cell_receptor" title="T-cell receptor">T cell receptor</a> activate NF-κB. In short, it was not clear what TLR4 recognized, nor what its function was. Separate publications, also based on transfection/overexpression studies, held that TLR2 rather than TLR4 was the LPS receptor.<a href="#cite_note-46">[46]</a><a href="#cite_note-47">[47]</a> The genetic evidence of Beutler and coworkers correctly identified TLR4 as the specific and non-redundant cell surface receptor for LPS, fully required for virtually all LPS activities. This suggested that other TLRs (of which ten are now known to exist in humans) might also act as sensors of infection in mammals,<a href="#cite_note-48">[48]</a> each detecting other signature molecules made by microbes whether or not they were <a href="/wiki/Pathogen" title="Pathogen">pathogens</a> in the classical sense of the term. The other TLRs, like TLR4, do indeed initiate innate immune responses. By promoting inflammatory signaling, TLRs can also mediate pathologic effects including <a href="/wiki/Fever" title="Fever">fever</a>, systemic inflammation, and shock. Sterile inflammatory and autoimmune diseases such as <a href="/wiki/Lupus" title="Lupus">systemic lupus erythematosus</a> also elicit TLR signaling, and disruption of signaling from the nucleic acid sensing TLRs can favorably modify the disease phenotype.<a href="#cite_note-49">[49]</a><a href="#cite_note-50">[50]</a><a href="#cite_note-51">[51]</a><a href="#cite_note-:8-52">[52]</a><a href="#cite_note-53">[53]</a><a href="#cite_note-54">[54]</a><a href="#cite_note-55">[55]</a><a href="#cite_note-56">[56]</a> <div class="mw-heading mw-heading3"><h3 id="Random_Germline_Mutagenesis/Forward_Genetics_in_the_mouse">Random Germline Mutagenesis/Forward Genetics in the mouse</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=6" title="Edit section: Random Germline Mutagenesis/Forward Genetics in the mouse">edit</a>]</div> After completing the positional cloning of the Lps locus in 1998, Beutler continued to apply a forward genetic approach to the analysis of immunity in mammals. In this process, germline mutations that alter immune function are created in mice through a random process using the alkylating agent <a href="/wiki/ENU" title="ENU">ENU</a>, detected by their phenotypic effects, and then isolated by positional cloning.<a href="#cite_note-57">[57]</a> This work disclosed numerous essential signaling molecules required for the innate immune response,<a href="#cite_note-:9-58">[58]</a><a href="#cite_note-59">[59]</a><a href="#cite_note-:10-60">[60]</a><a href="#cite_note-61">[61]</a><a href="#cite_note-62">[62]</a><a href="#cite_note-63">[63]</a><a href="#cite_note-64">[64]</a> and helped to delineate the biochemistry of innate immunity. Among the genes detected was Ticam1, implicated by an ENU-induced phenotype called Lps2.<a href="#cite_note-:9-58">[58]</a> The encoded protein <a href="/wiki/TRIF" class="mw-redirect" title="TRIF">TICAM1</a>, also known as TRIF, was a new adaptor molecule, binding to the cytoplasmic domains of both TLR3 and TLR4, and needed for signaling by each. Another phenotype, called 3d to connote a “triple defect” in TLR signaling, affected a gene of unknown function called Unc93b1.<a href="#cite_note-:10-60">[60]</a> TLRs 3, 7, and 9 (nucleic acid sensing TLRs) failed to signal in homozygotes for the mutation. These TLRs were found to be <a href="/wiki/Endosome" title="Endosome">endosomal</a>, and physically interact with the UNC93B1 protein which transports them to the endosomal compartment.<a href="#cite_note-65">[65]</a>  Humans with mutations in UNC93B1, the human ortholog of the same gene, were subsequently found to be susceptible to recurrent <a href="/wiki/Herpes_simplex_virus" title="Herpes simplex virus">Herpes simplex virus</a> (HSV) <a href="/wiki/Encephalitis" title="Encephalitis">encephalitis</a>, in which reactivation of latent virus occurs repeatedly in the <a href="/wiki/Trigeminal_ganglion" title="Trigeminal ganglion">trigeminal ganglion</a> at the base of the <a href="/wiki/Midbrain" title="Midbrain">midbrain</a>, leading to cortical neuron death.<a href="#cite_note-:11-66">[66]</a> Yet another protein needed to make the endosomal environment suitable for TLR signaling was SLC15A4, identified based on the phenotype feeble.<a href="#cite_note-67">[67]</a> feeble was identified in a screen in which immunostimulatory DNA was administered to mice intravenously with measurement of the systemic <a href="/wiki/Interferon_type_I" title="Interferon type I">type I interferon</a> response. Failure of this response, which is dependent on TLR9 signaling from <a href="/wiki/Plasmacytoid_dendritic_cell" title="Plasmacytoid dendritic cell">plasmacytoid dendritic cells</a> (pDC) was observed in homozygous mutants, and subsequently, failure of TLR7 (but not TLR3) signaling was observed as well. Because the feeble mutation suppressed SLE in mice,<a href="#cite_note-:8-52">[52]</a> the SLC15A4 protein has become a target of interest for drug development.<a href="#cite_note-68">[68]</a> In all, Beutler and colleagues detected 77 mutations in 36 genes in which ENU-induced mutations created defects of TLR signaling, detected due to faulty TNF and/or interferon responses. These genes encoded all TLRs kept under surveillance in screening, all of the four adapter proteins that signal from TLRs, kinases and other signaling proteins downstream, <a href="/wiki/Chaperone_(protein)" title="Chaperone (protein)">chaperones</a> needed to escort TLRs to their destinations, proteins that promote the availability of TLR ligands, proteins involved in vesicle transport, and proteins involved in <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcriptional</a> responses to TLR signaling, or the post-translational processing of TNF and/or type I interferons (the proteins assayed in screening). Beutler and colleagues also used ENU mutagenesis to study the global response to a defined infectious agent. They measured susceptibility to mouse <a href="/wiki/Cytomegalovirus" title="Cytomegalovirus">cytomegalovirus</a> (MCMV) and identified numerous genes that make a life-or-death difference during infection, terming this set of genes the MCMV "resistome".<a href="#cite_note-69">[69]</a><a href="#cite_note-70">[70]</a> These genes were grouped into "sensing," "signaling," "effector," "homeostatic," and "developmental" categories, some of which were wholly unexpected. In the homeostatic category, for example, <a href="/wiki/KCNJ8" title="KCNJ8">Kir6.1</a> <a href="/wiki/ATP-sensitive_potassium_channel" title="ATP-sensitive potassium channel">ATP-sensitive potassium channels</a> in the <a href="/wiki/Smooth_muscle" title="Smooth muscle">smooth muscle</a> of the <a href="/wiki/Coronary_circulation" title="Coronary circulation">coronary arteries</a> serve an essential role in the maintenance of blood flow during MCMV infection, and mutations that damage these channels cause sudden death during infection.<a href="#cite_note-71">[71]</a> Other genetic screens in the Beutler laboratory were used to identify genes that mediate homeostatic adaptations of the <a href="/wiki/Intestinal_epithelium" title="Intestinal epithelium">intestinal epithelium</a> following a cytotoxic insult;<a href="#cite_note-72">[72]</a><a href="#cite_note-73">[73]</a><a href="#cite_note-74">[74]</a><a href="#cite_note-75">[75]</a><a href="#cite_note-76">[76]</a><a href="#cite_note-77">[77]</a><a href="#cite_note-78">[78]</a> prevent allergic responses,<a href="#cite_note-79">[79]</a> diabetes,<a href="#cite_note-:12-80">[80]</a><a href="#cite_note-:13-81">[81]</a> or obesity;<a href="#cite_note-:14-82">[82]</a><a href="#cite_note-83">[83]</a><a href="#cite_note-:15-84">[84]</a> support normal hematopoiesis;<a href="#cite_note-:16-85">[85]</a><a href="#cite_note-86">[86]</a><a href="#cite_note-87">[87]</a><a href="#cite_note-:17-88">[88]</a><a href="#cite_note-:18-89">[89]</a><a href="#cite_note-:19-90">[90]</a><a href="#cite_note-:20-91">[91]</a><a href="#cite_note-:21-92">[92]</a><a href="#cite_note-:22-93">[93]</a><a href="#cite_note-:23-94">[94]</a> and enable humoral and cellular immunity.<a href="#cite_note-95">[95]</a><a href="#cite_note-96">[96]</a><a href="#cite_note-97">[97]</a><a href="#cite_note-:24-98">[98]</a> Some of these (beginning ~2015) were identified by a new process called automated meiotic mapping, which enabled greatly accelerated mutation identification compared to traditional genetic mapping (see below). In the course of their work, Beutler and his colleagues also discovered genes required for biological processes such as normal <a href="/wiki/Human_iron_metabolism" title="Human iron metabolism">iron absorption</a>,<a href="#cite_note-99">[99]</a> <a href="/wiki/Hearing" title="Hearing">hearing</a>,<a href="#cite_note-100">[100]</a> pigmentation,<a href="#cite_note-101">[101]</a><a href="#cite_note-102">[102]</a> metabolism,<a href="#cite_note-:14-82">[82]</a><a href="#cite_note-:15-84">[84]</a><a href="#cite_note-103">[103]</a><a href="#cite_note-:25-104">[104]</a><a href="#cite_note-:26-105">[105]</a> and <a href="/wiki/Embryogenesis" class="mw-redirect" title="Embryogenesis">embryonic development</a>.<a href="#cite_note-106">[106]</a> Many human diseases were ultimately linked to variants in the corresponding human genes after initial identification in the mouse by the Beutler laboratory,<a href="#cite_note-:11-66">[66]</a><a href="#cite_note-107">[107]</a><a href="#cite_note-108">[108]</a><a href="#cite_note-109">[109]</a> or by the laboratories of collaborating investigators.<a href="#cite_note-:27-110">[110]</a><a href="#cite_note-:28-111">[111]</a><a href="#cite_note-:29-112">[112]</a> <div class="mw-heading mw-heading3"><h3 id="Invention_of_Automated_Meiotic_Mapping">Invention of Automated Meiotic Mapping</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=7" title="Edit section: Invention of Automated Meiotic Mapping">edit</a>]</div> Prior to 2013, despite the development of methods for <a href="/wiki/Massive_parallel_sequencing" title="Massive parallel sequencing">massively parallel sequencing</a> and their application in finding induced germline mutations,<a href="#cite_note-113">[113]</a><a href="#cite_note-114">[114]</a><a href="#cite_note-115">[115]</a> positional cloning remained a slow process, limited by the need to genetically map mutations to chromosomal intervals to ascertain which induced mutation (among the average of approximately 60 changes in coding and splicing function induced per pedigree) was responsible for an observed phenotype. This required expansion of a mutant stock, outcrossing to a mapping strain, backcrossing, and genotypic and phenotypic analysis of F2 offspring. Moreover, when phenotypic screening was performed prior to positional cloning, only large effect size mutations (producing essentially qualitative phenotypes) were recoverable. Beutler invented a means of instantly identifying ENU-induced mutations that cause phenotypes.<a href="#cite_note-:30-116">[116]</a> The process, called automated meiotic mapping (AMM), eliminates the need to breed mutant mice to a mapping strain as required in classical <a href="/wiki/Genetic_linkage" title="Genetic linkage">genetic mapping</a> and flags causative mutations as soon as phenotypic assay data are collected. In a laboratory setting, it accelerates positional cloning approximately 200-fold, and permits ongoing measurement of genome saturation as mutagenesis progresses.<a href="#cite_note-117">[117]</a> Not only qualitative phenotypes, but subtle quantitative phenotypes, are detectable and mapped to individual mutations; hence the sensitivity of forward genetics is dramatically increased. AMM depends on statistical computation to detect associations between mutations in either the homozygous or heterozygous state and deviant phenotypes.<a href="#cite_note-:30-116">[116]</a> In addition, machine learning software, trained on the outcome of many thousands of experiments in which putative causative mutations were re-created and re-assayed for phenotype, is used to assess data quality.<a href="#cite_note-:31-118">[118]</a> As of 2022, more than 260,000 ENU-induced non-synonymous coding or splice site mutations had been assayed for phenotypic effects, and more than 5,800 mutations in approximately 2,500 genes had been declared causative of phenotype(s). For certain screens, such as <a href="/wiki/Flow_cytometry" title="Flow cytometry">flow cytometry</a> performed on the blood of germline mutant mice, more than 55% saturation of the genome has been achieved (i.e., more than 55% of all genes in which mutations will create flow cytometric aberrations in the peripheral blood have been detected, most of them based on assessment of multiple alleles, as of July 2021).<a href="#cite_note-:31-118">[118]</a> AMM led to the discovery of many new immunodeficiency disorders,<a href="#cite_note-:17-88">[88]</a><a href="#cite_note-:18-89">[89]</a><a href="#cite_note-:19-90">[90]</a><a href="#cite_note-:20-91">[91]</a><a href="#cite_note-:21-92">[92]</a><a href="#cite_note-:22-93">[93]</a><a href="#cite_note-:23-94">[94]</a><a href="#cite_note-:16-85">[85]</a><a href="#cite_note-:24-98">[98]</a> and disorders of bone morphology or mineral density,<a href="#cite_note-:28-111">[111]</a><a href="#cite_note-:29-112">[112]</a> vision,<a href="#cite_note-119">[119]</a> and metabolism.<a href="#cite_note-:14-82">[82]</a><a href="#cite_note-:15-84">[84]</a><a href="#cite_note-:25-104">[104]</a><a href="#cite_note-:26-105">[105]</a> Of note, AMM was used in the identification of a chemosensor that mediates innate fear behavior in mice and an autism gene found first in mice and then shown to cause autism in humans.<a href="#cite_note-:27-110">[110]</a><a href="#cite_note-120">[120]</a> AMM has also permitted high speed searches for mutations that suppress or augment disease phenotypes; for example, the development of autoimmune (Type 1) diabetes in mice of the NOD strain.<a href="#cite_note-:12-80">[80]</a><a href="#cite_note-:13-81">[81]</a> It offers a rational way to investigate the pathogenesis of complex disease phenotypes in general, in which many loci invariably contribute to susceptibility or resistance to disease, and disease occurs in those individuals with an unfavorable imbalance between these opposing influences. <div class="mw-heading mw-heading3"><h3 id="Developing_drugs_that_activate_TLRs">Developing drugs that activate TLRs</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=8" title="Edit section: Developing drugs that activate TLRs">edit</a>]</div> Beutler has collaborated with <a href="/wiki/Dale_L._Boger" title="Dale L. Boger">Dale L. Boger</a> and his research group to identify synthetic small molecule agonists of mammalian TLRs, which may be used in combination with defined molecular antigens to precisely target and coordinate innate and adaptive immune responses. Neoseptins, small molecules with no relationship to the structure of LPS, were shown to bind to the TLR4-MD2 complex in such a manner that two drug molecules trigger a conformational change similar to that elicited by an authentic LPS molecule. Diprovocims, which bear no structural similarity to bacterial lipopeptides, activate the TLR1-TLR2 heterodimer complex that normally acts as a receptor for tri-acylated lipopeptide molecules. These studies demonstrated that TLR2 and TLR4 can indeed respond to molecules other than classical microbial ligands, and set a new standard for verifying such interactions, in that <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a> was used to demonstrate the binding of neoseptins and diprovocims to their respective TLR targets at atomic level resolution. Beutler and colleagues also showed, again using X-ray crystallography combined with biological assays, that endogenous sulfatides are capable of binding to the TLR4-MD2 complex, causing its activation.<a href="#cite_note-121">[121]</a><a href="#cite_note-122">[122]</a><a href="#cite_note-123">[123]</a><a href="#cite_note-124">[124]</a><a href="#cite_note-125">[125]</a><a href="#cite_note-126">[126]</a><a href="#cite_note-127">[127]</a> <div class="mw-heading mw-heading2"><h2 id="Awards_and_recognition">Awards and recognition</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=9" title="Edit section: Awards and recognition">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-BLP_sources_section plainlinks metadata ambox ambox-content ambox-BLP_sources" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/40px-Ambox_important.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/60px-Ambox_important.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/80px-Ambox_important.svg.png 2x" data-file-width="40" data-file-height="40" /></div></td><td class="mbox-text"><div class="mbox-text-span">This section of a <a href="/wiki/Wikipedia:Biographies_of_living_persons" title="Wikipedia:Biographies of living persons">biography of a living person</a> needs additional <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">citations</a> for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a>. Please help by adding <a href="/wiki/Wikipedia:Reliable_sources" title="Wikipedia:Reliable sources">reliable sources</a>. <a href="/wiki/Wikipedia:Biographies_of_living_persons#Remove_contentious_material_that_is_unsourced_or_poorly_sourced" title="Wikipedia:Biographies of living persons">Contentious material</a> about living persons that is unsourced or <a href="/wiki/Wikipedia:Reliable_sources#Questionable_and_self-published_sources" title="Wikipedia:Reliable sources">poorly sourced</a> must be removed immediately from the article and its talk page, especially if potentially <a href="/wiki/Wikipedia:Libel" title="Wikipedia:Libel">libelous</a>. Find sources: <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Bruce+Beutler%22">"Bruce Beutler"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Bruce+Beutler%22+-wikipedia&tbs=ar:1">news</a> · <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Bruce+Beutler%22&tbs=bkt:s&tbm=bks">newspapers</a> · <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Bruce+Beutler%22+-wikipedia">books</a> · <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Bruce+Beutler%22">scholar</a> · <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Bruce+Beutler%22&acc=on&wc=on">JSTOR</a> (August 2023) (<a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a>)</div></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg/220px-Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg/330px-Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ac/Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg/440px-Nobel_Prize_2011-Press_Conference_KI-DSC_7609.jpg 2x" data-file-width="4723" data-file-height="3149" /></a><figcaption><a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules A. Hoffmann</a>, <a href="/wiki/G%C3%B6ran_K._Hansson" title="Göran K. Hansson">Göran K. Hansson</a> (chairman of the <a href="/wiki/Nobel_Committee_for_Physiology_or_Medicine" title="Nobel Committee for Physiology or Medicine">Nobel Committee for Physiology or Medicine</a>) and Beutler</figcaption></figure><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg/220px-Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg" decoding="async" width="220" height="330" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/97/Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg/330px-Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/97/Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg/440px-Nobel_Prize_2011-Press_Conference_KI-DSC_7568.jpg 2x" data-file-width="2175" data-file-height="3263" /></a><figcaption><a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules A. Hoffmann</a> (background) and Beutler</figcaption></figure><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg/220px-Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg/330px-Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg/440px-Nobel_Prize_2011-Press_Conference_KI-DSC_7512.jpg 2x" data-file-width="3250" data-file-height="2167" /></a><figcaption>Bruce Beutler at the Nobel Prize press conference at Karolinska, Solna</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Awards">Awards</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=10" title="Edit section: Awards">edit</a>]</div> <ul><li>1993 - <a href="/wiki/Alexander_von_Humboldt_Fellow" class="mw-redirect" title="Alexander von Humboldt Fellow">Alexander von Humboldt Fellow</a>; Germany</li> <li>1994 - Young Investigator Award (American Federation for Clinical Research); United States</li> <li>2001 - “Highly Cited” Researcher, <a href="/wiki/Institute_for_Scientific_Information" title="Institute for Scientific Information">Institute for Scientific Information</a>; United States</li> <li>2004 - <a href="/wiki/Robert_Koch_Medal_and_Award" title="Robert Koch Medal and Award">Robert Koch Prize</a> (Robert Koch Stiftung); Germany (shared with Jules A. Hoffmann and Shizuo Akira)</li> <li>2006 - <a href="/wiki/William_B._Coley_Award" title="William B. Coley Award">William B. Coley Award</a> (Cancer Research Institute); United States (shared with Shizuo Akira)</li> <li>2006 - <a href="/wiki/Grand_Prix_Charles-Leopold_Mayer" title="Grand Prix Charles-Leopold Mayer">Grand Prix Charles-Léopold Mayer</a> (Académie des Sciences); France</li> <li>2007 - Recipient of NIH/NIGMS MERIT Award; United States</li> <li>2007 - <a href="/wiki/Balzan_Prize" title="Balzan Prize">Balzan Prize</a> (International Balzan Foundation); Italy and Switzerland (shared with Jules A. Hoffmann)<a href="#cite_note-128">[128]</a></li> <li>2008 - Frederik B. Bang Award (The Stanley Watson Foundation); United States</li> <li>2008 - “<a href="/wiki/Clarivate_Citation_Laureates" title="Clarivate Citation Laureates">Citation Laureate</a>,” Thomson Reuters</li> <li>2009 - Will Rogers Institute Annual Prize for Scientific Research; United States</li> <li>2009 - <a href="/wiki/Albany_Medical_Center_Prize" title="Albany Medical Center Prize">Albany Medical Center Prize in Medicine and Biomedical Research</a>; United States (shared with <a href="/wiki/Charles_A._Dinarello" title="Charles A. Dinarello">Charles A. Dinarello</a> and Ralph M. Steinman)<a href="#cite_note-129">[129]</a></li> <li>2010 - University of Chicago, Professional Achievement Citation; United States</li> <li>2011 - <a href="/wiki/Shaw_Prize" title="Shaw Prize">Shaw Prize</a>; China (shared with Jules A. Hoffmann and Ruslan M. Medzhitov)<a href="#cite_note-130">[130]</a></li> <li>2011 - Nobel Prize in Physiology or Medicine; Sweden (shared with Jules A. Hoffmann and Ralph M. Steinman)<a href="#cite_note-Nobel-1">[1]</a></li> <li>2012 - Drexel Medicine Prize in Immunology; United States</li> <li>2013 - Rabbi Shai Shacknai Memorial Prize in Immunology and Cancer Research, The Hebrew University of Jerusalem; Israel</li> <li>2013 - Distinguished Service Award, University of Chicago; United States</li> <li>2013 - Korsmeyer Award; United States</li> <li>2016 - UCSD Distinguished Alumnus Award; United States</li></ul> <div class="mw-heading mw-heading3"><h3 id="Honorary_Doctoral_Degrees">Honorary Doctoral Degrees</h3>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=11" title="Edit section: Honorary Doctoral Degrees">edit</a>]</div> <ul><li>2007 - Doctor Med. Honoris Causa, <a href="/wiki/Technical_University_of_Munich" title="Technical University of Munich">Technical University of Munich</a>; Germany</li> <li>2009 - Honorary Doctoral Degree, <a href="/wiki/Xiamen_University" title="Xiamen University">Xiamen University</a>; China</li> <li>2012 - Honorary Professor, <a href="/wiki/Trinity_College_Dublin" title="Trinity College Dublin">Trinity College</a>; Ireland</li> <li>2013 - Honorary Professor, <a href="/wiki/Peking_University" title="Peking University">Peking University</a>; China</li> <li>2014 - Honorary Professor, Shanghai Jiao Tong University; China</li> <li>2014 - Chair of the Beutler Institute Council, Xiamen University; China</li> <li>2014 - Honorary Professor, Xiamen University; China</li> <li>2015 - Doctor Honoris Causa, University of Chile; Chile</li> <li>2015 - Doctor Honoris Causa, University of Marseille; France</li> <li>2015 - Doctor Honoris Causa, University of Brasilia; Brazil</li> <li>2015 - Doctor Honoris Causa,<a href="#cite_note-131">[131]</a> <a href="/wiki/Norwegian_University_of_Science_and_Technology" title="Norwegian University of Science and Technology">Norwegian University of Science and Technology</a> (NTNU); Norway</li> <li>2015 - Honorary Professor at Naresuan University; Thailand</li> <li>2016 - Honorary Doctorate, University of Athens; Greece</li> <li>2017 - Doctor Med. Honoris Causa, University of Ottawa; Canada</li> <li>2017 - Honorary Professor, Tianjin University; China</li> <li>2019 - Honorary Degree, Jewish Theological Seminary; United States</li> <li>2019 - Laurea Magistrale honoris causa in Medicina e Chirurgia (LM41),<a href="#cite_note-132">[132]</a> Universita Magna Grecia of Catanzaro; Italy</li></ul> <div class="mw-heading mw-heading2"><h2 id="Family">Family</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=12" title="Edit section: Family">edit</a>]</div> Bruce Beutler was the third son of Ernest Beutler (1928-2008) and Brondelle May Beutler (née Fleisher; 1928-2019). His siblings included two older brothers (Steven [b. 1952] and Earl [b. 1954]), and a younger sister, Deborah [b. 1962]).<a href="#cite_note-133">[133]</a> Ernest Beutler was a hematologist and medical geneticist famed for his studies of G-6-PD deficiency,<a href="#cite_note-134">[134]</a> other <a href="/wiki/Hemolytic_anemia" title="Hemolytic anemia">hemolytic anemias</a>,<a href="#cite_note-135">[135]</a><a href="#cite_note-136">[136]</a> iron metabolism,<a href="#cite_note-137">[137]</a> glycolipid storage diseases,<a href="#cite_note-138">[138]</a> and leukemias,<a href="#cite_note-139">[139]</a><a href="#cite_note-140">[140]</a> as well as his discovery of X chromosome inactivation.<a href="#cite_note-141">[141]</a> He was a Professor and department chairman at The Scripps Research Institute contemporaneously with Bruce. The two collaborated productively on several topics prior to Ernest Beutler’s death in 2008.<a href="#cite_note-:32-12">[12]</a><a href="#cite_note-:33-13">[13]</a><a href="#cite_note-142">[142]</a><a href="#cite_note-143">[143]</a><a href="#cite_note-144">[144]</a><a href="#cite_note-145">[145]</a> Both of Ernest Beutler’s parents were physicians.<a href="#cite_note-146">[146]</a> Bruce Beutler’s paternal grandmother, Kathe Beutler (née Italiener, daughter of Anna Rothstein, 1896-1999),<a href="#cite_note-147">[147]</a> was a pediatrician, trained at the Charité hospital in Berlin, earning her medical diploma in 1923. Käthe Italiener married Alfred Beutler in 1925. Also a physician, Alfred Beutler was a cousin to the spectral physicist, Hans G. Beutler (1896-1942), who worked at the Kaiser Wilhelm Institute and the University of Berlin before emigrating to the USA in 1936. He continued his work at the University of Chicago until his death.<a href="#cite_note-148">[148]</a> Bruce Beutler married Barbara Beutler (née Lanzl) in 1980 and divorced in 1988. Three sons were born to the couple.<a href="#cite_note-149">[149]</a><a href="#cite_note-150">[150]</a><a href="#cite_note-:0-11">[11]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=13" title="Edit section: See also">edit</a>]</div> <ul><li><a href="/wiki/List_of_Jewish_Nobel_laureates" title="List of Jewish Nobel laureates">List of Jewish Nobel laureates</a></li></ul> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=14" title="Edit section: References">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-Nobel-1">^ <a href="#cite_ref-Nobel_1-0">a</a> <a href="#cite_ref-Nobel_1-1">b</a> <a href="#cite_ref-Nobel_1-2">c</a> <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 class="citation pressrelease cs1"><a rel="nofollow" class="external text" href="https://www.nobelprize.org/nobel_prizes/medicine/laureates/2011/press.html">"Nobel Prize in Physiology or Medicine 2011"</a> (Press release). <a href="/wiki/Nobel_Foundation" title="Nobel Foundation">Nobel Foundation</a>. 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Haematologica. 94 (1): 154–156. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.3324%2Fhaematol.13863">10.3324/haematol.13863</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1592-8721">1592-8721</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2625414">2625414</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19118377">19118377</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:43531611">43531611</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Haematologica&rft.atitle=Ernest+Beutler+%281928%E2%80%932008%29&rft.volume=94&rft.issue=1&rft.pages=154-156&rft.date=2009-01-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2625414%23id-name%3DPMC&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A43531611%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.3324%2Fhaematol.13863&rft.issn=1592-8721&rft_id=info%3Apmid%2F19118377&rft.aulast=Beutler&rft.aufirst=Bruce&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2625414&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-134"><a href="#cite_ref-134">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutler1959" class="citation journal cs1">Beutler, E. (February 1959). <a rel="nofollow" class="external text" href="https://doi.org/10.1182%2Fblood.V14.2.103.103">"The hemolytic effect of primaquine and related compounds: a review"</a>. Blood. 14 (2): 103–139. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1182%2Fblood.V14.2.103.103">10.1182/blood.V14.2.103.103</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0006-4971">0006-4971</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/13618370">13618370</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Blood&rft.atitle=The+hemolytic+effect+of+primaquine+and+related+compounds%3A+a+review&rft.volume=14&rft.issue=2&rft.pages=103-139&rft.date=1959-02&rft.issn=0006-4971&rft_id=info%3Apmid%2F13618370&rft_id=info%3Adoi%2F10.1182%2Fblood.V14.2.103.103&rft.aulast=Beutler&rft.aufirst=E.&rft_id=https%3A%2F%2Fdoi.org%2F10.1182%252Fblood.V14.2.103.103&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-135"><a href="#cite_ref-135">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutler1971" class="citation book cs1">Beutler, Ernest (1971). Red Cell Metabolism: A Handbook of Biochemical Methods. New York: Grune and Stratton.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Red+Cell+Metabolism%3A+A+Handbook+of+Biochemical+Methods&rft.place=New+York&rft.pub=Grune+and+Stratton&rft.date=1971&rft.aulast=Beutler&rft.aufirst=Ernest&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-136"><a href="#cite_ref-136">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutler2006" class="citation book cs1">Beutler E (2006). "Disorders of red cells resulting from enzyme abnormalitites". In Lichtman MA, Beutler E, Kipps TJ, Seligsohn U, Kaushansky K, Prchal JT (eds.). Williams Hematology. New York: McGraw-Hill. pp. 603–632.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Disorders+of+red+cells+resulting+from+enzyme+abnormalitites&rft.btitle=Williams+Hematology&rft.place=New+York&rft.pages=603-632&rft.pub=McGraw-Hill&rft.date=2006&rft.aulast=Beutler&rft.aufirst=Ernest&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-137"><a href="#cite_ref-137">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutler1961" class="citation journal cs1">Beutler, E (February 1961). <a rel="nofollow" class="external text" href="https://www.annualreviews.org/doi/10.1146/annurev.me.12.020161.001211">"Hematology: Iron Metabolism"</a>. Annual Review of Medicine. 12 (1): 195–210. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.me.12.020161.001211">10.1146/annurev.me.12.020161.001211</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0066-4219">0066-4219</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Medicine&rft.atitle=Hematology%3A+Iron+Metabolism&rft.volume=12&rft.issue=1&rft.pages=195-210&rft.date=1961-02&rft_id=info%3Adoi%2F10.1146%2Fannurev.me.12.020161.001211&rft.issn=0066-4219&rft.aulast=Beutler&rft.aufirst=E&rft_id=https%3A%2F%2Fwww.annualreviews.org%2Fdoi%2F10.1146%2Fannurev.me.12.020161.001211&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-138"><a href="#cite_ref-138">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutler2006" class="citation journal cs1">Beutler, Ernest (July 2006). <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16515872">"Lysosomal storage diseases: natural history and ethical and economic aspects"</a>. Molecular Genetics and Metabolism. 88 (3): 208–215. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ymgme.2006.01.010">10.1016/j.ymgme.2006.01.010</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1096-7192">1096-7192</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16515872">16515872</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Genetics+and+Metabolism&rft.atitle=Lysosomal+storage+diseases%3A+natural+history+and+ethical+and+economic+aspects&rft.volume=88&rft.issue=3&rft.pages=208-215&rft.date=2006-07&rft.issn=1096-7192&rft_id=info%3Apmid%2F16515872&rft_id=info%3Adoi%2F10.1016%2Fj.ymgme.2006.01.010&rft.aulast=Beutler&rft.aufirst=Ernest&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F16515872&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-139"><a href="#cite_ref-139">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutlerBlumeBrossChillar1979" class="citation journal cs1">Beutler, E.; Blume, K. G.; Bross, K. J.; Chillar, R. K.; Ellington, O. B.; Fahey, J. L.; Farbstein, M. J.; Schmidt, G. M.; Spruce, W. E.; Turner, M. A. (1979). <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/398617">"Bone marrow transplantation as the treatment of choice for "good risk" adult patients with acute leukemia"</a>. Transactions of the Association of American Physicians. 92: 189–195. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0066-9458">0066-9458</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/398617">398617</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Transactions+of+the+Association+of+American+Physicians&rft.atitle=Bone+marrow+transplantation+as+the+treatment+of+choice+for+%22good+risk%22+adult+patients+with+acute+leukemia&rft.volume=92&rft.pages=189-195&rft.date=1979&rft_id=info%3Apmid%2F398617&rft.issn=0066-9458&rft.aulast=Beutler&rft.aufirst=E.&rft.au=Blume%2C+K.+G.&rft.au=Bross%2C+K.+J.&rft.au=Chillar%2C+R.+K.&rft.au=Ellington%2C+O.+B.&rft.au=Fahey%2C+J.+L.&rft.au=Farbstein%2C+M.+J.&rft.au=Schmidt%2C+G.+M.&rft.au=Spruce%2C+W.+E.&rft.au=Turner%2C+M.+A.&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F398617&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-140"><a href="#cite_ref-140">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPiroCarreraCarsonBeutler1990" class="citation journal cs1">Piro, L. D.; Carrera, C. J.; Carson, D. A.; Beutler, E. (April 19, 1990). <a rel="nofollow" class="external text" href="https://doi.org/10.1056%2FNEJM199004193221605">"Lasting remissions in hairy-cell leukemia induced by a single infusion of 2-chlorodeoxyadenosine"</a>. The New England Journal of Medicine. 322 (16): 1117–1121. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1056%2FNEJM199004193221605">10.1056/NEJM199004193221605</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0028-4793">0028-4793</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/1969613">1969613</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+New+England+Journal+of+Medicine&rft.atitle=Lasting+remissions+in+hairy-cell+leukemia+induced+by+a+single+infusion+of+2-chlorodeoxyadenosine&rft.volume=322&rft.issue=16&rft.pages=1117-1121&rft.date=1990-04-19&rft.issn=0028-4793&rft_id=info%3Apmid%2F1969613&rft_id=info%3Adoi%2F10.1056%2FNEJM199004193221605&rft.aulast=Piro&rft.aufirst=L.+D.&rft.au=Carrera%2C+C.+J.&rft.au=Carson%2C+D.+A.&rft.au=Beutler%2C+E.&rft_id=https%3A%2F%2Fdoi.org%2F10.1056%252FNEJM199004193221605&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-141"><a href="#cite_ref-141">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutlerYehFairbanks1962" class="citation journal cs1">Beutler, E.; Yeh, M.; Fairbanks, V. F. (January 15, 1962). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC285481">"The normal human female as a mosaic of X-chromosome activity: studies using the gene for C-6-PD-deficiency as a marker"</a>. Proceedings of the National Academy of Sciences of the United States of America. 48 (1): 9–16. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1962PNAS...48....9B">1962PNAS...48....9B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.48.1.9">10.1073/pnas.48.1.9</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0027-8424">0027-8424</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC285481">285481</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/13868717">13868717</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=The+normal+human+female+as+a+mosaic+of+X-chromosome+activity%3A+studies+using+the+gene+for+C-6-PD-deficiency+as+a+marker&rft.volume=48&rft.issue=1&rft.pages=9-16&rft.date=1962-01-15&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC285481%23id-name%3DPMC&rft_id=info%3Abibcode%2F1962PNAS...48....9B&rft_id=info%3Apmid%2F13868717&rft_id=info%3Adoi%2F10.1073%2Fpnas.48.1.9&rft.issn=0027-8424&rft.aulast=Beutler&rft.aufirst=E.&rft.au=Yeh%2C+M.&rft.au=Fairbanks%2C+V.+F.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC285481&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-142"><a href="#cite_ref-142">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutlerBeutler2002" class="citation journal cs1">Beutler, Bruce; Beutler, Ernest (December 12, 2002). <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12479194">"Toll-like receptor 4 polymorphisms and atherogenesis"</a>. The New England Journal of Medicine. 347 (24): 1978–1980, author reply 1978–1980. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1056%2FNEJM200212123472416">10.1056/NEJM200212123472416</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1533-4406">1533-4406</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12479194">12479194</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+New+England+Journal+of+Medicine&rft.atitle=Toll-like+receptor+4+polymorphisms+and+atherogenesis&rft.volume=347&rft.issue=24&rft.pages=1978-1980%2C+author+reply+1978-1980&rft.date=2002-12-12&rft.issn=1533-4406&rft_id=info%3Apmid%2F12479194&rft_id=info%3Adoi%2F10.1056%2FNEJM200212123472416&rft.aulast=Beutler&rft.aufirst=Bruce&rft.au=Beutler%2C+Ernest&rft_id=https%3A%2F%2Fpubmed.ncbi.nlm.nih.gov%2F12479194&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-143"><a href="#cite_ref-143">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutlerGelbartHanKoziol1989" class="citation journal cs1">Beutler, E.; Gelbart, T.; Han, J. H.; Koziol, J. A.; Beutler, B. (January 1989). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC286430">"Evolution of the genome and the genetic code: selection at the dinucleotide level by methylation and polyribonucleotide cleavage"</a>. Proceedings of the National Academy of Sciences of the United States of America. 86 (1): 192–196. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1989PNAS...86..192B">1989PNAS...86..192B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.86.1.192">10.1073/pnas.86.1.192</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0027-8424">0027-8424</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC286430">286430</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/2463621">2463621</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Evolution+of+the+genome+and+the+genetic+code%3A+selection+at+the+dinucleotide+level+by+methylation+and+polyribonucleotide+cleavage&rft.volume=86&rft.issue=1&rft.pages=192-196&rft.date=1989-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC286430%23id-name%3DPMC&rft_id=info%3Abibcode%2F1989PNAS...86..192B&rft_id=info%3Apmid%2F2463621&rft_id=info%3Adoi%2F10.1073%2Fpnas.86.1.192&rft.issn=0027-8424&rft.aulast=Beutler&rft.aufirst=E.&rft.au=Gelbart%2C+T.&rft.au=Han%2C+J.+H.&rft.au=Koziol%2C+J.+A.&rft.au=Beutler%2C+B.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC286430&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-144"><a href="#cite_ref-144">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTruksaGelbartPengBeutler2009" class="citation journal cs1">Truksa, Jaroslav; Gelbart, Terri; Peng, Hongfan; Beutler, Ernest; Beutler, Bruce; Lee, Pauline (November 2009). <a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2141.2009.07873.x">"Suppression of the hepcidin-encoding gene Hamp permits iron overload in mice lacking both hemojuvelin and matriptase-2/TMPRSS6"</a>. British Journal of Haematology. 147 (4): 571–581. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2141.2009.07873.x">10.1111/j.1365-2141.2009.07873.x</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1365-2141">1365-2141</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19751239">19751239</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:205266224">205266224</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=British+Journal+of+Haematology&rft.atitle=Suppression+of+the+hepcidin-encoding+gene+Hamp+permits+iron+overload+in+mice+lacking+both+hemojuvelin+and+matriptase-2%2FTMPRSS6&rft.volume=147&rft.issue=4&rft.pages=571-581&rft.date=2009-11&rft.issn=1365-2141&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A205266224%23id-name%3DS2CID&rft_id=info%3Apmid%2F19751239&rft_id=info%3Adoi%2F10.1111%2Fj.1365-2141.2009.07873.x&rft.aulast=Truksa&rft.aufirst=Jaroslav&rft.au=Gelbart%2C+Terri&rft.au=Peng%2C+Hongfan&rft.au=Beutler%2C+Ernest&rft.au=Beutler%2C+Bruce&rft.au=Lee%2C+Pauline&rft_id=https%3A%2F%2Fdoi.org%2F10.1111%252Fj.1365-2141.2009.07873.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-145"><a href="#cite_ref-145">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDuSheGelbartTruksa2008" class="citation journal cs1">Du, Xin; She, Ellen; Gelbart, Terri; Truksa, Jaroslav; Lee, Pauline; Xia, Yu; Khovananth, Kevin; Mudd, Suzanne; Mann, Navjiwan; Moresco, Eva Marie Y.; Beutler, Ernest; Beutler, Bruce (May 23, 2008). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2430097">"The serine protease TMPRSS6 is required to sense iron deficiency"</a>. Science. 320 (5879): 1088–1092. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008Sci...320.1088D">2008Sci...320.1088D</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1157121">10.1126/science.1157121</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1095-9203">1095-9203</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2430097">2430097</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18451267">18451267</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=The+serine+protease+TMPRSS6+is+required+to+sense+iron+deficiency&rft.volume=320&rft.issue=5879&rft.pages=1088-1092&rft.date=2008-05-23&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2430097%23id-name%3DPMC&rft_id=info%3Abibcode%2F2008Sci...320.1088D&rft_id=info%3Apmid%2F18451267&rft_id=info%3Adoi%2F10.1126%2Fscience.1157121&rft.issn=1095-9203&rft.aulast=Du&rft.aufirst=Xin&rft.au=She%2C+Ellen&rft.au=Gelbart%2C+Terri&rft.au=Truksa%2C+Jaroslav&rft.au=Lee%2C+Pauline&rft.au=Xia%2C+Yu&rft.au=Khovananth%2C+Kevin&rft.au=Mudd%2C+Suzanne&rft.au=Mann%2C+Navjiwan&rft.au=Moresco%2C+Eva+Marie+Y.&rft.au=Beutler%2C+Ernest&rft.au=Beutler%2C+Bruce&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2430097&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-146"><a href="#cite_ref-146">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWailoo" class="citation web cs1">Wailoo, Keith. <a rel="nofollow" class="external text" href="https://www.hematology.org/about/history/legends/ernest-beutler-bio/ernest-beutler-qa">"Ernest Beutler QA - Hematology.org"</a>. Retrieved March 9, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Ernest+Beutler+QA+-+Hematology.org&rft.aulast=Wailoo&rft.aufirst=Keith&rft_id=https%3A%2F%2Fwww.hematology.org%2Fabout%2Fhistory%2Flegends%2Fernest-beutler-bio%2Fernest-beutler-qa&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-147"><a href="#cite_ref-147">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHildebrandtKammertönsLechnerSchmitt2019" class="citation journal cs1">Hildebrandt, Sabine; Kammertöns, Thomas; Lechner, Christian; Schmitt, Philipp; Schumann, Ralf R. (2019). <a rel="nofollow" class="external text" href="https://biblioscout.net/article/10.25162/mhj-2019-0009">"Dr. Käthe Beutler, 1896–1999"</a>. Medizinhistorisches Journal. 54 (4): 294–346. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.25162%2Fmhj-2019-0009">10.25162/mhj-2019-0009</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0025-8431">0025-8431</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:213008951">213008951</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Medizinhistorisches+Journal&rft.atitle=Dr.+K%C3%A4the+Beutler%2C+1896%E2%80%931999&rft.volume=54&rft.issue=4&rft.pages=294-346&rft.date=2019&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A213008951%23id-name%3DS2CID&rft.issn=0025-8431&rft_id=info%3Adoi%2F10.25162%2Fmhj-2019-0009&rft.aulast=Hildebrandt&rft.aufirst=Sabine&rft.au=Kammert%C3%B6ns%2C+Thomas&rft.au=Lechner%2C+Christian&rft.au=Schmitt%2C+Philipp&rft.au=Schumann%2C+Ralf+R.&rft_id=https%3A%2F%2Fbiblioscout.net%2Farticle%2F10.25162%2Fmhj-2019-0009&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-148"><a href="#cite_ref-148">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation news cs1"><a rel="nofollow" class="external text" href="https://www.nytimes.com/1942/12/19/archives/hans-g-beutler-46-physicist-is-dead-research-aide-on-the-chicago-u.html">"HANS G. BEUTLER, 46, PHYSICIST, IS DEAD; Research Aide on the Chicago U. Faculty Was Spectroscopist"</a>. The New York Times. December 19, 1942. 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Retrieved November 30, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Lamplighter+Has+Ties+to+Nobel+Prize+Winner+-+People+Newspapers&rft.date=2011-10-05&rft_id=https%3A%2F%2Fwww.peoplenewspapers.com%2F2011%2F10%2F05%2Flamplighter-has-ties-to-nobel-prize-winner%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> <li id="cite_note-150"><a href="#cite_ref-150">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.shawprize.org/autobiography/bruce-a-beutler/">"Bruce A Beutler"</a>. The Shaw Prize. Retrieved November 30, 2023.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=The+Shaw+Prize&rft.atitle=Bruce+A+Beutler&rft_id=https%3A%2F%2Fwww.shawprize.org%2Fautobiography%2Fbruce-a-beutler%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABruce+Beutler" class="Z3988"> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Bruce_Beutler&action=edit&section=15" title="Edit section: External links">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output 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href="http://www.utsouthwestern.edu/labs/beutler/">Official website</a></li> <li><a href="/wiki/File:Wikisource-logo.svg" class="mw-file-description" title="Wikisource logo"><img alt="Wikisource logo" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/16px-Wikisource-logo.svg.png" decoding="async" width="16" height="17" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/24px-Wikisource-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Wikisource-logo.svg/32px-Wikisource-logo.svg.png 2x" data-file-width="410" data-file-height="430" /></a> Works on the topic <a href="https://en.wikisource.org/wiki/Portal:Bruce_Beutler" class="extiw" title="s:Portal:Bruce Beutler">Bruce Beutler</a> at <a href="/wiki/Wikisource" title="Wikisource">Wikisource</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links_2">External links</h2>[<a 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href="https://commons.wikimedia.org/wiki/Category:Bruce_Beutler" class="extiw" title="commons:Category:Bruce Beutler">Bruce Beutler</a>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://www.nobelprize.org/laureate/861">Bruce A. Beutler</a> on Nobelprize.org <a href="https://www.wikidata.org/wiki/Q104592#P8024" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a> including the Nobel lecture How Mammals Sense Infection: From Endotoxin to the Toll-like Receptors</li> <li><a rel="nofollow" class="external text" href="http://www.nobelprizeii.org/nobel-laureate/bruce-beutler/">Nobel Prize Inspiration Initiative</a></li> <li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/sites/entrez?db=pubmed&orig_db=pubmed&term=beutler%20b%20OR%20beutler%20ba&cmd=&cmd_current=">Scientific Publications</a> – All publications of articles by Bruce A. Beutler listed in PubMed.</li> <li><a rel="nofollow" class="external text" href="http://idw-online.de/pages/de/news237418">How we sense microbes: Genetic dissection of innate immunity in insects and mammals</a> – Brief review of recent work, written with Jules A. Hoffmann.</li> <li><a rel="nofollow" class="external text" href="http://circres.ahajournals.org/content/112/5/751.full">Persistent Prospector - MD. Bruce Beutler by Ruth Williams</a></li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=HNrx9-paRi0">2011 Video presentation by Dr. Bruce Beutler at University of Texas</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20170705095852/http://www.panoramio.com/photo/121662229">Discovery of TLR4 and his Nobel Prize Award displayed at Perot Museum of Nature and Science in Dallas, TX.</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 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navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Nobel_Prize_in_Physiology_or_Medicine" title="Template:Nobel Prize in Physiology or Medicine"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nobel_Prize_in_Physiology_or_Medicine" title="Template talk:Nobel Prize in Physiology or Medicine"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nobel_Prize_in_Physiology_or_Medicine" title="Special:EditPage/Template:Nobel Prize in Physiology or Medicine"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Laureates_of_the_Nobel_Prize_in_Physiology_or_Medicine" style="font-size:114%;margin:0 4em"><a href="/wiki/List_of_Nobel_laureates_in_Physiology_or_Medicine" title="List of Nobel laureates in Physiology or Medicine">Laureates</a> of the <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">1901–1925</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1901: <a href="/wiki/Emil_von_Behring" title="Emil von Behring">Emil Behring</a></li> <li>1902: <a href="/wiki/Ronald_Ross" title="Ronald Ross">Ronald Ross</a></li> <li>1903: <a href="/wiki/Niels_Ryberg_Finsen" title="Niels Ryberg Finsen">Niels Finsen</a></li> <li>1904: <a href="/wiki/Ivan_Pavlov" title="Ivan Pavlov">Ivan Pavlov</a></li> <li>1905: <a href="/wiki/Robert_Koch" title="Robert Koch">Robert Koch</a></li> <li>1906: <a href="/wiki/Camillo_Golgi" title="Camillo Golgi">Camillo Golgi</a> / <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a></li> <li>1907: <a href="/wiki/Charles_Louis_Alphonse_Laveran" title="Charles Louis Alphonse Laveran">Alphonse Laveran</a></li> <li>1908: <a href="/wiki/%C3%89lie_Metchnikoff" title="Élie Metchnikoff">Élie Metchnikoff</a> / <a href="/wiki/Paul_Ehrlich" title="Paul Ehrlich">Paul Ehrlich</a></li> <li>1909: <a href="/wiki/Emil_Theodor_Kocher" title="Emil Theodor Kocher">Emil Kocher</a></li> <li>1910: <a href="/wiki/Albrecht_Kossel" title="Albrecht Kossel">Albrecht Kossel</a></li> <li>1911: <a href="/wiki/Allvar_Gullstrand" title="Allvar Gullstrand">Allvar Gullstrand</a></li> <li>1912: <a href="/wiki/Alexis_Carrel" title="Alexis Carrel">Alexis Carrel</a></li> <li>1913: <a href="/wiki/Charles_Richet" title="Charles Richet">Charles Richet</a></li> <li>1914: <a href="/wiki/R%C3%B3bert_B%C3%A1r%C3%A1ny" class="mw-redirect" title="Róbert Bárány">Róbert Bárány</a></li> <li>1915</li> <li>1916</li> <li>1917</li> <li>1918</li> <li>1919: <a href="/wiki/Jules_Bordet" title="Jules Bordet">Jules Bordet</a></li> <li>1920: <a href="/wiki/August_Krogh" title="August Krogh">August Krogh</a></li> <li>1921</li> <li>1922: <a href="/wiki/Archibald_Hill" title="Archibald Hill">Archibald Hill</a> / <a href="/wiki/Otto_Fritz_Meyerhof" title="Otto Fritz Meyerhof">Otto Meyerhof</a></li> <li>1923: <a href="/wiki/Frederick_Banting" title="Frederick Banting">Frederick Banting</a> / <a href="/wiki/John_Macleod_(physiologist)" title="John Macleod (physiologist)">John Macleod</a></li> <li>1924: <a href="/wiki/Willem_Einthoven" title="Willem Einthoven">Willem Einthoven</a></li> <li>1925</li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/ed/Nobel_Prize.png/80px-Nobel_Prize.png" decoding="async" width="80" height="79" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/ed/Nobel_Prize.png/120px-Nobel_Prize.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/ed/Nobel_Prize.png/160px-Nobel_Prize.png 2x" data-file-width="500" data-file-height="492" /></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">1926–1950</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1926: <a href="/wiki/Johannes_Fibiger" title="Johannes Fibiger">Johannes Fibiger</a></li> <li>1927: <a href="/wiki/Julius_Wagner-Jauregg" title="Julius Wagner-Jauregg">Julius Wagner-Jauregg</a></li> <li>1928: <a href="/wiki/Charles_Nicolle" title="Charles Nicolle">Charles Nicolle</a></li> <li>1929: <a href="/wiki/Christiaan_Eijkman" title="Christiaan Eijkman">Christiaan Eijkman</a> / <a href="/wiki/Frederick_Gowland_Hopkins" title="Frederick Gowland Hopkins">Frederick Gowland Hopkins</a></li> <li>1930: <a href="/wiki/Karl_Landsteiner" title="Karl Landsteiner">Karl Landsteiner</a></li> <li>1931: <a href="/wiki/Otto_Heinrich_Warburg" title="Otto Heinrich Warburg">Otto Warburg</a></li> <li>1932: <a href="/wiki/Charles_Scott_Sherrington" title="Charles Scott Sherrington">Charles Scott Sherrington</a> / <a href="/wiki/Edgar_Adrian" title="Edgar Adrian">Edgar Adrian</a></li> <li>1933: <a href="/wiki/Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Morgan</a></li> <li>1934: <a href="/wiki/George_Whipple" title="George Whipple">George Whipple</a> / <a href="/wiki/George_Minot" title="George Minot">George Minot</a> / <a href="/wiki/William_P._Murphy" title="William P. Murphy">William Murphy</a></li> <li>1935: <a href="/wiki/Hans_Spemann" title="Hans Spemann">Hans Spemann</a></li> <li>1936: <a href="/wiki/Henry_Hallett_Dale" title="Henry Hallett Dale">Henry Dale</a> / <a href="/wiki/Otto_Loewi" title="Otto Loewi">Otto Loewi</a></li> <li>1937: <a href="/wiki/Albert_Szent-Gy%C3%B6rgyi" title="Albert Szent-Györgyi">Albert Szent-Györgyi</a></li> <li>1938: <a href="/wiki/Corneille_Heymans" title="Corneille Heymans">Corneille Heymans</a></li> <li>1939: <a href="/wiki/Gerhard_Domagk" title="Gerhard Domagk">Gerhard Domagk</a></li> <li>1940</li> <li>1941</li> <li>1942</li> <li>1943: <a href="/wiki/Henrik_Dam" title="Henrik Dam">Henrik Dam</a> / <a href="/wiki/Edward_Adelbert_Doisy" title="Edward Adelbert Doisy">Edward Doisy</a></li> <li>1944: <a href="/wiki/Joseph_Erlanger" title="Joseph Erlanger">Joseph Erlanger</a> / <a href="/wiki/Herbert_Spencer_Gasser" title="Herbert Spencer Gasser">Herbert Gasser</a></li> <li>1945: <a href="/wiki/Alexander_Fleming" title="Alexander Fleming">Alexander Fleming</a> / <a href="/wiki/Ernst_Chain" title="Ernst Chain">Ernst Chain</a> / <a href="/wiki/Howard_Florey" title="Howard Florey">Howard Florey</a></li> <li>1946: <a href="/wiki/Hermann_Joseph_Muller" title="Hermann Joseph Muller">Hermann Muller</a></li> <li>1947: <a href="/wiki/Carl_Ferdinand_Cori" title="Carl Ferdinand Cori">Carl Cori</a> / <a href="/wiki/Gerty_Cori" title="Gerty Cori">Gerty Cori</a> / <a href="/wiki/Bernardo_Houssay" title="Bernardo Houssay">Bernardo Houssay</a></li> <li>1948: <a href="/wiki/Paul_Hermann_M%C3%BCller" title="Paul Hermann Müller">Paul Müller</a></li> <li>1949: <a href="/wiki/Walter_Rudolf_Hess" title="Walter Rudolf Hess">Walter Hess</a> / <a href="/wiki/Ant%C3%B3nio_Egas_Moniz" title="António Egas Moniz">António Egas Moniz</a></li> <li>1950: <a href="/wiki/Edward_Calvin_Kendall" title="Edward Calvin Kendall">Edward Kendall</a> / <a href="/wiki/Tadeusz_Reichstein" title="Tadeusz Reichstein">Tadeusz Reichstein</a> / <a href="/wiki/Philip_Showalter_Hench" title="Philip Showalter Hench">Philip Hench</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">1951–1975</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1951: <a href="/wiki/Max_Theiler" title="Max Theiler">Max Theiler</a></li> <li>1952: <a href="/wiki/Selman_Waksman" title="Selman Waksman">Selman Waksman</a></li> <li>1953: <a href="/wiki/Hans_Krebs_(biochemist)" title="Hans Krebs (biochemist)">Hans Krebs</a> / <a href="/wiki/Fritz_Albert_Lipmann" title="Fritz Albert Lipmann">Fritz Lipmann</a></li> <li>1954: <a href="/wiki/John_Franklin_Enders" title="John Franklin Enders">John Enders</a> / <a href="/wiki/Thomas_Huckle_Weller" title="Thomas Huckle Weller">Thomas Weller</a> / <a href="/wiki/Frederick_Chapman_Robbins" title="Frederick Chapman Robbins">Frederick Robbins</a></li> <li>1955: <a href="/wiki/Hugo_Theorell" title="Hugo Theorell">Hugo Theorell</a></li> <li>1956: <a href="/wiki/Andr%C3%A9_Fr%C3%A9d%C3%A9ric_Cournand" title="André Frédéric Cournand">André Cournand</a> / <a href="/wiki/Werner_Forssmann" title="Werner Forssmann">Werner Forssmann</a> / <a href="/wiki/Dickinson_W._Richards" title="Dickinson W. Richards">Dickinson W. Richards</a></li> <li>1957: <a href="/wiki/Daniel_Bovet" title="Daniel Bovet">Daniel Bovet</a></li> <li>1958: <a href="/wiki/George_Beadle" title="George Beadle">George Beadle</a> / <a href="/wiki/Edward_Tatum" title="Edward Tatum">Edward Tatum</a> / <a href="/wiki/Joshua_Lederberg" title="Joshua Lederberg">Joshua Lederberg</a></li> <li>1959: <a href="/wiki/Severo_Ochoa" title="Severo Ochoa">Severo Ochoa</a> / <a href="/wiki/Arthur_Kornberg" title="Arthur Kornberg">Arthur Kornberg</a></li> <li>1960: <a href="/wiki/Macfarlane_Burnet" title="Macfarlane Burnet">Frank Burnet</a> / <a href="/wiki/Peter_Medawar" title="Peter Medawar">Peter Medawar</a></li> <li>1961: <a href="/wiki/Georg_von_B%C3%A9k%C3%A9sy" title="Georg von Békésy">Georg von Békésy</a></li> <li>1962: <a href="/wiki/Francis_Crick" title="Francis Crick">Francis Crick</a> / <a href="/wiki/James_Watson" title="James Watson">James Watson</a> / <a href="/wiki/Maurice_Wilkins" title="Maurice Wilkins">Maurice Wilkins</a></li> <li>1963: <a href="/wiki/John_Eccles_(neurophysiologist)" title="John Eccles (neurophysiologist)">John Eccles</a> / <a href="/wiki/Alan_Hodgkin" title="Alan Hodgkin">Alan Hodgkin</a> / <a href="/wiki/Andrew_Huxley" title="Andrew Huxley">Andrew Huxley</a></li> <li>1964: <a href="/wiki/Konrad_Emil_Bloch" title="Konrad Emil Bloch">Konrad Bloch</a> / <a href="/wiki/Feodor_Lynen" title="Feodor Lynen">Feodor Lynen</a></li> <li>1965: <a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> / <a href="/wiki/Andr%C3%A9_Michel_Lwoff" title="André Michel Lwoff">André Lwoff</a> / <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a></li> <li>1966: <a href="/wiki/Francis_Peyton_Rous" title="Francis Peyton Rous">Francis Rous</a> / <a href="/wiki/Charles_Brenton_Huggins" title="Charles Brenton Huggins">Charles B. Huggins</a></li> <li>1967: <a href="/wiki/Ragnar_Granit" title="Ragnar Granit">Ragnar Granit</a> / <a href="/wiki/Haldan_Keffer_Hartline" title="Haldan Keffer Hartline">Haldan Hartline</a> / <a href="/wiki/George_Wald" title="George Wald">George Wald</a></li> <li>1968: <a href="/wiki/Robert_W._Holley" title="Robert W. Holley">Robert W. Holley</a> / <a href="/wiki/Har_Gobind_Khorana" title="Har Gobind Khorana">Har Khorana</a> / <a href="/wiki/Marshall_Warren_Nirenberg" title="Marshall Warren Nirenberg">Marshall Nirenberg</a></li> <li>1969: <a href="/wiki/Max_Delbr%C3%BCck" title="Max Delbrück">Max Delbrück</a> / <a href="/wiki/Alfred_Hershey" title="Alfred Hershey">Alfred Hershey</a> / <a href="/wiki/Salvador_Luria" title="Salvador Luria">Salvador Luria</a></li> <li>1970: <a href="/wiki/Bernard_Katz" title="Bernard Katz">Bernard Katz</a> / <a href="/wiki/Ulf_von_Euler" title="Ulf von Euler">Ulf von Euler</a> / <a href="/wiki/Julius_Axelrod" title="Julius Axelrod">Julius Axelrod</a></li> <li>1971: <a href="/wiki/Earl_Wilbur_Sutherland_Jr." title="Earl Wilbur Sutherland Jr.">Earl Sutherland Jr.</a></li> <li>1972: <a href="/wiki/Gerald_Edelman" title="Gerald Edelman">Gerald Edelman</a> / <a href="/wiki/Rodney_Robert_Porter" class="mw-redirect" title="Rodney Robert Porter">Rodney Porter</a></li> <li>1973: <a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a> / <a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> / <a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Nikolaas Tinbergen</a></li> <li>1974: <a href="/wiki/Albert_Claude" title="Albert Claude">Albert Claude</a> / <a href="/wiki/Christian_de_Duve" title="Christian de Duve">Christian de Duve</a> / <a href="/wiki/George_Emil_Palade" title="George Emil Palade">George Palade</a></li> <li>1975: <a href="/wiki/David_Baltimore" title="David Baltimore">David Baltimore</a> / <a href="/wiki/Renato_Dulbecco" title="Renato Dulbecco">Renato Dulbecco</a> / <a href="/wiki/Howard_Martin_Temin" title="Howard Martin Temin">Howard Temin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">1976–2000</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1976: <a href="/wiki/Baruch_Samuel_Blumberg" title="Baruch Samuel Blumberg">Baruch Blumberg</a> / <a href="/wiki/Daniel_Carleton_Gajdusek" title="Daniel Carleton Gajdusek">Daniel Gajdusek</a></li> <li>1977: <a href="/wiki/Roger_Guillemin" title="Roger Guillemin">Roger Guillemin</a> / <a href="/wiki/Andrew_Schally" title="Andrew Schally">Andrew Schally</a> / <a href="/wiki/Rosalyn_Sussman_Yalow" title="Rosalyn Sussman Yalow">Rosalyn Yalow</a></li> <li>1978: <a href="/wiki/Werner_Arber" title="Werner Arber">Werner Arber</a> / <a href="/wiki/Daniel_Nathans" title="Daniel Nathans">Daniel Nathans</a> / <a href="/wiki/Hamilton_O._Smith" title="Hamilton O. Smith">Hamilton O. Smith</a></li> <li>1979: <a href="/wiki/Allan_MacLeod_Cormack" title="Allan MacLeod Cormack">Allan Cormack</a> / <a href="/wiki/Godfrey_Hounsfield" title="Godfrey Hounsfield">Godfrey Hounsfield</a></li> <li>1980: <a href="/wiki/Baruj_Benacerraf" title="Baruj Benacerraf">Baruj Benacerraf</a> / <a href="/wiki/Jean_Dausset" title="Jean Dausset">Jean Dausset</a> / <a href="/wiki/George_Davis_Snell" title="George Davis Snell">George Snell</a></li> <li>1981: <a href="/wiki/Roger_Wolcott_Sperry" title="Roger Wolcott Sperry">Roger Sperry</a> / <a href="/wiki/David_H._Hubel" title="David H. Hubel">David H. Hubel</a> / <a href="/wiki/Torsten_Wiesel" title="Torsten Wiesel">Torsten Wiesel</a></li> <li>1982: <a href="/wiki/Sune_Bergstr%C3%B6m" title="Sune Bergström">Sune Bergström</a> / <a href="/wiki/Bengt_I._Samuelsson" title="Bengt I. Samuelsson">Bengt I. Samuelsson</a> / <a href="/wiki/John_Vane" title="John Vane">John Vane</a></li> <li>1983: <a href="/wiki/Barbara_McClintock" title="Barbara McClintock">Barbara McClintock</a></li> <li>1984: <a href="/wiki/Niels_Kaj_Jerne" title="Niels Kaj Jerne">Niels Jerne</a> / <a href="/wiki/Georges_J._F._K%C3%B6hler" title="Georges J. F. Köhler">Georges Köhler</a> / <a href="/wiki/C%C3%A9sar_Milstein" title="César Milstein">César Milstein</a></li> <li>1985: <a href="/wiki/Michael_Stuart_Brown" title="Michael Stuart Brown">Michael Brown</a> / <a href="/wiki/Joseph_L._Goldstein" title="Joseph L. Goldstein">Joseph L. Goldstein</a></li> <li>1986: <a href="/wiki/Stanley_Cohen_(biochemist)" title="Stanley Cohen (biochemist)">Stanley Cohen</a> / <a href="/wiki/Rita_Levi-Montalcini" title="Rita Levi-Montalcini">Rita Levi-Montalcini</a></li> <li>1987: <a href="/wiki/Susumu_Tonegawa" title="Susumu Tonegawa">Susumu Tonegawa</a></li> <li>1988: <a href="/wiki/James_Black_(pharmacologist)" title="James Black (pharmacologist)">James W. Black</a> / <a href="/wiki/Gertrude_B._Elion" title="Gertrude B. Elion">Gertrude B. Elion</a> / <a href="/wiki/George_H._Hitchings" title="George H. Hitchings">George H. Hitchings</a></li> <li>1989: <a href="/wiki/J._Michael_Bishop" title="J. Michael Bishop">J. Michael Bishop</a> / <a href="/wiki/Harold_E._Varmus" title="Harold E. Varmus">Harold E. Varmus</a></li> <li>1990: <a href="/wiki/Joseph_Murray" class="mw-redirect" title="Joseph Murray">Joseph Murray</a> / <a href="/wiki/E._Donnall_Thomas" title="E. Donnall Thomas">E. Donnall Thomas</a></li> <li>1991: <a href="/wiki/Erwin_Neher" title="Erwin Neher">Erwin Neher</a> / <a href="/wiki/Bert_Sakmann" title="Bert Sakmann">Bert Sakmann</a></li> <li>1992: <a href="/wiki/Edmond_H._Fischer" title="Edmond H. Fischer">Edmond Fischer</a> / <a href="/wiki/Edwin_G._Krebs" title="Edwin G. Krebs">Edwin G. Krebs</a></li> <li>1993: <a href="/wiki/Richard_J._Roberts" title="Richard J. Roberts">Richard J. Roberts</a> / <a href="/wiki/Phillip_Allen_Sharp" title="Phillip Allen Sharp">Phillip Sharp</a></li> <li>1994: <a href="/wiki/Alfred_G._Gilman" title="Alfred G. Gilman">Alfred G. Gilman</a> / <a href="/wiki/Martin_Rodbell" title="Martin Rodbell">Martin Rodbell</a></li> <li>1995: <a href="/wiki/Edward_B._Lewis" title="Edward B. Lewis">Edward B. Lewis</a> / <a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a> / <a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li>1996: <a href="/wiki/Peter_C._Doherty" class="mw-redirect" title="Peter C. Doherty">Peter C. Doherty</a> / <a href="/wiki/Rolf_M._Zinkernagel" title="Rolf M. Zinkernagel">Rolf M. Zinkernagel</a></li> <li>1997: <a href="/wiki/Stanley_B._Prusiner" title="Stanley B. Prusiner">Stanley B. Prusiner</a></li> <li>1998: <a href="/wiki/Robert_F._Furchgott" title="Robert F. Furchgott">Robert F. Furchgott</a> / <a href="/wiki/Louis_Ignarro" title="Louis Ignarro">Louis Ignarro</a> / <a href="/wiki/Ferid_Murad" title="Ferid Murad">Ferid Murad</a></li> <li>1999: <a href="/wiki/G%C3%BCnter_Blobel" title="Günter Blobel">Günter Blobel</a></li> <li>2000: <a href="/wiki/Arvid_Carlsson" title="Arvid Carlsson">Arvid Carlsson</a> / <a href="/wiki/Paul_Greengard" title="Paul Greengard">Paul Greengard</a> / <a href="/wiki/Eric_Kandel" title="Eric Kandel">Eric Kandel</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">2001–present</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>2001: <a href="/wiki/Leland_H._Hartwell" title="Leland H. Hartwell">Leland H. Hartwell</a> / <a href="/wiki/Tim_Hunt" title="Tim Hunt">Tim Hunt</a> / <a href="/wiki/Paul_Nurse" title="Paul Nurse">Paul Nurse</a></li> <li>2002: <a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Sydney Brenner</a> / <a href="/wiki/H._Robert_Horvitz" title="H. Robert Horvitz">H. Robert Horvitz</a> / <a href="/wiki/John_Sulston" title="John Sulston">John E. Sulston</a></li> <li>2003: <a href="/wiki/Paul_Lauterbur" title="Paul Lauterbur">Paul Lauterbur</a> / <a href="/wiki/Peter_Mansfield" title="Peter Mansfield">Peter Mansfield</a></li> <li>2004: <a href="/wiki/Richard_Axel" title="Richard Axel">Richard Axel</a> / <a href="/wiki/Linda_B._Buck" title="Linda B. Buck">Linda B. Buck</a></li> <li>2005: <a href="/wiki/Barry_Marshall" title="Barry Marshall">Barry Marshall</a> / <a href="/wiki/Robin_Warren" title="Robin Warren">Robin Warren</a></li> <li>2006: <a href="/wiki/Andrew_Fire" title="Andrew Fire">Andrew Fire</a> / <a href="/wiki/Craig_Mello" title="Craig Mello">Craig Mello</a></li> <li>2007: <a href="/wiki/Mario_Capecchi" title="Mario Capecchi">Mario Capecchi</a> / <a href="/wiki/Martin_Evans" title="Martin Evans">Martin Evans</a> / <a href="/wiki/Oliver_Smithies" title="Oliver Smithies">Oliver Smithies</a></li> <li>2008: <a href="/wiki/Harald_zur_Hausen" title="Harald zur Hausen">Harald zur Hausen</a> / <a href="/wiki/Luc_Montagnier" title="Luc Montagnier">Luc Montagnier</a> / <a href="/wiki/Fran%C3%A7oise_Barr%C3%A9-Sinoussi" title="Françoise Barré-Sinoussi">Françoise Barré-Sinoussi</a></li> <li>2009: <a href="/wiki/Elizabeth_Blackburn" title="Elizabeth Blackburn">Elizabeth Blackburn</a> / <a href="/wiki/Carol_W._Greider" title="Carol W. Greider">Carol W. Greider</a> / <a href="/wiki/Jack_W._Szostak" title="Jack W. Szostak">Jack W. Szostak</a></li> <li>2010: <a href="/wiki/Robert_Edwards_(physiologist)" title="Robert Edwards (physiologist)">Robert G. Edwards</a></li> <li>2011: <a class="mw-selflink selflink">Bruce Beutler</a> / <a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules A. Hoffmann</a> / <a href="/wiki/Ralph_M._Steinman" title="Ralph M. Steinman">Ralph M. Steinman</a> (posthumously)</li> <li>2012: <a href="/wiki/John_Gurdon" title="John Gurdon">John Gurdon</a> / <a href="/wiki/Shinya_Yamanaka" title="Shinya Yamanaka">Shinya Yamanaka</a></li> <li>2013: <a href="/wiki/James_Rothman" title="James Rothman">James Rothman</a> / <a href="/wiki/Randy_Schekman" title="Randy Schekman">Randy Schekman</a> / <a href="/wiki/Thomas_C._S%C3%BCdhof" title="Thomas C. Südhof">Thomas C. Südhof</a></li> <li>2014: <a href="/wiki/John_O%27Keefe_(neuroscientist)" title="John O'Keefe (neuroscientist)">John O'Keefe</a> / <a href="/wiki/May-Britt_Moser" title="May-Britt Moser">May-Britt Moser</a> / <a href="/wiki/Edvard_Moser" title="Edvard Moser">Edvard Moser</a></li> <li>2015: <a href="/wiki/William_C._Campbell_(scientist)" title="William C. Campbell (scientist)">William C. Campbell</a> / <a href="/wiki/Satoshi_%C5%8Cmura" title="Satoshi Ōmura">Satoshi Ōmura</a> / <a href="/wiki/Tu_Youyou" title="Tu Youyou">Tu Youyou</a></li> <li>2016: <a href="/wiki/Yoshinori_Ohsumi" title="Yoshinori Ohsumi">Yoshinori Ohsumi</a></li> <li>2017: <a href="/wiki/Jeffrey_C._Hall" title="Jeffrey C. Hall">Jeffrey C. Hall</a> / <a href="/wiki/Michael_Rosbash" title="Michael Rosbash">Michael Rosbash</a> / <a href="/wiki/Michael_W._Young" title="Michael W. Young">Michael W. Young</a></li> <li>2018: <a href="/wiki/James_P._Allison" title="James P. Allison">James P. Allison</a> / <a href="/wiki/Tasuku_Honjo" title="Tasuku Honjo">Tasuku Honjo</a></li> <li>2019: <a href="/wiki/Gregg_L._Semenza" title="Gregg L. Semenza">Gregg L. Semenza</a> / <a href="/wiki/Peter_J._Ratcliffe" title="Peter J. Ratcliffe">Peter J. Ratcliffe</a> / <a href="/wiki/William_Kaelin_Jr." title="William Kaelin Jr.">William Kaelin Jr.</a></li> <li><a href="/wiki/2020_Nobel_Prize_in_Physiology_or_Medicine" title="2020 Nobel Prize in Physiology or Medicine">2020</a>: <a href="/wiki/Harvey_J._Alter" title="Harvey J. Alter">Harvey J. Alter</a> / <a href="/wiki/Michael_Houghton_(virologist)" class="mw-redirect" title="Michael Houghton (virologist)">Michael Houghton</a> / <a href="/wiki/Charles_M._Rice" title="Charles M. Rice">Charles M. Rice</a></li> <li><a href="/wiki/2021_Nobel_Prize_in_Physiology_or_Medicine" title="2021 Nobel Prize in Physiology or Medicine">2021</a>: <a href="/wiki/David_Julius" title="David Julius">David Julius</a> / <a href="/wiki/Ardem_Patapoutian" title="Ardem Patapoutian">Ardem Patapoutian</a></li> <li><a href="/wiki/2022_Nobel_Prize_in_Physiology_or_Medicine" title="2022 Nobel Prize in Physiology or Medicine">2022</a>: <a href="/wiki/Svante_P%C3%A4%C3%A4bo" title="Svante Pääbo">Svante Pääbo</a></li> <li><a href="/wiki/2023_Nobel_Prize_in_Physiology_or_Medicine" title="2023 Nobel Prize in Physiology or Medicine">2023</a>: <a href="/wiki/Katalin_Karik%C3%B3" title="Katalin Karikó">Katalin Karikó</a> / <a href="/wiki/Drew_Weissman" title="Drew Weissman">Drew Weissman</a></li> <li>2024: <a href="/wiki/Victor_Ambros" title="Victor Ambros">Victor Ambros</a> / <a href="/wiki/Gary_Ruvkun" title="Gary Ruvkun">Gary Ruvkun</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:r886047488">.mw-parser-output .nobold{font-weight:normal}</style></div><div role="navigation" class="navbox" aria-labelledby="2011_Nobel_Prize_laureates" style="padding:3px"><table class="nowraplinks hlist 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:2011_Nobel_Prize_winners" title="Template:2011 Nobel Prize winners"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:2011_Nobel_Prize_winners" title="Template talk:2011 Nobel Prize winners"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:2011_Nobel_Prize_winners" title="Special:EditPage/Template:2011 Nobel Prize winners"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="2011_Nobel_Prize_laureates" style="font-size:114%;margin:0 4em">2011 <a href="/wiki/Nobel_Prize" title="Nobel Prize">Nobel Prize</a> laureates</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Chemistry</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/Dan_Shechtman" title="Dan Shechtman">Dan Shechtman</a> (Israel)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Prize_in_Literature" title="Nobel Prize in Literature">Literature</a> (<a href="/wiki/2011_Nobel_Prize_in_Literature" title="2011 Nobel Prize in Literature">2011</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/Tomas_Transtr%C3%B6mer" title="Tomas Tranströmer">Tomas Tranströmer</a> (Sweden)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Peace_Prize" title="Nobel Peace Prize">Peace</a> (<a href="/wiki/2011_Nobel_Peace_Prize" title="2011 Nobel Peace Prize">2011</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/Ellen_Johnson_Sirleaf" title="Ellen Johnson Sirleaf">Ellen Johnson Sirleaf</a> (Liberia)</li> <li><a href="/wiki/Leymah_Gbowee" title="Leymah Gbowee">Leymah Gbowee</a> (Liberia)</li> <li><a href="/wiki/Tawakkol_Karman" title="Tawakkol Karman">Tawakkul Karman</a> (Yemen)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Prize_in_Physics" title="Nobel Prize in Physics">Physics</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/Saul_Perlmutter" title="Saul Perlmutter">Saul Perlmutter</a> (United States)</li> <li><a href="/wiki/Adam_Riess" title="Adam Riess">Adam Riess</a> (United States)</li> <li><a href="/wiki/Brian_Schmidt" title="Brian Schmidt">Brian P. Schmidt</a> (Australia/United States)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Physiology or Medicine</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 class="mw-selflink selflink">Bruce A. Beutler</a> (United States)</li> <li><a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules A. Hoffmann</a> (France)</li> <li><a href="/wiki/Ralph_M._Steinman" title="Ralph M. Steinman">Ralph M. Steinman</a> (Canada/United States)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nobel_Memorial_Prize_in_Economic_Sciences" title="Nobel Memorial Prize in Economic Sciences">Economic Sciences</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/Christopher_A._Sims" title="Christopher A. Sims">Christopher A. Sims</a> (United States)</li> <li><a href="/wiki/Thomas_J._Sargent" title="Thomas J. Sargent">Thomas J. Sargent</a> (United States)</li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <dl><dt><a href="/wiki/List_of_Nobel_laureates" title="List of Nobel laureates">Nobel Prize recipients</a></dt> <dd><a href="/wiki/Template:2006_Nobel_Prize_winners" title="Template:2006 Nobel Prize winners">2006</a></dd> <dd><a href="/wiki/Template:2007_Nobel_Prize_winners" title="Template:2007 Nobel Prize winners">2007</a></dd> <dd><a href="/wiki/Template:2008_Nobel_Prize_winners" title="Template:2008 Nobel Prize winners">2008</a></dd> <dd><a href="/wiki/Template:2009_Nobel_Prize_winners" title="Template:2009 Nobel Prize winners">2009</a></dd> <dd><a href="/wiki/Template:2010_Nobel_Prize_winners" title="Template:2010 Nobel Prize winners">2010</a></dd> <dd>2011 </dd> <dd><a href="/wiki/Template:2012_Nobel_Prize_winners" title="Template:2012 Nobel Prize winners">2012</a></dd> <dd><a href="/wiki/Template:2013_Nobel_Prize_winners" title="Template:2013 Nobel Prize winners">2013</a></dd> <dd><a href="/wiki/Template:2014_Nobel_Prize_winners" title="Template:2014 Nobel Prize winners">2014</a></dd> <dd><a href="/wiki/Template:2015_Nobel_Prize_winners" title="Template:2015 Nobel Prize winners">2015</a></dd> <dd><a href="/wiki/Template:2016_Nobel_Prize_winners" title="Template:2016 Nobel Prize winners">2016</a></dd></dl> </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="Shaw_Prize_laureates" 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="background:#d3d3d3;"><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:Shaw_Prize_laureates" title="Template:Shaw Prize laureates"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Shaw_Prize_laureates" title="Template talk:Shaw Prize laureates"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Shaw_Prize_laureates" title="Special:EditPage/Template:Shaw Prize laureates"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Shaw_Prize_laureates" style="font-size:114%;margin:0 4em"><a href="/wiki/Shaw_Prize" title="Shaw Prize">Shaw Prize</a> laureates</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#d3d3d3;">Astronomy</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/Jim_Peebles" title="Jim Peebles">Jim Peebles</a> (2004)</li> <li><a href="/wiki/Geoffrey_Marcy" title="Geoffrey Marcy">Geoffrey Marcy</a> and <a href="/wiki/Michel_Mayor" title="Michel Mayor">Michel Mayor</a> (2005)</li> <li><a href="/wiki/Saul_Perlmutter" title="Saul Perlmutter">Saul Perlmutter</a>, <a href="/wiki/Adam_Riess" title="Adam Riess">Adam Riess</a> and <a href="/wiki/Brian_Schmidt" title="Brian Schmidt">Brian Schmidt</a> (2006)</li> <li><a href="/wiki/Peter_Goldreich" title="Peter Goldreich">Peter Goldreich</a> (2007)</li> <li><a href="/wiki/Reinhard_Genzel" title="Reinhard Genzel">Reinhard Genzel</a> (2008)</li> <li><a href="/wiki/Frank_Shu" title="Frank Shu">Frank Shu</a> (2009)</li> <li><a href="/wiki/Charles_L._Bennett" title="Charles L. Bennett">Charles Bennett</a>, <a href="/wiki/Lyman_Page" title="Lyman Page">Lyman Page</a> and <a href="/wiki/David_Spergel" title="David Spergel">David Spergel</a> (2010)</li> <li><a href="/wiki/Enrico_Costa_(physicist)" title="Enrico Costa (physicist)">Enrico Costa</a> and <a href="/wiki/Gerald_J._Fishman" title="Gerald J. Fishman">Gerald Fishman</a> (2011)</li> <li><a href="/wiki/David_C._Jewitt" title="David C. Jewitt">David C. Jewitt</a> and <a href="/wiki/Jane_Luu" title="Jane Luu">Jane Luu</a> (2012)</li> <li><a href="/wiki/Steven_Balbus" title="Steven Balbus">Steven Balbus</a> and <a href="/wiki/John_F._Hawley" title="John F. Hawley">John F. Hawley</a> (2013)</li> <li><a href="/wiki/Daniel_Eisenstein" title="Daniel Eisenstein">Daniel Eisenstein</a>, <a href="/wiki/Shaun_Cole" title="Shaun Cole">Shaun Cole</a> and <a href="/wiki/John_A._Peacock" title="John A. Peacock">John A. Peacock</a> (2014)</li> <li><a href="/wiki/William_J._Borucki" title="William J. Borucki">William J. Borucki</a> (2015)</li> <li><a href="/wiki/Ronald_Drever" title="Ronald Drever">Ronald Drever</a>, <a href="/wiki/Kip_Thorne" title="Kip Thorne">Kip Thorne</a> and <a href="/wiki/Rainer_Weiss" title="Rainer Weiss">Rainer Weiss</a> (2016)</li> <li><a href="/wiki/Simon_White" title="Simon White">Simon White</a> (2017)</li> <li><a href="/wiki/Jean-Loup_Puget" title="Jean-Loup Puget">Jean-Loup Puget</a> (2018)</li> <li><a href="/wiki/Edward_C._Stone" title="Edward C. Stone">Edward C. Stone</a> (2019)</li> <li><a href="/wiki/Roger_Blandford" title="Roger Blandford">Roger Blandford</a> (2020)</li> <li><a href="/wiki/Victoria_Kaspi" title="Victoria Kaspi">Victoria Kaspi</a> and <a href="/wiki/Chryssa_Kouveliotou" title="Chryssa Kouveliotou">Chryssa Kouveliotou</a> (2021)</li> <li><a href="/wiki/Lennart_Lindegren" title="Lennart Lindegren">Lennart Lindegren</a> and <a href="/wiki/Michael_Perryman" title="Michael Perryman">Michael Perryman</a> (2022)</li> <li><a href="/wiki/Matthew_Bailes" title="Matthew Bailes">Matthew Bailes</a>, <a href="/wiki/Duncan_Lorimer" title="Duncan Lorimer">Duncan Lorimer</a> and <a href="/wiki/Maura_McLaughlin" title="Maura McLaughlin">Maura McLaughlin</a> (2023)</li> <li><a href="/wiki/Shrinivas_R._Kulkarni" class="mw-redirect" title="Shrinivas R. Kulkarni">Shrinivas R. Kulkarni</a> (2024)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#d3d3d3;">Life science and medicine</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/Stanley_Norman_Cohen" title="Stanley Norman Cohen">Stanley Norman Cohen</a>, <a href="/wiki/Herbert_Boyer" title="Herbert Boyer">Herbert Boyer</a>, <a href="/wiki/Yuet_Wai_Kan" title="Yuet Wai Kan">Yuet-Wai Kan</a> and <a href="/wiki/Richard_Doll" title="Richard Doll">Richard Doll</a> (2004)</li> <li><a href="/wiki/Michael_Berridge" title="Michael Berridge">Michael Berridge</a> (2005)</li> <li><a href="/wiki/Xiaodong_Wang_(biochemist)" title="Xiaodong Wang (biochemist)">Xiaodong Wang</a> (2006)</li> <li><a href="/wiki/Robert_Lefkowitz" title="Robert Lefkowitz">Robert Lefkowitz</a> (2007)</li> <li><a href="/wiki/Ian_Wilmut" title="Ian Wilmut">Ian Wilmut</a>, <a href="/wiki/Keith_Campbell_(biologist)" title="Keith Campbell (biologist)">Keith H. S. Campbell</a> and <a href="/wiki/Shinya_Yamanaka" title="Shinya Yamanaka">Shinya Yamanaka</a> (2008)</li> <li><a href="/wiki/Douglas_L._Coleman" title="Douglas L. Coleman">Douglas Coleman</a> and <a href="/wiki/Jeffrey_M._Friedman" title="Jeffrey M. Friedman">Jeffrey Friedman</a> (2009)</li> <li><a href="/wiki/David_Julius" title="David Julius">David Julius</a> (2010)</li> <li><a href="/wiki/Jules_A._Hoffmann" title="Jules A. Hoffmann">Jules Hoffmann</a>, <a href="/wiki/Ruslan_Medzhitov" title="Ruslan Medzhitov">Ruslan Medzhitov</a> and <a class="mw-selflink selflink">Bruce Beutler</a> (2011)</li> <li><a href="/wiki/Franz-Ulrich_Hartl" title="Franz-Ulrich Hartl">Franz-Ulrich Hartl</a> and <a href="/wiki/Arthur_L._Horwich" title="Arthur L. Horwich">Arthur L. Horwich</a> (2012)</li> <li><a href="/wiki/Jeffrey_C._Hall" title="Jeffrey C. Hall">Jeffrey C. Hall</a>, <a href="/wiki/Michael_Rosbash" title="Michael Rosbash">Michael Rosbash</a> and <a href="/wiki/Michael_W._Young" title="Michael W. Young">Michael W. Young</a> (2013)</li> <li><a href="/wiki/Kazutoshi_Mori" title="Kazutoshi Mori">Kazutoshi Mori</a> and <a href="/wiki/Peter_Walter" title="Peter Walter">Peter Walter</a> (2014)</li> <li><a href="/wiki/Bonnie_Bassler" title="Bonnie Bassler">Bonnie Bassler</a> and <a href="/wiki/Everett_Peter_Greenberg" title="Everett Peter Greenberg">Everett Peter Greenberg</a> (2015)</li> <li><a href="/wiki/Adrian_Bird" title="Adrian Bird">Adrian Bird</a> and <a href="/wiki/Huda_Zoghbi" title="Huda Zoghbi">Huda Zoghbi</a> (2016)</li> <li><a href="/wiki/Ian_R._Gibbons" title="Ian R. Gibbons">Ian R. Gibbons</a> and <a href="/wiki/Ronald_Vale" title="Ronald Vale">Ronald Vale</a> (2017)</li> <li><a href="/wiki/Mary-Claire_King" title="Mary-Claire King">Mary-Claire King</a> (2018)</li> <li><a href="/wiki/Maria_Jasin" title="Maria Jasin">Maria Jasin</a> (2019)</li> <li><a href="/wiki/Gero_Miesenb%C3%B6ck" title="Gero Miesenböck">Gero Miesenböck</a>, <a href="/wiki/Peter_Hegemann" title="Peter Hegemann">Peter Hegemann</a> and <a href="/wiki/Georg_Nagel" title="Georg Nagel">Georg Nagel</a> (2020)</li> <li><a href="/wiki/Scott_D._Emr" title="Scott D. Emr">Scott D. Emr</a> (2021)</li> <li><a href="/wiki/Paul_Negulescu" title="Paul Negulescu">Paul A. Negulescu</a> and <a href="/wiki/Michael_J._Welsh_(biologist)" title="Michael J. Welsh (biologist)">Michael J. Welsh</a> (2022)</li> <li><a href="/wiki/Patrick_Cramer" title="Patrick Cramer">Patrick Cramer</a> and <a href="/wiki/Eva_Nogales" title="Eva Nogales">Eva Nogales</a> (2023)</li> <li><a href="/wiki/Stuart_H._Orkin" class="mw-redirect" title="Stuart H. Orkin">Stuart H. Orkin</a> and <a href="/wiki/Swee_Lay_Thein" title="Swee Lay Thein">Swee Lay Thein</a> (2024)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#d3d3d3;">Mathematical science</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/Shiing-Shen_Chern" title="Shiing-Shen Chern">Shiing-Shen Chern</a> (2004)</li> <li><a href="/wiki/Andrew_Wiles" title="Andrew Wiles">Andrew Wiles</a> (2005)</li> <li><a href="/wiki/David_Mumford" title="David Mumford">David Mumford</a> and <a href="/wiki/Wu_Wenjun" title="Wu Wenjun">Wentsun Wu</a> (2006)</li> <li><a href="/wiki/Robert_Langlands" title="Robert Langlands">Robert Langlands</a> and <a href="/wiki/Richard_Taylor_(mathematician)" title="Richard Taylor (mathematician)">Richard Taylor</a> (2007)</li> <li><a href="/wiki/Vladimir_Arnold" title="Vladimir Arnold">Vladimir Arnold</a> and <a href="/wiki/Ludvig_Faddeev" title="Ludvig Faddeev">Ludwig Faddeev</a> (2008)</li> <li><a href="/wiki/Simon_Donaldson" title="Simon Donaldson">Simon Donaldson</a> and <a href="/wiki/Clifford_Taubes" title="Clifford Taubes">Clifford Taubes</a> (2009)</li> <li><a href="/wiki/Jean_Bourgain" title="Jean Bourgain">Jean Bourgain</a> (2010)</li> <li><a href="/wiki/Demetrios_Christodoulou" title="Demetrios Christodoulou">Demetrios Christodoulou</a> and <a href="/wiki/Richard_S._Hamilton" title="Richard S. Hamilton">Richard S. Hamilton</a> (2011)</li> <li><a href="/wiki/Maxim_Kontsevich" title="Maxim Kontsevich">Maxim Kontsevich</a> (2012)</li> <li><a href="/wiki/David_Donoho" title="David Donoho">David Donoho</a> (2013)</li> <li><a href="/wiki/George_Lusztig" title="George Lusztig">George Lusztig</a> (2014)</li> <li><a href="/wiki/Gerd_Faltings" title="Gerd Faltings">Gerd Faltings</a> and <a href="/wiki/Henryk_Iwaniec" title="Henryk Iwaniec">Henryk Iwaniec</a> (2015)</li> <li><a href="/wiki/Nigel_Hitchin" title="Nigel Hitchin">Nigel Hitchin</a> (2016)</li> <li><a href="/wiki/J%C3%A1nos_Koll%C3%A1r" title="János Kollár">János Kollár</a> and <a href="/wiki/Claire_Voisin" title="Claire Voisin">Claire Voisin</a> (2017)</li> <li><a href="/wiki/Luis_Caffarelli" title="Luis Caffarelli">Luis Caffarelli</a> (2018)</li> <li><a href="/wiki/Michel_Talagrand" title="Michel Talagrand">Michel Talagrand</a> (2019)</li> <li><a href="/wiki/Alexander_Beilinson" title="Alexander Beilinson">Alexander Beilinson</a> and <a href="/wiki/David_Kazhdan" title="David Kazhdan">David Kazhdan</a> (2020)</li> <li><a href="/wiki/Jean-Michel_Bismut" title="Jean-Michel Bismut">Jean-Michel Bismut</a> and <a href="/wiki/Jeff_Cheeger" title="Jeff Cheeger">Jeff Cheeger</a> (2021)</li> <li><a href="/wiki/Noga_Alon" title="Noga Alon">Noga Alon</a> and <a href="/wiki/Ehud_Hrushovski" title="Ehud Hrushovski">Ehud Hrushovski</a> (2022)</li> <li><a href="/wiki/Vladimir_Drinfeld" title="Vladimir Drinfeld">Vladimir Drinfeld</a> and <a href="/wiki/Shing-Tung_Yau" title="Shing-Tung Yau">Shing-Tung Yau</a> (2023)</li> <li><a href="/wiki/Peter_Sarnak" title="Peter Sarnak">Peter Sarnak</a> (2024)</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 authority-control" 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href="https://id.oclc.org/worldcat/entity/E39PBJjMG77xpYkctFYmGyj9jC">WorldCat</a></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</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 rel="nofollow" class="external text" href="https://d-nb.info/gnd/1031955119">Germany</a></li><li><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/n89143955">United States</a></li><li><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb15939506n">France</a></li><li><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb15939506n">BnF data</a></li><li><a rel="nofollow" class="external text" href="http://data.bibliotheken.nl/id/thes/p101319622">Netherlands</a></li><li><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007335604105171">Israel</a></li><li><a rel="nofollow" class="external text" href="https://opac.kbr.be/LIBRARY/doc/AUTHORITY/14056528">Belgium</a></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Academics</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 rel="nofollow" class="external text" href="https://orcid.org/0000-0002-3639-246X">ORCID</a></li><li><a rel="nofollow" class="external text" href="https://www.leopoldina.org/mitgliederverzeichnis/mitglieder/member/Member/show/bruce-beutler/">Leopoldina</a></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">People</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 rel="nofollow" class="external text" href="https://www.deutsche-digitale-bibliothek.de/person/gnd/1031955119">DDB</a></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><a rel="nofollow" class="external text" href="https://www.idref.fr/06902572X">IdRef</a></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Bruce_Beutler&oldid=1250047553">https://en.wikipedia.org/w/index.php?title=Bruce_Beutler&oldid=1250047553</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a 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