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Stem cell - Wikipedia
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class="vector-toc-link" href="#Identification"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Identification</span> </div> </a> <ul id="toc-Identification-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Embryonic" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Embryonic"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Embryonic</span> </div> </a> <button aria-controls="toc-Embryonic-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Embryonic subsection</span> </button> <ul id="toc-Embryonic-sublist" class="vector-toc-list"> <li id="toc-Mesenchymal_stem_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mesenchymal_stem_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Mesenchymal stem cells</span> </div> </a> <ul id="toc-Mesenchymal_stem_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cell_cycle_control" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cell_cycle_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Cell cycle control</span> </div> </a> <ul id="toc-Cell_cycle_control-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Fetal" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Fetal"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Fetal</span> </div> </a> <ul id="toc-Fetal-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Adult" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Adult"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Adult</span> </div> </a> <button aria-controls="toc-Adult-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Adult subsection</span> </button> <ul id="toc-Adult-sublist" class="vector-toc-list"> <li id="toc-Hematopoietic_stem_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Hematopoietic_stem_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Hematopoietic stem cells</span> </div> </a> <ul id="toc-Hematopoietic_stem_cells-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Amniotic" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Amniotic"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Amniotic</span> </div> </a> <ul id="toc-Amniotic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Induced_pluripotent" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Induced_pluripotent"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Induced pluripotent</span> </div> </a> <button aria-controls="toc-Induced_pluripotent-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Induced pluripotent subsection</span> </button> <ul id="toc-Induced_pluripotent-sublist" class="vector-toc-list"> <li id="toc-Cell_cycle_control_2" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cell_cycle_control_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Cell cycle control</span> </div> </a> <ul id="toc-Cell_cycle_control_2-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Lineage" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Lineage"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Lineage</span> </div> </a> <ul id="toc-Lineage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Therapies" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Therapies"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Therapies</span> </div> </a> <button aria-controls="toc-Therapies-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Therapies subsection</span> </button> <ul id="toc-Therapies-sublist" class="vector-toc-list"> <li id="toc-Advantages" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Advantages"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.1</span> <span>Advantages</span> </div> </a> <ul id="toc-Advantages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Disadvantages" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Disadvantages"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.2</span> <span>Disadvantages</span> </div> </a> <ul id="toc-Disadvantages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stem_cell_tourism" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Stem_cell_tourism"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.3</span> <span>Stem cell tourism</span> </div> </a> <ul id="toc-Stem_cell_tourism-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Research" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Research"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Research</span> </div> </a> <button aria-controls="toc-Research-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Research subsection</span> </button> <ul id="toc-Research-sublist" class="vector-toc-list"> <li id="toc-Conditions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Conditions"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Conditions</span> </div> </a> <ul id="toc-Conditions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Production" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Production"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.2</span> <span>Production</span> </div> </a> <ul id="toc-Production-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Organoids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Organoids"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.3</span> <span>Organoids</span> </div> </a> <ul id="toc-Organoids-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Risks" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Risks"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.4</span> <span>Risks</span> </div> </a> <ul id="toc-Risks-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dormancy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dormancy"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.5</span> <span>Dormancy</span> </div> </a> <ul id="toc-Dormancy-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">14</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Stem cell</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 85 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-85" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">85 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Stamsel" title="Stamsel – Afrikaans" lang="af" hreflang="af" data-title="Stamsel" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AE%D9%84%D9%8A%D8%A9_%D8%AC%D8%B0%D8%B9%D9%8A%D8%A9" title="خلية جذعية – Arabic" lang="ar" hreflang="ar" data-title="خلية جذعية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/K%C3%B6k_h%C3%BCceyr%C9%99l%C9%99r" title="Kök hüceyrələr – Azerbaijani" lang="az" hreflang="az" data-title="Kök hüceyrələr" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-azb mw-list-item"><a href="https://azb.wikipedia.org/wiki/%DA%A9%D8%A4%DA%A9_%D8%B3%D9%84%D9%84%D9%88%D9%84" title="کؤک سللول – South Azerbaijani" lang="azb" hreflang="azb" data-title="کؤک سللول" data-language-autonym="تۆرکجه" data-language-local-name="South Azerbaijani" class="interlanguage-link-target"><span>تۆرکجه</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%AE%E0%A6%BE%E0%A6%A4%E0%A7%83%E0%A6%95%E0%A7%8B%E0%A6%B7" title="মাতৃকোষ – Bangla" lang="bn" hreflang="bn" data-title="মাতৃকোষ" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/K%C3%A0n-s%C3%A8-pau" title="Kàn-sè-pau – Minnan" lang="nan" hreflang="nan" data-title="Kàn-sè-pau" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%A1%D1%82%D0%B2%D0%B0%D0%BB%D0%BE%D0%B2%D1%8B%D1%8F_%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D1%96" title="Стваловыя клеткі – Belarusian" lang="be" hreflang="be" data-title="Стваловыя клеткі" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A1%D1%82%D0%B2%D0%BE%D0%BB%D0%BE%D0%B2%D0%B0_%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="Стволова клетка – Bulgarian" lang="bg" hreflang="bg" data-title="Стволова клетка" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Mati%C4%8Dna_%C4%87elija" title="Matična ćelija – Bosnian" lang="bs" hreflang="bs" data-title="Matična ćelija" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/C%C3%A8l%C2%B7lula_mare" title="Cèl·lula mare – Catalan" lang="ca" hreflang="ca" data-title="Cèl·lula mare" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Kmenov%C3%A1_bu%C5%88ka" title="Kmenová buňka – Czech" lang="cs" hreflang="cs" data-title="Kmenová buňka" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Stamcelle" title="Stamcelle – Danish" lang="da" hreflang="da" data-title="Stamcelle" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-se mw-list-item"><a href="https://se.wikipedia.org/wiki/Vuo%C4%91%C4%91osealla" title="Vuođđosealla – Northern Sami" lang="se" hreflang="se" data-title="Vuođđosealla" data-language-autonym="Davvisámegiella" data-language-local-name="Northern Sami" class="interlanguage-link-target"><span>Davvisámegiella</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Stammzelle" title="Stammzelle – German" lang="de" hreflang="de" data-title="Stammzelle" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/T%C3%BCvirakk" title="Tüvirakk – Estonian" lang="et" hreflang="et" data-title="Tüvirakk" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%92%CE%BB%CE%B1%CF%83%CF%84%CE%BF%CE%BA%CF%8D%CF%84%CF%84%CE%B1%CF%81%CE%BF" title="Βλαστοκύτταρο – Greek" lang="el" hreflang="el" data-title="Βλαστοκύτταρο" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/C%C3%A9lula_madre" title="Célula madre – Spanish" lang="es" hreflang="es" data-title="Célula madre" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Pra%C4%89elo" title="Praĉelo – Esperanto" lang="eo" hreflang="eo" data-title="Praĉelo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Zelula_ama" title="Zelula ama – Basque" lang="eu" hreflang="eu" data-title="Zelula ama" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D9%84%D9%88%D9%84_%D8%A8%D9%86%DB%8C%D8%A7%D8%AF%DB%8C" title="سلول بنیادی – Persian" lang="fa" hreflang="fa" data-title="سلول بنیادی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Cellule_souche" title="Cellule souche – French" lang="fr" hreflang="fr" data-title="Cellule souche" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Gaschill" title="Gaschill – Irish" lang="ga" hreflang="ga" data-title="Gaschill" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/C%C3%A9lula_nai" title="Célula nai – Galician" lang="gl" hreflang="gl" data-title="Célula nai" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/Kon-se-p%C3%A2u" title="Kon-se-pâu – Hakka Chinese" lang="hak" hreflang="hak" data-title="Kon-se-pâu" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target"><span>客家語 / Hak-kâ-ngî</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%A4%84%EA%B8%B0%EC%84%B8%ED%8F%AC" title="줄기세포 – Korean" lang="ko" hreflang="ko" data-title="줄기세포" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%91%D5%B8%D5%B2%D5%B8%D6%82%D5%B6%D5%A1%D5%B5%D5%AB%D5%B6_%D5%A2%D5%BB%D5%AB%D5%BB%D5%B6%D5%A5%D6%80" title="Ցողունային բջիջներ – Armenian" lang="hy" hreflang="hy" data-title="Ցողունային բջիջներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B8%E0%A5%8D%E0%A4%9F%E0%A5%87%E0%A4%AE_%E0%A4%95%E0%A5%8B%E0%A4%B6%E0%A4%BF%E0%A4%95%E0%A4%BE" title="स्टेम कोशिका – Hindi" lang="hi" hreflang="hi" data-title="स्टेम कोशिका" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Mati%C4%8Dne_stanice" title="Matične stanice – Croatian" lang="hr" hreflang="hr" data-title="Matične stanice" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Sel_punca" title="Sel punca – Indonesian" lang="id" hreflang="id" data-title="Sel punca" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Cellula_originator" title="Cellula originator – Interlingua" lang="ia" hreflang="ia" data-title="Cellula originator" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target"><span>Interlingua</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Stofnfruma" title="Stofnfruma – Icelandic" lang="is" hreflang="is" data-title="Stofnfruma" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Cellula_staminale" title="Cellula staminale – Italian" lang="it" hreflang="it" data-title="Cellula staminale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%AA%D7%90_%D7%92%D7%96%D7%A2" title="תא גזע – Hebrew" lang="he" hreflang="he" data-title="תא גזע" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Sel_Punca" title="Sel Punca – Javanese" lang="jv" hreflang="jv" data-title="Sel Punca" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%86%E0%B2%95%E0%B2%B0_%E0%B2%95%E0%B3%8B%E0%B2%B6" title="ಆಕರ ಕೋಶ – Kannada" lang="kn" hreflang="kn" data-title="ಆಕರ ಕೋಶ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A6%E1%83%94%E1%83%A0%E1%83%9D%E1%83%95%E1%83%90%E1%83%9C%E1%83%98_%E1%83%A3%E1%83%AF%E1%83%A0%E1%83%94%E1%83%93%E1%83%98" title="ღეროვანი უჯრედი – Georgian" lang="ka" hreflang="ka" data-title="ღეროვანი უჯრედი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%91%D0%B0%D2%93%D0%B0%D0%BD%D0%B0%D0%BB%D1%8B_%D0%B6%D0%B0%D1%81%D1%83%D1%88%D0%B0" title="Бағаналы жасуша – Kazakh" lang="kk" hreflang="kk" data-title="Бағаналы жасуша" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Cellula_praecursoria" title="Cellula praecursoria – Latin" lang="la" hreflang="la" data-title="Cellula praecursoria" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Cilmes_%C5%A1%C5%ABna" title="Cilmes šūna – Latvian" lang="lv" hreflang="lv" data-title="Cilmes šūna" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Kamienin%C4%97_l%C4%85stel%C4%97" title="Kamieninė ląstelė – Lithuanian" lang="lt" hreflang="lt" data-title="Kamieninė ląstelė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/%C5%90ssejt" title="Őssejt – Hungarian" lang="hu" hreflang="hu" data-title="Őssejt" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9C%D0%B0%D1%82%D0%B8%D1%87%D0%BD%D0%B0_%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="Матична клетка – Macedonian" lang="mk" hreflang="mk" data-title="Матична клетка" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B5%E0%B4%BF%E0%B4%A4%E0%B5%8D%E0%B4%A4%E0%B5%81%E0%B4%95%E0%B5%8B%E0%B4%B6%E0%B4%99%E0%B5%8D%E0%B4%99%E0%B5%BE" title="വിത്തുകോശങ്ങൾ – Malayalam" lang="ml" hreflang="ml" data-title="വിത്തുകോശങ്ങൾ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%AE%E0%A5%82%E0%A4%B2%E0%A4%AA%E0%A5%87%E0%A4%B6%E0%A5%80" title="मूलपेशी – Marathi" lang="mr" hreflang="mr" data-title="मूलपेशी" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target"><span>मराठी</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Sel_dasar" title="Sel dasar – Malay" lang="ms" hreflang="ms" data-title="Sel dasar" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D2%AE%D2%AF%D0%B4%D1%8D%D0%BB_%D1%8D%D1%81" title="Үүдэл эс – Mongolian" lang="mn" hreflang="mn" data-title="Үүдэл эс" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%95%E1%80%84%E1%80%BA%E1%80%99%E1%80%86%E1%80%B2%E1%80%9C%E1%80%BA" title="ပင်မဆဲလ် – Burmese" lang="my" hreflang="my" data-title="ပင်မဆဲလ်" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target"><span>မြန်မာဘာသာ</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Stamcel" title="Stamcel – Dutch" lang="nl" hreflang="nl" data-title="Stamcel" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E5%B9%B9%E7%B4%B0%E8%83%9E" title="幹細胞 – Japanese" lang="ja" hreflang="ja" data-title="幹細胞" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Stamceller" title="Stamceller – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Stamceller" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Stamcelle" title="Stamcelle – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Stamcelle" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%B8%E0%A8%9F%E0%A9%88%E0%A9%B1%E0%A8%AE_%E0%A8%B8%E0%A9%88%E0%A9%B1%E0%A8%B2" title="ਸਟੈੱਮ ਸੈੱਲ – Punjabi" lang="pa" hreflang="pa" data-title="ਸਟੈੱਮ ਸੈੱਲ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target"><span>ਪੰਜਾਬੀ</span></a></li><li class="interlanguage-link interwiki-blk mw-list-item"><a href="https://blk.wikipedia.org/wiki/%E1%80%9E%E1%80%B2%E1%80%9C%E1%80%BA%E1%80%99%E1%80%BD%E1%80%B0%E1%80%B8%E1%80%94%E1%80%99%E1%80%BA%E1%80%B8" title="သဲလ်မွူးနမ်း – Pa'O" lang="blk" hreflang="blk" data-title="သဲလ်မွူးနမ်း" data-language-autonym="ပအိုဝ်ႏဘာႏသာႏ" data-language-local-name="Pa'O" class="interlanguage-link-target"><span>ပအိုဝ်ႏဘာႏသာႏ</span></a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%A7%D8%B5%D9%84%D9%8A_%D8%AD%D8%AC%D8%B1%D9%87" title="اصلي حجره – Pashto" lang="ps" hreflang="ps" data-title="اصلي حجره" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-km mw-list-item"><a href="https://km.wikipedia.org/wiki/%E1%9E%80%E1%9F%84%E1%9E%9F%E1%9E%B7%E1%9E%80%E1%9E%B6%E2%80%8B%E1%9E%8A%E1%9E%BE%E1%9E%98" title="កោសិកាដើម – Khmer" lang="km" hreflang="km" data-title="កោសិកាដើម" data-language-autonym="ភាសាខ្មែរ" data-language-local-name="Khmer" class="interlanguage-link-target"><span>ភាសាខ្មែរ</span></a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Stammzell" title="Stammzell – Low German" lang="nds" hreflang="nds" data-title="Stammzell" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target"><span>Plattdüütsch</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Kom%C3%B3rki_macierzyste" title="Komórki macierzyste – Polish" lang="pl" hreflang="pl" data-title="Komórki macierzyste" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/C%C3%A9lula_estaminal" title="Célula estaminal – Portuguese" lang="pt" hreflang="pt" data-title="Célula estaminal" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Celule_stem" title="Celule stem – Romanian" lang="ro" hreflang="ro" data-title="Celule stem" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A1%D1%82%D0%B2%D0%BE%D0%BB%D0%BE%D0%B2%D1%8B%D0%B5_%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B8" title="Стволовые клетки – Russian" lang="ru" hreflang="ru" data-title="Стволовые клетки" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Stem_cell" title="Stem cell – Scots" lang="sco" hreflang="sco" data-title="Stem cell" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target"><span>Scots</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Qelizat_staminale" title="Qelizat staminale – Albanian" lang="sq" hreflang="sq" data-title="Qelizat staminale" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Stem_cell" title="Stem cell – Simple English" lang="en-simple" hreflang="en-simple" data-title="Stem cell" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Kme%C5%88ov%C3%A1_bunka" title="Kmeňová bunka – Slovak" lang="sk" hreflang="sk" data-title="Kmeňová bunka" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mati%C4%8Dna_celica" title="Matična celica – Slovenian" lang="sl" hreflang="sl" data-title="Matična celica" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AE%D8%A7%D9%86%DB%95%DB%8C_%D8%A8%D9%86%DB%95%DA%95%DB%95%D8%AA%DB%8C" title="خانەی بنەڕەتی – Central Kurdish" lang="ckb" hreflang="ckb" data-title="خانەی بنەڕەتی" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%98%D0%B7%D0%B2%D0%BE%D1%80%D0%BD%D0%B0_%D1%9B%D0%B5%D0%BB%D0%B8%D1%98%D0%B0" title="Изворна ћелија – Serbian" lang="sr" hreflang="sr" data-title="Изворна ћелија" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Mati%C4%8Dna_%C4%87elija" title="Matična ćelija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Matična ćelija" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/S%C3%A9l_punca" title="Sél punca – Sundanese" lang="su" hreflang="su" data-title="Sél punca" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Kantasolu" title="Kantasolu – Finnish" lang="fi" hreflang="fi" data-title="Kantasolu" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Stamcell" title="Stamcell – Swedish" lang="sv" hreflang="sv" data-title="Stamcell" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%95%E0%AF%81%E0%AE%B0%E0%AF%81%E0%AE%A4%E0%AF%8D%E0%AE%A4%E0%AE%A3%E0%AF%81" title="குருத்தணு – Tamil" lang="ta" hreflang="ta" data-title="குருத்தணு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-te mw-list-item"><a href="https://te.wikipedia.org/wiki/%E0%B0%AE%E0%B1%82%E0%B0%B2_%E0%B0%95%E0%B0%A3%E0%B0%82" title="మూల కణం – Telugu" lang="te" hreflang="te" data-title="మూల కణం" data-language-autonym="తెలుగు" data-language-local-name="Telugu" class="interlanguage-link-target"><span>తెలుగు</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B9%80%E0%B8%8B%E0%B8%A5%E0%B8%A5%E0%B9%8C%E0%B8%95%E0%B9%89%E0%B8%99%E0%B8%81%E0%B8%B3%E0%B9%80%E0%B8%99%E0%B8%B4%E0%B8%94" title="เซลล์ต้นกำเนิด – Thai" lang="th" hreflang="th" data-title="เซลล์ต้นกำเนิด" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/K%C3%B6k_h%C3%BCcre" title="Kök hücre – Turkish" lang="tr" hreflang="tr" data-title="Kök hücre" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D1%82%D0%BE%D0%B2%D0%B1%D1%83%D1%80%D0%BE%D0%B2%D1%96_%D0%BA%D0%BB%D1%96%D1%82%D0%B8%D0%BD%D0%B8" title="Стовбурові клітини – Ukrainian" lang="uk" 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class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Undifferentiated biological cells that can differentiate into specialized cells</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about the cell type. For the medical therapy, see <a href="/wiki/Stem-cell_therapy" title="Stem-cell therapy">stem-cell therapy</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Stem cells" and "Stem cell research" redirect here. For the journals, see <a href="/wiki/Stem_Cells_(journal)" title="Stem Cells (journal)"><i>Stem Cells</i> (journal)</a> and <a href="/wiki/Stem_Cell_Research_(journal)" title="Stem Cell Research (journal)"><i>Stem Cell Research</i> (journal)</a>.</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:thistle">Stem cell</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:MSC_high_magnification.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/MSC_high_magnification.jpg/250px-MSC_high_magnification.jpg" decoding="async" width="250" height="333" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/MSC_high_magnification.jpg/375px-MSC_high_magnification.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/MSC_high_magnification.jpg/500px-MSC_high_magnification.jpg 2x" data-file-width="1803" data-file-height="2400" /></a></span><div class="infobox-caption"><a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">Transmission</a> <a href="/wiki/Electron_micrograph" class="mw-redirect" title="Electron micrograph">electron micrograph</a> of a <a href="/wiki/Mesenchymal_stem_cell" title="Mesenchymal stem cell">mesenchymal stem cell</a> displaying typical <a href="/wiki/Ultrastructural" class="mw-redirect" title="Ultrastructural">ultrastructural</a> characteristics</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>cellula praecursoria</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D013234">D013234</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Histologica" title="Terminologia Histologica">TH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/PDF/TH%20Chapter%20H1.00.pdf">H1.00.01.0.00028, H2.00.01.0.00001 </a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma63368">63368</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terminology" title="Anatomical terminology">Anatomical terminology</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q48196" class="extiw" title="d:Q48196">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>In <a href="/wiki/Multicellular_organisms" class="mw-redirect" title="Multicellular organisms">multicellular organisms</a>, <b>stem cells</b> are <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">undifferentiated</a> or partially differentiated <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a> that can change into various <a href="/wiki/Types_of_cells" class="mw-redirect" title="Types of cells">types of cells</a> and <a href="/wiki/Cell_proliferation" title="Cell proliferation">proliferate</a> indefinitely to produce more of the same stem cell. They are the earliest type of cell in a <a href="/wiki/Cell_lineage" title="Cell lineage">cell lineage</a>.<sup id="cite_ref-:7_1-0" class="reference"><a href="#cite_note-:7-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> They are found in both <a href="/wiki/Embryo" title="Embryo">embryonic</a> and adult organisms, but they have slightly different properties in each. They are usually distinguished from <a href="/wiki/Progenitor_cells" class="mw-redirect" title="Progenitor cells">progenitor cells</a>, which cannot divide indefinitely, and <a href="/wiki/Precursor_cell" title="Precursor cell">precursor</a> or blast cells, which are usually committed to differentiating into one cell type. </p><p>In <a href="/wiki/Mammals" class="mw-redirect" title="Mammals">mammals</a>, roughly 50 to 150 cells make up the <a href="/wiki/Inner_cell_mass" title="Inner cell mass">inner cell mass</a> during the <a href="/wiki/Blastocyst" title="Blastocyst">blastocyst</a> stage of <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a>, around days 5–14. These have stem-cell capability. <i><a href="/wiki/In_vivo" title="In vivo">In vivo</a></i>, they eventually differentiate into all of the body's cell types (making them <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a>). This process starts with the differentiation into the three <a href="/wiki/Germ_layers" class="mw-redirect" title="Germ layers">germ layers</a> – the <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a>, <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a> and <a href="/wiki/Endoderm" title="Endoderm">endoderm</a> – at the <a href="/wiki/Gastrulation" title="Gastrulation">gastrulation</a> stage. However, when they are isolated and <a href="/wiki/Cell_culture" title="Cell culture">cultured</a> <i><a href="/wiki/In_vitro" title="In vitro">in vitro</a></i>, they can be kept in the stem-cell stage and are known as <a href="/wiki/Embryonic_stem_cells" class="mw-redirect" title="Embryonic stem cells">embryonic stem cells</a> (ESCs). </p><p><a href="/wiki/Adult_stem_cells" class="mw-redirect" title="Adult stem cells">Adult stem cells</a> are found in a few select locations in the body, known as <a href="/wiki/Stem-cell_niche" title="Stem-cell niche">niches</a>, such as those in the <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a> or <a href="/wiki/Gonads" class="mw-redirect" title="Gonads">gonads</a>. They exist to replenish rapidly lost cell types and are <a href="/wiki/Multipotent" class="mw-redirect" title="Multipotent">multipotent</a> or unipotent, meaning they only differentiate into a few cell types or one type of cell. In mammals, they include, among others, <a href="/wiki/Hematopoietic_stem_cells" class="mw-redirect" title="Hematopoietic stem cells">hematopoietic stem cells</a>, which replenish blood and immune cells, <a href="/wiki/Stratum_basale" title="Stratum basale">basal cells</a>, which maintain the skin <a href="/wiki/Epithelium" title="Epithelium">epithelium</a>, and <a href="/wiki/Mesenchymal_stem_cells" class="mw-redirect" title="Mesenchymal stem cells">mesenchymal stem cells</a>, which maintain bone, <a href="/wiki/Cartilage" title="Cartilage">cartilage</a>, muscle and fat cells. Adult stem cells are a small minority of cells; they are vastly outnumbered by the progenitor cells and terminally differentiated cells that they differentiate into.<sup id="cite_ref-:7_1-1" class="reference"><a href="#cite_note-:7-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Research into stem cells grew out of findings by Canadian biologists <a href="/wiki/Ernest_McCulloch" title="Ernest McCulloch">Ernest McCulloch</a>, <a href="/wiki/James_Till" title="James Till">James Till</a> and Andrew J. Becker at the <a href="/wiki/University_of_Toronto" title="University of Toronto">University of Toronto</a> and the Ontario Cancer Institute in the 1960s.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> As of 2016<sup class="plainlinks noexcerpt noprint asof-tag update" style="display:none;"><a class="external text" href="https://en.wikipedia.org/w/index.php?title=Stem_cell&action=edit">[update]</a></sup>, the only established <a href="/wiki/Stem-cell_therapy" title="Stem-cell therapy">medical therapy using stem cells</a> is <a href="/wiki/Hematopoietic_stem_cell_transplantation" title="Hematopoietic stem cell transplantation">hematopoietic stem cell transplantation</a>,<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> first performed in 1958 by French oncologist <a href="/wiki/Georges_Math%C3%A9" title="Georges Mathé">Georges Mathé</a>. Since 1998 however, it has been possible to culture and differentiate human embryonic stem cells (in <a href="/wiki/Stem-cell_lines" class="mw-redirect" title="Stem-cell lines">stem-cell lines</a>). The process of isolating these cells has been <a href="/wiki/Stem_cell_controversy" title="Stem cell controversy">controversial</a>, because it typically results in the destruction of the embryo. Sources for isolating ESCs have been <a href="/wiki/Stem_cell_laws" title="Stem cell laws">restricted</a> in some European countries and Canada, but others such as the UK and China have promoted the research.<sup id="cite_ref-Pew_Intl_5-0" class="reference"><a href="#cite_note-Pew_Intl-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Somatic_cell_nuclear_transfer" title="Somatic cell nuclear transfer">Somatic cell nuclear transfer</a> is a <a href="/wiki/Cloning" title="Cloning">cloning</a> method that can be used to create a cloned embryo for the use of its embryonic stem cells in stem cell therapy.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In 2006, a Japanese team led by <a href="/wiki/Shinya_Yamanaka" title="Shinya Yamanaka">Shinya Yamanaka</a> discovered a method to convert mature body cells back into stem cells. These were termed <a href="/wiki/Induced_pluripotent_stem_cells" class="mw-redirect" title="Induced pluripotent stem cells">induced pluripotent stem cells</a> (iPSCs).<sup id="cite_ref-:6_7-0" class="reference"><a href="#cite_note-:6-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term <i>stem cell</i> was coined by <a href="/wiki/Theodor_Boveri" title="Theodor Boveri">Theodor Boveri</a> and <a href="/wiki/Valentin_Haecker" title="Valentin Haecker">Valentin Haecker</a> in late 19th century.<sup id="cite_ref-origin_8-0" class="reference"><a href="#cite_note-origin-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Pioneering works in theory of blood stem cell were conducted in the beginning of 20th century by <a href="/wiki/Artur_Pappenheim" title="Artur Pappenheim">Artur Pappenheim</a>, <a href="/wiki/Alexander_A._Maximow" title="Alexander A. Maximow">Alexander A. Maximow</a>, <a href="/wiki/Franz_Ernst_Christian_Neumann" class="mw-redirect" title="Franz Ernst Christian Neumann">Franz Ernst Christian Neumann</a>.<sup id="cite_ref-origin_8-1" class="reference"><a href="#cite_note-origin-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>The key properties of a stem cell were first defined by <a href="/wiki/Ernest_McCulloch" title="Ernest McCulloch">Ernest McCulloch</a> and <a href="/wiki/James_Till" title="James Till">James Till</a> at the University of Toronto and the Ontario Cancer Institute in the early 1960s. They discovered the blood-forming stem cell, the <a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">hematopoietic stem cell</a> (HSC), through their pioneering work in mice. McCulloch and Till began a series of experiments in which bone marrow cells were injected into irradiated mice. They observed lumps in the spleens of the mice that were linearly proportional to the number of bone marrow cells injected. They hypothesized that each lump (colony) was a clone arising from a single marrow cell (stem cell). In subsequent work, McCulloch and Till, joined by graduate student <a href="/w/index.php?title=Andrew_John_Becker&action=edit&redlink=1" class="new" title="Andrew John Becker (page does not exist)">Andrew John Becker</a> and senior scientist <a href="/wiki/Louis_Siminovitch" title="Louis Siminovitch">Louis Siminovitch</a>, confirmed that each lump did in fact arise from a single cell. Their results were published in <i>Nature</i> in 1963. In that same year, Siminovitch was a lead investigator for studies that found colony-forming cells were capable of self-renewal, which is a key defining property of stem cells that Till and McCulloch had theorized.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>The first therapy using stem cells was a <a href="/wiki/Bone_marrow_transplant" class="mw-redirect" title="Bone marrow transplant">bone marrow transplant</a> performed by French oncologist <a href="/wiki/Georges_Math%C3%A9" title="Georges Mathé">Georges Mathé</a> in 1956 on five workers at the <a href="/wiki/Vin%C4%8Da_Nuclear_Institute" title="Vinča Nuclear Institute">Vinča Nuclear Institute</a> in <a href="/wiki/Socialist_Federal_Republic_of_Yugoslavia" title="Socialist Federal Republic of Yugoslavia">Yugoslavia</a> who had been affected by a <a href="/wiki/Criticality_accident" title="Criticality accident">criticality accident</a>. The workers all survived.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>In 1981, embryonic stem (ES) cells were first isolated and successfully cultured using mouse blastocysts by British biologists <a href="/wiki/Martin_Evans" title="Martin Evans">Martin Evans</a> and <a href="/wiki/Matthew_Kaufman" title="Matthew Kaufman">Matthew Kaufman</a>. This allowed the formation of murine genetic models, a system in which the genes of mice are deleted or altered in order to study their function in pathology. In 1991, a process that allowed the human stem cell to be isolated was patented by Ann Tsukamoto. By 1998, human embryonic stem cells were first isolated by American biologist <a href="/wiki/James_Thomson_(cell_biologist)" title="James Thomson (cell biologist)">James Thomson</a>, which made it possible to have new transplantation methods or various cell types for testing new treatments. In 2006, <a href="/wiki/Shinya_Yamanaka" title="Shinya Yamanaka">Shinya Yamanaka</a>'s team in Kyoto, Japan converted fibroblasts into pluripotent stem cells by modifying the expression of only four genes. The feat represents the origin of induced pluripotent stem cells, known as iPS cells.<sup id="cite_ref-:6_7-1" class="reference"><a href="#cite_note-:6-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>In 2011, a female <a href="/wiki/Maned_wolf" title="Maned wolf">maned wolf</a>, run over by a truck, underwent stem cell treatment at the <a href="/w/index.php?title=Zoo_Bras%C3%ADlia&action=edit&redlink=1" class="new" title="Zoo Brasília (page does not exist)">Zoo Brasília</a><span class="noprint" style="font-size:85%; font-style: normal;"> [<a href="https://pt.wikipedia.org/wiki/Jardim_Zool%C3%B3gico_de_Bras%C3%ADlia" class="extiw" title="pt:Jardim Zoológico de Brasília">pt</a>]</span>, this being the first recorded case of the use of stem cells to heal injuries in a wild animal.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Properties">Properties</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=2" title="Edit section: Properties"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The classical definition of a stem cell requires that it possesses two properties: </p> <ul><li>Self-renewal: the ability to go through numerous <a href="/wiki/Cell_cycle" title="Cell cycle">cycles</a> of <a href="/wiki/Cell_growth" title="Cell growth">cell growth</a> and <a href="/wiki/Cell_division" title="Cell division">cell division</a>, known as <a href="/wiki/Cell_proliferation" title="Cell proliferation">cell proliferation</a>, while maintaining the undifferentiated state.</li> <li><a href="/wiki/Cell_potency" title="Cell potency">Potency</a>: the capacity to <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiate</a> into specialized cell types. In the strictest sense, this requires stem cells to be either <a href="/wiki/Totipotency" class="mw-redirect" title="Totipotency">totipotent</a> or <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotent</a>—to be able to give rise to any mature cell type, although <a href="/wiki/Multipotent" class="mw-redirect" title="Multipotent">multipotent</a> or <a href="/wiki/Unipotent_cell" class="mw-redirect" title="Unipotent cell">unipotent</a> <a href="/wiki/Progenitor_cell" title="Progenitor cell">progenitor cells</a> are sometimes referred to as stem cells. Apart from this, it is said that stem cell function is regulated in a <a href="/wiki/Feedback" title="Feedback">feedback</a> mechanism.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Self-renewal">Self-renewal</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=3" title="Edit section: Self-renewal"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Two mechanisms ensure that a stem cell population is maintained (does not shrink in size): </p><p>1. <a href="/wiki/Asymmetric_cell_division" title="Asymmetric cell division">Asymmetric cell division</a>: a stem cell divides into one mother cell, which is identical to the original stem cell, and another daughter cell, which is differentiated. </p><p>When a stem cell self-renews, it divides and disrupts the undifferentiated state. This self-renewal demands control of cell cycle as well as upkeep of multipotency or pluripotency, which all depends on the stem cell.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>H. </p><p>Stem cells use <a href="/wiki/Telomerase" title="Telomerase">telomerase</a>, a protein that restores <a href="/wiki/Telomeres" class="mw-redirect" title="Telomeres">telomeres</a>, to protect their DNA and extend their cell division limit (the <a href="/wiki/Hayflick_limit" title="Hayflick limit">Hayflick limit</a>).<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Potency_meaning">Potency meaning</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=4" title="Edit section: Potency meaning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cell_potency" title="Cell potency">Cell potency</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Stem_cells_diagram.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Stem_cells_diagram.png/330px-Stem_cells_diagram.png" decoding="async" width="330" height="301" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Stem_cells_diagram.png/495px-Stem_cells_diagram.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Stem_cells_diagram.png/660px-Stem_cells_diagram.png 2x" data-file-width="1015" data-file-height="927" /></a><figcaption>Pluripotent, embryonic stem cells originate as inner cell mass (ICM) cells within a blastocyst. These stem cells can become any tissue in the body, excluding a placenta. Only cells from an earlier stage of the embryo, known as the <a href="/wiki/Morula" class="mw-redirect" title="Morula">morula</a>, are totipotent, able to become all tissues in the body and the extraembryonic placenta.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_embryonic_stem_cells.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Human_embryonic_stem_cells.png/220px-Human_embryonic_stem_cells.png" decoding="async" width="220" height="414" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Human_embryonic_stem_cells.png/330px-Human_embryonic_stem_cells.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/65/Human_embryonic_stem_cells.png/440px-Human_embryonic_stem_cells.png 2x" data-file-width="1010" data-file-height="1899" /></a><figcaption>Human <a href="/wiki/Embryo" title="Embryo">embryonic</a> stem cells <br /> A: Stem cell colonies that are not yet differentiated. <br /> B: <a href="/wiki/Nerve" title="Nerve">Nerve</a> cells, an example of a <a href="/wiki/Cell_type" title="Cell type">cell type</a> after differentiation.</figcaption></figure> <p><a href="/wiki/Cell_potency" title="Cell potency">Potency</a> specifies the differentiation potential (the potential to differentiate into different cell types) of the stem cell.<sup id="cite_ref-Schoeler_15-0" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <ul><li><a href="/wiki/Totipotency" class="mw-redirect" title="Totipotency">Totipotent</a> (also known as omnipotent) stem cells can differentiate into embryonic and extraembryonic cell types. Such cells can construct a complete, viable organism.<sup id="cite_ref-Schoeler_15-1" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> These cells are produced from the <a href="/wiki/Cell_fusion" title="Cell fusion">fusion</a> of an egg and sperm cell. Cells produced by the first few divisions of the fertilized egg are also totipotent.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">Pluripotent</a> stem cells are the descendants of totipotent cells and can differentiate into nearly all cells,<sup id="cite_ref-Schoeler_15-2" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> i.e. cells derived from any of the three <a href="/wiki/Germ_layer" title="Germ layer">germ layers</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Multipotency" class="mw-redirect" title="Multipotency">Multipotent</a> stem cells can differentiate into a number of cell types, but only those of a closely related family of cells.<sup id="cite_ref-Schoeler_15-3" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Oligopotency" class="mw-redirect" title="Oligopotency">Oligopotent</a> stem cells can differentiate into only a few cell types, such as lymphoid or myeloid stem cells.<sup id="cite_ref-Schoeler_15-4" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Unipotency" class="mw-redirect" title="Unipotency">Unipotent</a> cells can produce only one cell type, their own,<sup id="cite_ref-Schoeler_15-5" class="reference"><a href="#cite_note-Schoeler-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> but have the property of self-renewal, which distinguishes them from non-stem cells</li></ul> <div class="mw-heading mw-heading3"><h3 id="Identification">Identification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=5" title="Edit section: Identification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In practice, stem cells are identified by whether they can regenerate tissue. For example, the defining test for bone marrow or <a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">hematopoietic stem cells</a> (HSCs) is the ability to transplant the cells and save an individual without HSCs. This demonstrates that the cells can produce new blood cells over a long term. It should also be possible to isolate stem cells from the transplanted individual, which can themselves be transplanted into another individual without HSCs, demonstrating that the stem cell was able to self-renew. </p><p>Properties of stem cells can be illustrated <i><a href="/wiki/In_vitro" title="In vitro">in vitro</a></i>, using methods such as <a href="/wiki/Clonogenic_assay" title="Clonogenic assay">clonogenic assays</a>, in which single cells are assessed for their ability to differentiate and self-renew.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Stem cells can also be isolated by their possession of a distinctive set of cell surface markers. However, <i>in vitro</i> culture conditions can alter the behavior of cells, making it unclear whether the cells shall behave in a similar manner <i><a href="/wiki/In_vivo" title="In vivo">in vivo</a></i>. There is considerable debate as to whether some proposed adult cell populations are truly stem cells.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Embryonic">Embryonic</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=6" title="Edit section: Embryonic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Embryonic_stem_cell" title="Embryonic stem cell">Embryonic stem cell</a></div> <p><a href="/wiki/Embryonic_stem_cell" title="Embryonic stem cell">Embryonic stem cells</a> (ESCs) are the cells of the <a href="/wiki/Inner_cell_mass" title="Inner cell mass">inner cell mass</a> of a <a href="/wiki/Blastocyst" title="Blastocyst">blastocyst</a>, formed prior to <a href="/wiki/Implantation_(human_embryo)" class="mw-redirect" title="Implantation (human embryo)">implantation</a> in the uterus.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Human_embryonic_development" title="Human embryonic development">human embryonic development</a> the <a href="/wiki/Blastocyst" title="Blastocyst">blastocyst</a> stage is reached 4–5 days after <a href="/wiki/Human_fertilization" title="Human fertilization">fertilization</a>, at which time it consists of 50–150 cells. ESCs are <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a> and give rise during development to all derivatives of the three <a href="/wiki/Germ_layer" title="Germ layer">germ layers</a>: <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a>, <a href="/wiki/Endoderm" title="Endoderm">endoderm</a> and <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a>. In other words, they can develop into each of the more than 200 cell types of the adult <a href="/wiki/Human_body" title="Human body">body</a> when given sufficient and necessary stimulation for a specific cell type. They do not contribute to the <a href="/wiki/Extraembryonic_membrane" title="Extraembryonic membrane">extraembryonic membranes</a> or to the <a href="/wiki/Placenta" title="Placenta">placenta</a>. </p><p>During embryonic development the cells of the inner cell mass continuously divide and become more specialized. For example, a portion of the ectoderm in the dorsal part of the embryo specializes as '<a href="/wiki/Neurectoderm" class="mw-redirect" title="Neurectoderm">neurectoderm</a>', which will become the future <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> (CNS).<sup id="cite_ref-Developmental_biology_22-0" class="reference"><a href="#cite_note-Developmental_biology-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Later in development, <a href="/wiki/Neurulation" title="Neurulation">neurulation</a> causes the neurectoderm to form the <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>. At the neural tube stage, the anterior portion undergoes <a href="/wiki/Encephalization" class="mw-redirect" title="Encephalization">encephalization</a> to generate or 'pattern' the basic form of the brain. At this stage of development, the principal cell type of the CNS is considered a <a href="/wiki/Neural_stem_cell" title="Neural stem cell">neural stem cell</a>. </p><p>The neural stem cells self-renew and at some point transition into <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glial progenitor cells</a> (RGPs). Early-formed RGPs self-renew by symmetrical division to form a reservoir group of <a href="/wiki/Progenitor_cell" title="Progenitor cell">progenitor cells</a>. These cells transition to a <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenic</a> state and start to divide <a href="/wiki/Asymmetric_cell_division" title="Asymmetric cell division">asymmetrically</a> to produce a large diversity of many different neuron types, each with unique gene expression, morphological, and functional characteristics. The process of generating neurons from radial glial cells is called <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a>. The radial glial cell, has a distinctive bipolar morphology with highly elongated processes spanning the thickness of the neural tube wall. It shares some <a href="/wiki/Glial" class="mw-redirect" title="Glial">glial</a> characteristics, most notably the expression of <a href="/wiki/Glial_fibrillary_acidic_protein" title="Glial fibrillary acidic protein">glial fibrillary acidic protein</a> (GFAP).<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> The radial glial cell is the primary neural stem cell of the developing <a href="/wiki/Vertebrate" title="Vertebrate">vertebrate</a> CNS, and its cell body resides in the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a>, adjacent to the developing <a href="/wiki/Ventricular_system" title="Ventricular system">ventricular system</a>. Neural stem cells are committed to the neuronal lineages (<a href="/wiki/Neuron" title="Neuron">neurons</a>, <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a>, and <a href="/wiki/Oligodendrocyte" title="Oligodendrocyte">oligodendrocytes</a>), and thus their potency is restricted.<sup id="cite_ref-Developmental_biology_22-1" class="reference"><a href="#cite_note-Developmental_biology-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>Nearly all research to date has made use of mouse embryonic stem cells (mES) or <b>human embryonic stem cells </b> <b>(hES)</b> derived from the early inner cell mass. Both have the essential stem cell characteristics, yet they require very different environments in order to maintain an undifferentiated state. Mouse ES cells are grown on a layer of <a href="/wiki/Gelatin" title="Gelatin">gelatin</a> as an <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> (for support) and require the presence of <a href="/wiki/Leukemia_inhibitory_factor" title="Leukemia inhibitory factor">leukemia inhibitory factor</a> (LIF) in serum media. A drug cocktail containing inhibitors to <a href="/wiki/Glycogen_synthase_kinase-3_beta" title="Glycogen synthase kinase-3 beta">GSK3B</a> and the <a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK pathway</a>, called 2i, has also been shown to maintain pluripotency in stem cell culture.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Human ESCs are grown on a feeder layer of mouse embryonic <a href="/wiki/Fibroblasts" class="mw-redirect" title="Fibroblasts">fibroblasts</a> and require the presence of basic fibroblast growth factor (bFGF or FGF-2).<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Without optimal culture conditions or genetic manipulation,<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> embryonic stem cells will rapidly differentiate. </p><p>A human embryonic stem cell is also defined by the expression of several transcription factors and cell surface proteins. The transcription factors <a href="/wiki/Oct-4" title="Oct-4">Oct-4</a>, <a href="/wiki/Homeobox_protein_NANOG" title="Homeobox protein NANOG">Nanog</a>, and <a href="/wiki/Sox2" class="mw-redirect" title="Sox2">Sox2</a> form the core regulatory network that ensures the suppression of genes that lead to differentiation and the maintenance of pluripotency.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> The cell surface antigens most commonly used to identify hES cells are the glycolipids <a href="/wiki/Stage_specific_embryonic_antigen_3" class="mw-redirect" title="Stage specific embryonic antigen 3">stage specific embryonic antigen 3</a> and 4, and the keratan sulfate antigens Tra-1-60 and Tra-1-81. The molecular definition of a stem cell includes many more proteins and continues to be a topic of research.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p>By using human embryonic stem cells to produce specialized cells like nerve cells or heart cells in the lab, scientists can gain access to adult human cells without taking tissue from patients. They can then study these specialized adult cells in detail to try to discern complications of diseases, or to study cell reactions to proposed new drugs. </p><p>Because of their combined abilities of unlimited expansion and pluripotency, embryonic stem cells remain a theoretically potential source for <a href="/wiki/Regenerative_medicine" title="Regenerative medicine">regenerative medicine</a> and tissue replacement after injury or disease.,<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> however, there are currently no approved treatments using ES cells. The first human trial was approved by the US Food and Drug Administration in January 2009.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> However, the human trial was not initiated until October 13, 2010 in Atlanta for <a href="/wiki/Spinal_cord_injury_research" title="Spinal cord injury research">spinal cord injury research</a>. On November 14, 2011 the company conducting the trial (<a href="/wiki/Geron_Corporation" title="Geron Corporation">Geron Corporation</a>) announced that it will discontinue further development of its stem cell programs.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Differentiating ES cells into usable cells while avoiding transplant rejection are just a few of the hurdles that embryonic stem cell researchers still face.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> Embryonic stem cells, being pluripotent, require specific signals for correct differentiation – if injected directly into another body, ES cells will differentiate into many different types of cells, causing a <a href="/wiki/Teratoma" title="Teratoma">teratoma</a>. Ethical considerations regarding the use of unborn human tissue are another reason for the lack of approved treatments using embryonic stem cells. Many nations currently have <a href="/wiki/Moratorium_(law)" title="Moratorium (law)">moratoria</a> or limitations on either human ES cell research or the production of new human ES cell lines. </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Mouse_embryonic_stem_cells.jpg" class="mw-file-description" title="Mouse embryonic stem cells with fluorescent marker"><img alt="Mouse embryonic stem cells with fluorescent marker" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Mouse_embryonic_stem_cells.jpg/120px-Mouse_embryonic_stem_cells.jpg" decoding="async" width="120" height="101" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Mouse_embryonic_stem_cells.jpg/180px-Mouse_embryonic_stem_cells.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Mouse_embryonic_stem_cells.jpg/240px-Mouse_embryonic_stem_cells.jpg 2x" data-file-width="600" data-file-height="507" /></a></span></div> <div class="gallerytext"> <a href="/wiki/Mus_musculus" class="mw-redirect" title="Mus musculus">Mouse</a> <a href="/wiki/Mammalian_embryogenesis" title="Mammalian embryogenesis">embryonic</a> stem cells with fluorescent marker</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Human_embryonic_stem_cell_colony_phase.jpg" class="mw-file-description" title="Human embryonic stem cell colony on mouse embryonic fibroblast feeder layer"><img alt="Human embryonic stem cell colony on mouse embryonic fibroblast feeder layer" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Human_embryonic_stem_cell_colony_phase.jpg/120px-Human_embryonic_stem_cell_colony_phase.jpg" decoding="async" width="120" height="105" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Human_embryonic_stem_cell_colony_phase.jpg/180px-Human_embryonic_stem_cell_colony_phase.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Human_embryonic_stem_cell_colony_phase.jpg/240px-Human_embryonic_stem_cell_colony_phase.jpg 2x" data-file-width="1024" data-file-height="894" /></a></span></div> <div class="gallerytext"> Human embryonic stem cell colony on mouse embryonic fibroblast feeder layer</div> </li> </ul> <div class="mw-heading mw-heading3"><h3 id="Mesenchymal_stem_cells">Mesenchymal stem cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=7" title="Edit section: Mesenchymal stem cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Mesenchymal_stem_cell" title="Mesenchymal stem cell">Mesenchymal stem cell</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_mesenchymal_stem_cells.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Human_mesenchymal_stem_cells.gif/220px-Human_mesenchymal_stem_cells.gif" decoding="async" width="220" height="219" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/04/Human_mesenchymal_stem_cells.gif/330px-Human_mesenchymal_stem_cells.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/04/Human_mesenchymal_stem_cells.gif/440px-Human_mesenchymal_stem_cells.gif 2x" data-file-width="600" data-file-height="598" /></a><figcaption>Human mesenchymal stem cells</figcaption></figure> <p>Mesenchymal stem cells (MSC) or mesenchymal stromal cells, also known as medicinal signaling cells are known to be multipotent, which can be found in adult tissues, for example, in the muscle, liver, bone marrow and adipose tissue. Mesenchymal stem cells usually function as structural support in various organs as mentioned above, and control the movement of substances. MSC can differentiate into numerous cell categories as an illustration of adipocytes, osteocytes, and chondrocytes, derived by the mesodermal layer.<sup id="cite_ref-Mesenchymal_and_induced_pluripotent_34-0" class="reference"><a href="#cite_note-Mesenchymal_and_induced_pluripotent-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Where the mesoderm layer provides an increase to the body's skeletal elements, such as relating to the cartilage or bone. The term "meso" means middle, infusion originated from the Greek, signifying that mesenchymal cells are able to range and travel in early embryonic growth among the ectodermal and endodermal layers. This mechanism helps with space-filling thus, key for repairing wounds in adult organisms that have to do with mesenchymal cells in the dermis (skin), bone, or muscle.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Mesenchymal stem cells are known to be essential for regenerative medicine. They are broadly studied in <a href="/wiki/List_of_countries_by_stem_cell_research_trials" title="List of countries by stem cell research trials">clinical trials</a>. Since they are easily isolated and obtain high yield, high plasticity, which makes able to facilitate inflammation and encourage cell growth, cell differentiation, and restoring tissue derived from immunomodulation and immunosuppression. MSC comes from the bone marrow, which requires an aggressive procedure when it comes to isolating the quantity and quality of the isolated cell, and it varies by how old the donor. When comparing the rates of MSC in the bone marrow aspirates and bone marrow stroma, the aspirates tend to have lower rates of MSC than the stroma. MSC are known to be heterogeneous, and they express a high level of pluripotent markers when compared to other types of stem cells, such as embryonic stem cells.<sup id="cite_ref-Mesenchymal_and_induced_pluripotent_34-1" class="reference"><a href="#cite_note-Mesenchymal_and_induced_pluripotent-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> MSCs injection leads to wound healing primarily through stimulation of angiogenesis.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cell_cycle_control">Cell cycle control</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=8" title="Edit section: Cell cycle control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Cell_cycle" title="Cell cycle">Cell cycle</a></div> <p>Embryonic stem cells (ESCs) have the ability to divide indefinitely while keeping their <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a>, which is made possible through specialized mechanisms of <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> control.<sup id="cite_ref-:0_37-0" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Compared to proliferating <a href="/wiki/Somatic_cell" title="Somatic cell">somatic cells</a>, ESCs have unique cell cycle characteristics—such as rapid cell division caused by shortened <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a>, absent <a href="/wiki/G2_phase" title="G2 phase">G0 phase</a>, and modifications in <a href="/wiki/Cell_cycle_checkpoint" title="Cell cycle checkpoint">cell cycle checkpoints</a>—which leaves the cells mostly in <a href="/wiki/S_phase" title="S phase">S phase</a> at any given time.<sup id="cite_ref-:0_37-1" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_38-0" class="reference"><a href="#cite_note-:1-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> ESCs' rapid division is demonstrated by their short doubling time, which ranges from 8 to 10 hours, whereas somatic cells have doubling time of approximately 20 hours or longer.<sup id="cite_ref-:2_39-0" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> As cells differentiate, these properties change: G1 and G2 phases lengthen, leading to longer cell division cycles. This suggests that a specific cell cycle structure may contribute to the establishment of pluripotency.<sup id="cite_ref-:0_37-2" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>Particularly because G1 phase is the phase in which cells have increased sensitivity to differentiation, shortened G1 is one of the key characteristics of ESCs and plays an important role in maintaining undifferentiated <a href="/wiki/Phenotype" title="Phenotype">phenotype</a>. Although the exact molecular mechanism remains only partially understood, several studies have shown insight on how ESCs progress through G1—and  potentially other phases—so rapidly.<sup id="cite_ref-:1_38-1" class="reference"><a href="#cite_note-:1-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p><p>The cell cycle is regulated by complex network of <a href="/wiki/Cyclin" title="Cyclin">cyclins</a>, <a href="/wiki/Cyclin-dependent_kinase" title="Cyclin-dependent kinase">cyclin-dependent kinases</a> (Cdk), <a href="/wiki/Cyclin-dependent_kinase_inhibitor" class="mw-redirect" title="Cyclin-dependent kinase inhibitor">cyclin-dependent kinase inhibitors</a> (Cdkn), pocket proteins of the retinoblastoma (Rb) family, and other accessory factors.<sup id="cite_ref-:2_39-1" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Foundational insight into the distinctive regulation of ESC cell cycle was gained by studies on mouse ESCs (mESCs).<sup id="cite_ref-:1_38-2" class="reference"><a href="#cite_note-:1-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> mESCs showed a cell cycle with highly abbreviated G1 phase, which enabled cells to rapidly alternate between M phase and S phase. In a somatic cell cycle, oscillatory activity of Cyclin-Cdk complexes is observed in sequential action, which controls crucial regulators of the cell cycle to induce unidirectional transitions between phases: <a href="/wiki/Cyclin_D" title="Cyclin D">Cyclin D</a> and Cdk4/6 are active in the G1 phase, while <a href="/wiki/Cyclin_E" title="Cyclin E">Cyclin E</a> and <a href="/wiki/Cyclin-dependent_kinase_2" title="Cyclin-dependent kinase 2">Cdk2</a> are active during the late G1 phase and S phase; and <a href="/wiki/Cyclin_A" title="Cyclin A">Cyclin A</a> and Cdk2 are active in the S phase and G2, while <a href="/wiki/Cyclin_B" title="Cyclin B">Cyclin B</a> and <a href="/wiki/Cyclin-dependent_kinase_1" title="Cyclin-dependent kinase 1">Cdk1</a> are active in G2 and M phase.<sup id="cite_ref-:2_39-2" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> However, in mESCs, this typically ordered and oscillatory activity of Cyclin-Cdk complexes is absent. Rather, the Cyclin E/Cdk2 complex is constitutively active throughout the cycle, keeping <a href="/wiki/Retinoblastoma_protein" title="Retinoblastoma protein">retinoblastoma protein</a> (pRb) <a href="/wiki/Hyperphosphorylated" class="mw-redirect" title="Hyperphosphorylated">hyperphosphorylated</a> and thus inactive. This allows for direct transition from M phase to the late G1 phase, leading to absence of D-type cyclins and therefore a shortened G1 phase.<sup id="cite_ref-:1_38-3" class="reference"><a href="#cite_note-:1-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Cdk2 activity is crucial for both cell cycle regulation and cell-fate decisions in mESCs; downregulation of Cdk2 activity prolongs G1 phase progression, establishes a somatic cell-like cell cycle, and induces expression of differentiation markers.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p><p>In human ESCs (hESCs), the duration of G1 is dramatically shortened. This has been attributed to high mRNA levels of G1-related Cyclin D2 and Cdk4 genes and low levels of cell cycle regulatory proteins that inhibit cell cycle progression at G1, such as <a href="/wiki/P21Cip1" class="mw-redirect" title="P21Cip1">p21<sup>CipP1</sup></a>, p27<sup>Kip1</sup>, and p57<sup>Kip2</sup>.<sup id="cite_ref-:0_37-3" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Furthermore, regulators of Cdk4 and Cdk6 activity, such as members of the Ink family of inhibitors (p15, p16, p18, and p19), are expressed at low levels or not at all. Thus, similar to mESCs, hESCs show high Cdk activity, with Cdk2 exhibiting the highest kinase activity. Also similar to mESCs, hESCs demonstrate the importance of Cdk2 in G1 phase regulation by showing that G1 to S transition is delayed when Cdk2 activity is inhibited and G1 is arrest when Cdk2 is knocked down.<sup id="cite_ref-:0_37-4" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> However unlike mESCs, hESCs have a functional G1 phase. hESCs show that the activities of Cyclin E/Cdk2 and Cyclin A/Cdk2 complexes are cell cycle-dependent and the Rb checkpoint in G1 is functional.<sup id="cite_ref-:2_39-3" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p><p>ESCs are also characterized by G1 checkpoint non-functionality, even though the G1 checkpoint is crucial for maintaining genomic stability. In response to <a href="/wiki/DNA_damage" class="mw-redirect" title="DNA damage">DNA damage</a>, ESCs do not stop in G1 to repair DNA damages but instead, depend on S and G2/M checkpoints or undergo apoptosis. The absence of G1 checkpoint in ESCs allows for the removal of cells with damaged DNA, hence avoiding potential mutations from inaccurate DNA repair.<sup id="cite_ref-:0_37-5" class="reference"><a href="#cite_note-:0-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Consistent with this idea, ESCs are hypersensitive to DNA damage to minimize mutations passed onto the next generation.<sup id="cite_ref-:2_39-4" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Fetal">Fetal</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=9" title="Edit section: Fetal"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The primitive stem cells located in the organs of <a href="/wiki/Fetuses" class="mw-redirect" title="Fetuses">fetuses</a> are referred to as fetal stem cells.<sup id="cite_ref-isbn981-256-126-9_42-0" class="reference"><a href="#cite_note-isbn981-256-126-9-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>There are two types of fetal stem cells: </p> <ol><li>Fetal proper stem cells come from the tissue of the fetus proper and are generally obtained after an <a href="/wiki/Abortion" title="Abortion">abortion</a>. These stem cells are not immortal but have a high level of division and are multipotent.</li> <li>Extraembryonic fetal stem cells come from <a href="/wiki/Extraembryonic_membrane" title="Extraembryonic membrane">extraembryonic membranes</a>, and are generally not distinguished from adult stem cells. These stem cells are acquired after birth, they are not immortal but have a high level of cell division, and are pluripotent.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup></li></ol> <div class="mw-heading mw-heading2"><h2 id="Adult">Adult</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=10" title="Edit section: Adult"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Adult_stem_cell" title="Adult stem cell">Adult stem cell</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stem_cell_division_and_differentiation.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Stem_cell_division_and_differentiation.svg/220px-Stem_cell_division_and_differentiation.svg.png" decoding="async" width="220" height="440" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/18/Stem_cell_division_and_differentiation.svg/330px-Stem_cell_division_and_differentiation.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/18/Stem_cell_division_and_differentiation.svg/440px-Stem_cell_division_and_differentiation.svg.png 2x" data-file-width="200" data-file-height="400" /></a><figcaption>Stem cell division and differentiation A: stem cell; B: progenitor cell; C: differentiated cell; 1: symmetric stem cell division; 2: asymmetric stem cell division; 3: progenitor division; 4: terminal differentiation</figcaption></figure> <p>Adult stem cells, also called <a href="/wiki/Somatic_(biology)" title="Somatic (biology)">somatic</a> (from Greek σωματικóς, "of the body") stem cells, are stem cells which maintain and repair the tissue in which they are found.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>There are three known accessible sources of <a href="/wiki/Autologous" class="mw-redirect" title="Autologous">autologous</a> adult stem cells in humans: </p> <ol><li><a href="/wiki/Bone_marrow" title="Bone marrow">Bone marrow</a>, which requires extraction by <i>harvesting</i>, usually from pelvic bones via surgery.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup></li> <li>Adipose tissue (fat cells), which requires extraction by liposuction.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup></li> <li>Blood, which requires extraction through <a href="/wiki/Apheresis" title="Apheresis">apheresis</a>, wherein blood is drawn from the donor (similar to a blood donation), and passed through a machine that extracts the stem cells and returns other portions of the blood to the donor.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup></li></ol> <p>Stem cells can also be taken from <a href="/wiki/Umbilical_cord_blood" class="mw-redirect" title="Umbilical cord blood">umbilical cord blood</a> just after birth. Of all stem cell types, autologous harvesting involves the least risk. By definition, autologous cells are obtained from one's own body, just as one may bank their own blood for elective surgical procedures.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2021)">citation needed</span></a></i>]</sup> </p><p>Pluripotent adult stem cells are rare and generally small in number, but they can be found in umbilical cord blood and other tissues.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Bone_marrow" title="Bone marrow">Bone marrow</a> is a rich source of adult stem cells,<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> which have been used in treating several conditions including liver cirrhosis,<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> chronic limb ischemia<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> and endstage heart failure.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> The quantity of bone marrow stem cells declines with age and is greater in males than females during reproductive years.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> Much adult stem cell research to date has aimed to characterize their potency and self-renewal capabilities.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> DNA damage accumulates with age in both stem cells and the cells that comprise the stem cell environment. This accumulation is considered to be responsible, at least in part, for increasing stem cell dysfunction with aging (see <a href="/wiki/DNA_damage_theory_of_aging" title="DNA damage theory of aging">DNA damage theory of aging</a>).<sup id="cite_ref-pmid24576896_55-0" class="reference"><a href="#cite_note-pmid24576896-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p><p>Most adult stem cells are lineage-restricted (<a href="/wiki/Multipotent" class="mw-redirect" title="Multipotent">multipotent</a>) and are generally referred to by their tissue origin (<a href="/wiki/Mesenchymal_stem_cell" title="Mesenchymal stem cell">mesenchymal stem cell</a>, adipose-derived stem cell, <a href="/wiki/Endothelial_stem_cell" title="Endothelial stem cell">endothelial stem cell</a>, <a href="/wiki/Dental_pulp_stem_cell" title="Dental pulp stem cell">dental pulp stem cell</a>, etc.).<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Muse_cell" title="Muse cell">Muse cells</a> (multi-lineage differentiating stress enduring cells) are a recently discovered pluripotent stem cell type found in multiple adult tissues, including adipose, dermal fibroblasts, and bone marrow. While rare, muse cells are identifiable by their expression of <a href="/wiki/SSEA-3" class="mw-redirect" title="SSEA-3">SSEA-3</a>, a marker for undifferentiated stem cells, and general mesenchymal stem cells markers such as CD90, <a href="/wiki/CD105" class="mw-redirect" title="CD105">CD105</a>. When subjected to single cell suspension culture, the cells will generate clusters that are similar to embryoid bodies in morphology as well as gene expression, including canonical pluripotency markers <a href="/wiki/Oct4" class="mw-redirect" title="Oct4">Oct4</a>, <a href="/wiki/Sox2" class="mw-redirect" title="Sox2">Sox2</a>, and <a href="/wiki/Homeobox_protein_NANOG" title="Homeobox protein NANOG">Nanog</a>.<sup id="cite_ref-Kuroda_58-0" class="reference"><a href="#cite_note-Kuroda-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p><p>Adult stem cell treatments have been successfully used for many years to treat leukemia and related bone/blood cancers through bone marrow transplants.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Adult stem cells are also used in veterinary medicine to treat tendon and ligament injuries in horses.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p><p>The use of adult stem cells in research and therapy is not as <a href="/wiki/Stem_cell_controversy" title="Stem cell controversy">controversial</a> as the use of <a href="/wiki/Embryonic_stem_cell" title="Embryonic stem cell">embryonic stem cells</a>, because the production of adult stem cells does not require the destruction of an <a href="/wiki/Embryo" title="Embryo">embryo</a>. Additionally, in instances where adult stem cells are obtained from the intended recipient (an <a href="/wiki/Autograft" class="mw-redirect" title="Autograft">autograft</a>), the risk of rejection is essentially non-existent. Consequently, more US government funding is being provided for adult stem cell research.<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p><p>With the increasing demand of human adult stem cells for both research and clinical purposes (typically 1–5 million cells per kg of body weight are required per treatment) it becomes of utmost importance to bridge the gap between the need to expand the cells in vitro and the capability of harnessing the factors underlying replicative senescence. Adult stem cells are known to have a limited lifespan in vitro and to enter replicative senescence almost undetectably upon starting in vitro culturing.<sup id="cite_ref-Oliveira_62-0" class="reference"><a href="#cite_note-Oliveira-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Hematopoietic_stem_cells">Hematopoietic stem cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=11" title="Edit section: Hematopoietic stem cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">Hematopoietic stem cells</a> (HSCs) are vulnerable to <a href="/wiki/DNA_damage_(naturally_occurring)" title="DNA damage (naturally occurring)">DNA damage</a> and <a href="/wiki/Mutation" title="Mutation">mutations</a> that increase with age.<sup id="cite_ref-Zhang2020_63-0" class="reference"><a href="#cite_note-Zhang2020-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> This vulnerability may explain the increased risk of slow growing blood cancers (myeloid malignancies) in the elderly.<sup id="cite_ref-Zhang2020_63-1" class="reference"><a href="#cite_note-Zhang2020-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> Several factors appear to influence HSC aging including responses to the production of <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> that may cause DNA damage and genetic mutations as well as altered <a href="/wiki/Epigenetics" title="Epigenetics">epigenetic</a> profiling.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Amniotic">Amniotic</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=12" title="Edit section: Amniotic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Also called perinatal stem cells, these multipotent stem cells are found in <a href="/wiki/Amniotic_fluid" title="Amniotic fluid">amniotic fluid</a> and umbilical cord blood. These stem cells are very active, expand extensively without feeders and are not tumorigenic. <a href="/wiki/Amniotic_stem_cells" title="Amniotic stem cells">Amniotic stem cells</a> are multipotent and can differentiate in cells of adipogenic, osteogenic, myogenic, endothelial, hepatic and also neuronal lines.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> Amniotic stem cells are a topic of active research. </p><p>Use of stem cells from <a href="/wiki/Amniotic_fluid" title="Amniotic fluid">amniotic fluid</a> overcomes the ethical objections to using human embryos as a source of cells. <a href="/wiki/Roman_Catholic" class="mw-redirect" title="Roman Catholic">Roman Catholic</a> teaching forbids the use of embryonic stem cells in experimentation; accordingly, the <a href="/wiki/Holy_See" title="Holy See">Vatican</a> newspaper "<a href="/wiki/Osservatore_Romano" class="mw-redirect" title="Osservatore Romano">Osservatore Romano</a>" called amniotic stem cells "the future of medicine".<sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> </p><p>It is possible to collect amniotic stem cells for donors or for autologous use: the first US amniotic stem cells bank<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> was opened in 2009 in Medford, MA, by <a href="/wiki/Biocell_Center" title="Biocell Center">Biocell Center</a> Corporation<sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> and collaborates with various hospitals and universities all over the world.<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Induced_pluripotent">Induced pluripotent</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=13" title="Edit section: Induced pluripotent"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Induced_pluripotent_stem_cell" title="Induced pluripotent stem cell">Induced pluripotent stem cell</a></div> <p>Adult stem cells have limitations with their potency; unlike <a href="/wiki/Embryonic_stem_cell" title="Embryonic stem cell">embryonic stem cells</a> (ESCs), they are not able to differentiate into cells from all three <a href="/wiki/Germ_layers" class="mw-redirect" title="Germ layers">germ layers</a>. As such, they are deemed <a href="/wiki/Cell_potency" title="Cell potency">multipotent</a>. </p><p>However, <a href="/wiki/Reprogramming" title="Reprogramming">reprogramming</a> allows for the creation of pluripotent cells, <a href="/wiki/Induced_pluripotent_stem_cell" title="Induced pluripotent stem cell">induced pluripotent stem cells</a> (iPSCs), from adult cells. These are not adult stem cells, but somatic cells (e.g. epithelial cells) reprogrammed to give rise to cells with pluripotent capabilities. Using genetic reprogramming with protein <a href="/wiki/Transcription_factors" class="mw-redirect" title="Transcription factors">transcription factors</a>, pluripotent stem cells with ESC-like capabilities have been derived.<sup id="cite_ref-Economist2007_11_22_73-0" class="reference"><a href="#cite_note-Economist2007_11_22-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> The first demonstration of induced pluripotent stem cells was conducted by <a href="/wiki/Shinya_Yamanaka" title="Shinya Yamanaka">Shinya Yamanaka</a> and his colleagues at <a href="/wiki/Kyoto_University" title="Kyoto University">Kyoto University</a>.<sup id="cite_ref-overview_76-0" class="reference"><a href="#cite_note-overview-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> They used the transcription factors <a href="/wiki/Oct-4" title="Oct-4">Oct3/4</a>, <a href="/wiki/Sox2" class="mw-redirect" title="Sox2">Sox2</a>, <a href="/wiki/C-Myc" class="mw-redirect" title="C-Myc">c-Myc</a>, and <a href="/wiki/Klf4" class="mw-redirect" title="Klf4">Klf4</a> to reprogram mouse fibroblast cells into pluripotent cells.<sup id="cite_ref-Economist2007_11_22_73-1" class="reference"><a href="#cite_note-Economist2007_11_22-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-og_Yak_77-0" class="reference"><a href="#cite_note-og_Yak-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> Subsequent work used these factors to induce pluripotency in human fibroblast cells.<sup id="cite_ref-2007_Yak_78-0" class="reference"><a href="#cite_note-2007_Yak-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Junying_Yu" title="Junying Yu">Junying Yu</a>, <a href="/wiki/James_Thomson_(cell_biologist)" title="James Thomson (cell biologist)">James Thomson</a>, and their colleagues at the <a href="/wiki/University_of_Wisconsin%E2%80%93Madison" title="University of Wisconsin–Madison">University of Wisconsin–Madison</a> used a different set of factors, Oct4, Sox2, Nanog and Lin28, and carried out their experiments using cells from human <a href="/wiki/Foreskin" title="Foreskin">foreskin</a>.<sup id="cite_ref-Economist2007_11_22_73-2" class="reference"><a href="#cite_note-Economist2007_11_22-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-wisconsin_79-0" class="reference"><a href="#cite_note-wisconsin-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> However, they were able to replicate <a href="/wiki/Yamanaka" title="Yamanaka">Yamanaka</a>'s finding that inducing pluripotency in human cells was possible. </p><p>Induced pluripotent stem cells differ from embryonic stem cells. They share many similar properties, such as <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a> and differentiation potential, the expression of <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a> genes, <a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">epigenetic</a> patterns, <a href="/wiki/Embryoid_body" title="Embryoid body">embryoid body</a> and <a href="/wiki/Teratoma" title="Teratoma">teratoma</a> formation, and viable <a href="/wiki/Chimera_(genetics)" title="Chimera (genetics)">chimera</a> formation,<sup id="cite_ref-overview_76-1" class="reference"><a href="#cite_note-overview-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-og_Yak_77-1" class="reference"><a href="#cite_note-og_Yak-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> but there are many differences within these properties. The chromatin of iPSCs appears to be more "closed" or methylated than that of ESCs.<sup id="cite_ref-overview_76-2" class="reference"><a href="#cite_note-overview-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-og_Yak_77-2" class="reference"><a href="#cite_note-og_Yak-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> Similarly, the gene expression pattern between ESCs and iPSCs, or even iPSCs sourced from different origins.<sup id="cite_ref-overview_76-3" class="reference"><a href="#cite_note-overview-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> There are thus questions about the "completeness" of <a href="/wiki/Reprogramming" title="Reprogramming">reprogramming</a> and the somatic memory of induced pluripotent stem cells. Despite this, inducing somatic cells to be pluripotent appears to be viable. </p><p>As a result of the success of these experiments, <a href="/wiki/Ian_Wilmut" title="Ian Wilmut">Ian Wilmut</a>, who helped create the first cloned animal <a href="/wiki/Dolly_the_Sheep" class="mw-redirect" title="Dolly the Sheep">Dolly the Sheep</a>, has announced that he will abandon <a href="/wiki/Somatic_cell_nuclear_transfer" title="Somatic cell nuclear transfer">somatic cell nuclear transfer</a> as an avenue of research.<sup id="cite_ref-80" class="reference"><a href="#cite_note-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p><p>The ability to induce pluripotency benefits developments in <a href="/wiki/Tissue_engineering" title="Tissue engineering">tissue engineering</a>. By providing a suitable scaffold and microenvironment, iPSC can be differentiated into cells of therapeutic application, and for <i>in vitro</i> models to study toxins and pathogenesis.<sup id="cite_ref-81" class="reference"><a href="#cite_note-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p><p>Induced pluripotent stem cells provide several therapeutic advantages. Like ESCs, they are <a href="/wiki/Pluripotent" class="mw-redirect" title="Pluripotent">pluripotent</a>. They thus have great differentiation potential; theoretically, they could produce any cell within the human body (if <a href="/wiki/Reprogramming" title="Reprogramming">reprogramming</a> to pluripotency was "complete").<sup id="cite_ref-overview_76-4" class="reference"><a href="#cite_note-overview-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> Moreover, unlike ESCs, they potentially could allow doctors to create a pluripotent stem cell line for each individual patient.<sup id="cite_ref-R&D_82-0" class="reference"><a href="#cite_note-R&D-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> Frozen blood samples can be used as a valuable source of induced pluripotent stem cells.<sup id="cite_ref-pmid20621045_83-0" class="reference"><a href="#cite_note-pmid20621045-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> Patient specific stem cells allow for the screening for side effects before drug treatment, as well as the reduced risk of transplantation rejection.<sup id="cite_ref-R&D_82-1" class="reference"><a href="#cite_note-R&D-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> Despite their current limited use therapeutically, iPSCs hold great potential for future use in medical treatment and research. </p> <div class="mw-heading mw-heading3"><h3 id="Cell_cycle_control_2">Cell cycle control</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=14" title="Edit section: Cell cycle control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The key factors controlling the cell cycle also regulate <a href="/wiki/Pluripotency" class="mw-redirect" title="Pluripotency">pluripotency</a>. Thus, manipulation of relevant genes can maintain pluripotency and reprogram somatic cells to an induced pluripotent state.<sup id="cite_ref-:2_39-5" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> However, reprogramming of somatic cells is often low in efficiency and considered <a href="/wiki/Stochastic" title="Stochastic">stochastic</a>.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> </p><p>With the idea that a more rapid cell cycle is a key component of pluripotency, reprogramming efficiency can be improved. Methods for improving pluripotency through manipulation of cell cycle regulators include: overexpression of Cyclin D/Cdk4, phosphorylation of <a href="/wiki/SOX2" title="SOX2">Sox2</a> at S39 and S253, overexpression of Cyclin A and Cyclin E, knockdown of Rb, and knockdown of members of the <a href="/wiki/CIP/KIP" title="CIP/KIP">Cip/Kip</a> family or the Ink family.<sup id="cite_ref-:2_39-6" class="reference"><a href="#cite_note-:2-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Furthermore, reprogramming efficiency is correlated with the number of cell divisions happened during the stochastic phase, which is suggested by the growing inefficiency of reprogramming of older or slow diving cells.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Lineage">Lineage</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=15" title="Edit section: Lineage"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Stem_cell_line" class="mw-redirect" title="Stem cell line">Stem cell line</a></div> <p>Lineage is an important procedure to analyze developing embryos. Since cell lineages shows the relationship between cells at each division. This helps in analyzing stem cell lineages along the way which helps recognize stem cell effectiveness, lifespan, and other factors. With the technique of cell lineage mutant genes can be analyzed in stem cell clones that can help in genetic pathways. These pathways can regulate how the stem cell perform.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> </p><p>To ensure self-renewal, stem cells undergo two types of cell division (see <i>Stem cell division and differentiation</i> diagram). Symmetric division gives rise to two identical daughter cells both endowed with stem cell properties. Asymmetric division, on the other hand, produces only one stem cell and a <a href="/wiki/Progenitor_cell" title="Progenitor cell">progenitor cell</a> with limited self-renewal potential. Progenitors can go through several rounds of cell division before terminally <a href="/wiki/Cell_differentiation" class="mw-redirect" title="Cell differentiation">differentiating</a> into a mature cell. It is possible that the molecular distinction between symmetric and asymmetric divisions lies in differential segregation of cell membrane proteins (such as <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a>) between the daughter cells.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p><p>An alternative theory is that stem cells remain undifferentiated due to environmental cues in their particular <a href="/wiki/Stem-cell_niche" title="Stem-cell niche">niche</a>. Stem cells differentiate when they leave that niche or no longer receive those signals. Studies in <i>Drosophila</i> germarium have identified the signals <a href="/wiki/Decapentaplegic" title="Decapentaplegic">decapentaplegic</a> and adherens junctions that prevent germarium stem cells from differentiating.<sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> </p><p>In the United States, Executive Order 13505 established that federal money can be used for research in which approved human embryonic stem-cell (hESC) lines are used, but it cannot be used to derive new lines.<sup id="cite_ref-90" class="reference"><a href="#cite_note-90"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup> The National Institutes of Health (NIH) Guidelines on Human Stem Cell Research, effective July 7, 2009, implemented the Executive Order 13505 by establishing criteria which hESC lines must meet to be approved for funding.<sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> The NIH Human Embryonic Stem Cell Registry can be accessed online and has updated information on cell lines eligible for NIH funding.<sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> There are 486 approved lines as of January 2022.<sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Therapies">Therapies</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=16" title="Edit section: Therapies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Stem_cell_therapy" class="mw-redirect" title="Stem cell therapy">Stem cell therapy</a></div> <p>Stem cell therapy is the use of stem cells to treat or prevent a disease or condition. <a href="/wiki/Bone_marrow_transplant" class="mw-redirect" title="Bone marrow transplant">Bone marrow transplant</a> is a form of stem cell therapy that has been used for many years because it has proven to be effective in clinical trials.<sup id="cite_ref-Why_Perform_a_Stem_Cell_Transplant_94-0" class="reference"><a href="#cite_note-Why_Perform_a_Stem_Cell_Transplant-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NCI_95-0" class="reference"><a href="#cite_note-NCI-95"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup> Stem cell implantation may help in strengthening the left-ventricle of the heart, as well as retaining the heart tissue to patients who have suffered from heart attacks in the past.<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> </p><p>For over 50 years, <a href="/wiki/Hematopoietic_stem_cell_transplantation" title="Hematopoietic stem cell transplantation">hematopoietic stem cell transplantation</a> (HSCT) has been used to treat people with conditions such as <a href="/wiki/Leukaemia" class="mw-redirect" title="Leukaemia">leukaemia</a> and <a href="/wiki/Lymphoma" title="Lymphoma">lymphoma</a>; this is the only widely practiced form of stem-cell therapy.<sup id="cite_ref-Why_Perform_a_Stem_Cell_Transplant_94-1" class="reference"><a href="#cite_note-Why_Perform_a_Stem_Cell_Transplant-94"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NCI2_97-0" class="reference"><a href="#cite_note-NCI2-97"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid18721775_98-0" class="reference"><a href="#cite_note-pmid18721775-98"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup> As of 2016<sup class="plainlinks noexcerpt noprint asof-tag update" style="display:none;"><a class="external text" href="https://en.wikipedia.org/w/index.php?title=Stem_cell&action=edit">[update]</a></sup>, the only established therapy using stem cells is <a href="/wiki/Hematopoietic_stem_cell_transplantation" title="Hematopoietic stem cell transplantation">hematopoietic stem cell transplantation</a>.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup> This usually takes the form of a <a href="/wiki/Bone_marrow" title="Bone marrow">bone-marrow</a> transplantation, but the cells can also be derived from <a href="/wiki/Umbilical_cord_blood" class="mw-redirect" title="Umbilical cord blood">umbilical cord blood</a>. Research is underway to develop various sources for stem cells as well as to apply stem-cell treatments for <a href="/wiki/Neurodegenerative_diseases" class="mw-redirect" title="Neurodegenerative diseases">neurodegenerative diseases</a><sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-102" class="reference"><a href="#cite_note-102"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> and conditions such as <a href="/wiki/Diabetes" title="Diabetes">diabetes</a> and <a href="/wiki/Heart_disease" class="mw-redirect" title="Heart disease">heart disease</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Advantages">Advantages</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=17" title="Edit section: Advantages"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stem cell treatments may lower symptoms of the disease or condition that is being treated. The lowering of symptoms may allow patients to reduce the drug intake of the disease or condition. Stem cell treatment may also provide knowledge for society to further stem cell understanding and future treatments.<sup id="cite_ref-103" class="reference"><a href="#cite_note-103"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> The physicians' creed would be to do no injury, and stem cells make that simpler than ever before. Surgical processes by their character are harmful. Tissue has to be dropped as a way to reach a successful outcome. One may prevent the dangers of surgical interventions using stem cells. Additionally, there's a possibility of disease, and whether the procedure fails, further surgery may be required. Risks associated with anesthesia can also be eliminated with stem cells.<sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> On top of that, stem cells have been harvested from the patient's body and redeployed in which they're wanted. Since they come from the patient's own body, this is referred to as an autologous treatment. Autologous remedies are thought to be the safest because there's likely zero probability of donor substance rejection. </p> <div class="mw-heading mw-heading3"><h3 id="Disadvantages">Disadvantages</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=18" title="Edit section: Disadvantages"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stem cell treatments may require <a href="/wiki/Immunosuppression" title="Immunosuppression">immunosuppression</a> because of a requirement for radiation before the transplant to remove the person's previous cells, or because the patient's immune system may target the stem cells. One approach to avoid the second possibility is to use stem cells from the same patient who is being treated. </p><p>Pluripotency in certain stem cells could also make it difficult to obtain a specific cell type. It is also difficult to obtain the exact cell type needed, because not all cells in a population differentiate uniformly. Undifferentiated cells can create tissues other than desired types.<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup> </p><p>Some stem cells form tumors after transplantation;<sup id="cite_ref-usn2009_106-0" class="reference"><a href="#cite_note-usn2009-106"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> pluripotency is linked to tumor formation especially in embryonic stem cells, fetal proper stem cells, induced pluripotent stem cells. Fetal proper stem cells form tumors despite multipotency.<sup id="cite_ref-107" class="reference"><a href="#cite_note-107"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> </p><p>Ethical concerns are also raised about the practice of using or researching embryonic stem cells. Harvesting cells from the blastocyst results in the death of the blastocyst. The concern is whether or not the blastocyst should be considered as a human life.<sup id="cite_ref-108" class="reference"><a href="#cite_note-108"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> The debate on this issue is mainly a philosophical one, not a scientific one. </p> <div class="mw-heading mw-heading3"><h3 id="Stem_cell_tourism">Stem cell tourism</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=19" title="Edit section: Stem cell tourism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stem cell tourism is the part of the <a href="/wiki/Medical_tourism" title="Medical tourism">medical tourism</a> industry in which patients travel to obtain stem cell procedures.<sup id="cite_ref-109" class="reference"><a href="#cite_note-109"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> </p><p>The United States has had an explosion of "stem cell clinics".<sup id="cite_ref-110" class="reference"><a href="#cite_note-110"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> Stem cell procedures are highly profitable for clinics. The advertising sounds authoritative but the efficacy and safety of the procedures is unproven. Patients sometimes experience complications, such as spinal tumors<sup id="cite_ref-Berkowitz_et_al_2016_111-0" class="reference"><a href="#cite_note-Berkowitz_et_al_2016-111"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> and death. The high expense can also lead to financial problems.<sup id="cite_ref-Berkowitz_et_al_2016_111-1" class="reference"><a href="#cite_note-Berkowitz_et_al_2016-111"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> According to researchers, there is a need to educate the public, patients, and doctors about this issue.<sup id="cite_ref-112" class="reference"><a href="#cite_note-112"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> </p><p>According to the <a href="/wiki/International_Society_for_Stem_Cell_Research" title="International Society for Stem Cell Research">International Society for Stem Cell Research</a>, the largest academic organization that advocates for stem cell research, stem cell therapies are under development and cannot yet be said to be proven.<sup id="cite_ref-113" class="reference"><a href="#cite_note-113"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup> Doctors should inform patients that clinical trials continue to investigate whether these therapies are safe and effective but that unethical clinics present them as proven.<sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Research">Research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=20" title="Edit section: Research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Consumer_Watchdog_vs._Wisconsin_Alumni_Research_Foundation" class="mw-redirect" title="Consumer Watchdog vs. Wisconsin Alumni Research Foundation">Consumer Watchdog vs. Wisconsin Alumni Research Foundation</a></div> <p>Some of the fundamental <a href="/wiki/Patent" title="Patent">patents</a> covering human embryonic stem cells are owned by the <a href="/wiki/Wisconsin_Alumni_Research_Foundation" title="Wisconsin Alumni Research Foundation">Wisconsin Alumni Research Foundation</a> (WARF) – they are patents 5,843,780, 6,200,806, and 7,029,913 invented by <a href="/wiki/James_Thomson_(cell_biologist)" title="James Thomson (cell biologist)">James A. Thomson</a>. WARF does not enforce these patents against academic scientists, but does enforce them against companies.<sup id="cite_ref-stemcellPatent_116-0" class="reference"><a href="#cite_note-stemcellPatent-116"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup> </p><p>In 2006, a request for the <a href="/wiki/US_Patent_and_Trademark_Office" class="mw-redirect" title="US Patent and Trademark Office">US Patent and Trademark Office</a> (USPTO) to re-examine the three patents was filed by the Public Patent Foundation<sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> on behalf of its client, the non-profit patent-watchdog group <a href="/wiki/Consumer_Watchdog" title="Consumer Watchdog">Consumer Watchdog</a> (formerly the Foundation for Taxpayer and Consumer Rights).<sup id="cite_ref-stemcellPatent_116-1" class="reference"><a href="#cite_note-stemcellPatent-116"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup> In the re-examination process, which involves several rounds of discussion between the USPTO and the parties, the USPTO initially agreed with Consumer Watchdog and rejected all the claims in all three patents,<sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup> however in response, WARF amended the claims of all three patents to make them more narrow, and in 2008 the USPTO found the amended claims in all three patents to be patentable. The decision on one of the patents (7,029,913) was appealable, while the decisions on the other two were not.<sup id="cite_ref-119" class="reference"><a href="#cite_note-119"><span class="cite-bracket">[</span>119<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">[</span>120<span class="cite-bracket">]</span></a></sup> Consumer Watchdog appealed the granting of the '913 patent to the USPTO's Board of Patent Appeals and Interferences (BPAI) which granted the appeal, and in 2010 the BPAI decided that the amended claims of the '913 patent were not patentable.<sup id="cite_ref-121" class="reference"><a href="#cite_note-121"><span class="cite-bracket">[</span>121<span class="cite-bracket">]</span></a></sup> However, WARF was able to re-open prosecution of the case and did so, amending the claims of the '913 patent again to make them more narrow, and in January 2013 the amended claims were allowed.<sup id="cite_ref-122" class="reference"><a href="#cite_note-122"><span class="cite-bracket">[</span>122<span class="cite-bracket">]</span></a></sup> </p><p>In July 2013, Consumer Watchdog announced that it would appeal the decision to allow the claims of the '913 patent to the US <a href="/wiki/Court_of_Appeals_for_the_Federal_Circuit" class="mw-redirect" title="Court of Appeals for the Federal Circuit">Court of Appeals for the Federal Circuit</a> (CAFC), the federal appeals court that hears patent cases.<sup id="cite_ref-123" class="reference"><a href="#cite_note-123"><span class="cite-bracket">[</span>123<span class="cite-bracket">]</span></a></sup> At a hearing in December 2013, the CAFC raised the question of whether Consumer Watchdog had <a href="/wiki/Legal_standing" class="mw-redirect" title="Legal standing">legal standing</a> to appeal; the case could not proceed until that issue was resolved.<sup id="cite_ref-124" class="reference"><a href="#cite_note-124"><span class="cite-bracket">[</span>124<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Conditions">Conditions</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=21" title="Edit section: Conditions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stem_cell_treatments.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Stem_cell_treatments.svg/330px-Stem_cell_treatments.svg.png" decoding="async" width="330" height="271" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Stem_cell_treatments.svg/495px-Stem_cell_treatments.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/bd/Stem_cell_treatments.svg/660px-Stem_cell_treatments.svg.png 2x" data-file-width="1450" data-file-height="1190" /></a><figcaption>Diseases and conditions where stem cell treatment is being investigated</figcaption></figure> <p>Diseases and conditions where stem cell treatment is being investigated include: </p> <ul><li><a href="/wiki/Diabetes" title="Diabetes">Diabetes</a><sup id="cite_ref-nih_125-0" class="reference"><a href="#cite_note-nih-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Pattern_hair_loss" title="Pattern hair loss">Androgenic Alopecia and hair loss</a><sup id="cite_ref-126" class="reference"><a href="#cite_note-126"><span class="cite-bracket">[</span>126<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-127" class="reference"><a href="#cite_note-127"><span class="cite-bracket">[</span>127<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Rheumatoid_arthritis" title="Rheumatoid arthritis">Rheumatoid arthritis</a><sup id="cite_ref-nih_125-1" class="reference"><a href="#cite_note-nih-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a><sup id="cite_ref-nih_125-2" class="reference"><a href="#cite_note-nih-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Alzheimer%27s_disease" title="Alzheimer's disease">Alzheimer's disease</a><sup id="cite_ref-nih_125-3" class="reference"><a href="#cite_note-nih-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Respiratory_disease" title="Respiratory disease">Respiratory disease</a><sup id="cite_ref-128" class="reference"><a href="#cite_note-128"><span class="cite-bracket">[</span>128<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Osteoarthritis" title="Osteoarthritis">Osteoarthritis</a><sup id="cite_ref-nih_125-4" class="reference"><a href="#cite_note-nih-125"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Stroke" title="Stroke">Stroke</a> and <a href="/wiki/Traumatic_brain_injury" title="Traumatic brain injury">traumatic brain injury</a> repair<sup id="cite_ref-Steinberg2000_129-0" class="reference"><a href="#cite_note-Steinberg2000-129"><span class="cite-bracket">[</span>129<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Learning_disability" title="Learning disability">Learning disability</a> due to <a href="/wiki/Congenital_disorder" class="mw-redirect" title="Congenital disorder">congenital disorder</a><sup id="cite_ref-130" class="reference"><a href="#cite_note-130"><span class="cite-bracket">[</span>130<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Spinal_cord_injury" title="Spinal cord injury">Spinal cord injury</a> repair<sup id="cite_ref-131" class="reference"><a href="#cite_note-131"><span class="cite-bracket">[</span>131<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Heart_infarction" class="mw-redirect" title="Heart infarction">Heart infarction</a><sup id="cite_ref-132" class="reference"><a href="#cite_note-132"><span class="cite-bracket">[</span>132<span class="cite-bracket">]</span></a></sup></li> <li>Anti-<a href="/wiki/Cancer" title="Cancer">cancer</a> treatments<sup id="cite_ref-Steinberg2000_129-1" class="reference"><a href="#cite_note-Steinberg2000-129"><span class="cite-bracket">[</span>129<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Baldness" class="mw-redirect" title="Baldness">Baldness</a> reversal<sup id="cite_ref-133" class="reference"><a href="#cite_note-133"><span class="cite-bracket">[</span>133<span class="cite-bracket">]</span></a></sup></li> <li>Replace missing <a href="/wiki/Teeth" class="mw-redirect" title="Teeth">teeth</a><sup id="cite_ref-134" class="reference"><a href="#cite_note-134"><span class="cite-bracket">[</span>134<span class="cite-bracket">]</span></a></sup></li> <li>Repair <a href="/wiki/Hearing" title="Hearing">hearing</a><sup id="cite_ref-135" class="reference"><a href="#cite_note-135"><span class="cite-bracket">[</span>135<span class="cite-bracket">]</span></a></sup></li> <li>Restore <a href="/wiki/Visual_system" title="Visual system">vision</a><sup id="cite_ref-136" class="reference"><a href="#cite_note-136"><span class="cite-bracket">[</span>136<span class="cite-bracket">]</span></a></sup> and repair damage to the <a href="/wiki/Cornea" title="Cornea">cornea</a><sup id="cite_ref-137" class="reference"><a href="#cite_note-137"><span class="cite-bracket">[</span>137<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Amyotrophic_lateral_sclerosis" class="mw-redirect" title="Amyotrophic lateral sclerosis">Amyotrophic lateral sclerosis</a><sup id="cite_ref-138" class="reference"><a href="#cite_note-138"><span class="cite-bracket">[</span>138<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Crohn%27s_disease" title="Crohn's disease">Crohn's disease</a><sup id="cite_ref-139" class="reference"><a href="#cite_note-139"><span class="cite-bracket">[</span>139<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Wound_healing" title="Wound healing">Wound healing</a><sup id="cite_ref-140" class="reference"><a href="#cite_note-140"><span class="cite-bracket">[</span>140<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Male_infertility" title="Male infertility">Male infertility</a> due to absence of spermatogonial stem cells.<sup id="cite_ref-141" class="reference"><a href="#cite_note-141"><span class="cite-bracket">[</span>141<span class="cite-bracket">]</span></a></sup> In recent studies, scientists have found a way to solve this problem by reprogramming a cell and turning it into a spermatozoon. Other studies have proven the restoration of spermatogenesis by introducing human iPSC cells in mice testicles. This could mean the end of <a href="/wiki/Azoospermia" title="Azoospermia">azoospermia</a>.<sup id="cite_ref-142" class="reference"><a href="#cite_note-142"><span class="cite-bracket">[</span>142<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Female_infertility" title="Female infertility">Female infertility</a>: oocytes made from embryonic stem cells. Scientists have found the ovarian stem cells, a rare type of cells (0.014%) found in the ovary. They could be used as a treatment not only for infertility, but also for premature ovarian insufficiency (POI).<sup id="cite_ref-143" class="reference"><a href="#cite_note-143"><span class="cite-bracket">[</span>143<span class="cite-bracket">]</span></a></sup> New research posted in Science Direct suggests that ovarian follicles could be triggered to grow in the ovarian environment by using stem cells present in <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a>. This study was conducted by infusing human bone marrow stem cells into immune-deficient mice to imrpove fertilization.<sup id="cite_ref-144" class="reference"><a href="#cite_note-144"><span class="cite-bracket">[</span>144<span class="cite-bracket">]</span></a></sup> Another study conducted using mice with damanged ovarian function from <a href="/wiki/Chemotherapy" title="Chemotherapy">chemothearpy</a> found that <a href="/wiki/In_vivo" title="In vivo">in vivo</a> thearpy with bone marrow stem cells can heal the damaged ovaries. <sup id="cite_ref-145" class="reference"><a href="#cite_note-145"><span class="cite-bracket">[</span>145<span class="cite-bracket">]</span></a></sup> Both of these studies are <a href="/wiki/Proof_of_concept" title="Proof of concept">proof-of-concept</a> and need to be furthered tested, but they have the possibility improve fertility for individuals who have POI from chemothearpy treatment.</li> <li>Critical Limb Ischemia<sup id="cite_ref-146" class="reference"><a href="#cite_note-146"><span class="cite-bracket">[</span>146<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Production">Production</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=22" title="Edit section: Production"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Research is underway to develop various sources for stem cells.<sup id="cite_ref-pmid23131007_147-0" class="reference"><a href="#cite_note-pmid23131007-147"><span class="cite-bracket">[</span>147<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Organoids">Organoids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=23" title="Edit section: Organoids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Research is attempting to generating <a href="/wiki/Organoid" title="Organoid">organoids</a> using stem cells, which would allow for further understanding of human development, <a href="/wiki/Organogenesis" title="Organogenesis">organogenesis</a>, and modeling of human diseases.<sup id="cite_ref-pmid25033469_148-0" class="reference"><a href="#cite_note-pmid25033469-148"><span class="cite-bracket">[</span>148<span class="cite-bracket">]</span></a></sup> Engineered ‘synthetic organizer’ (SO) cells can instruct stem cells to grow into specific tissues and organs. The program used native and synthetic <a href="/wiki/Cell_adhesion_molecules" class="mw-redirect" title="Cell adhesion molecules">cell adhesion protein molecules</a> (CAMs) that help make cells sticky. The organizer cells self-assembled around mouse ESCs. These cells were engineered to produce <a href="/wiki/Morphogens" class="mw-redirect" title="Morphogens">morphogens</a> (signaling molecules) that direct cellular development based on their concentration. Delivered morphogens disperse, leaving higher concentrations closer to the source and lower concentrations further away. These gradients signal cells' ultimate roles, such as nerve, skin cell, or connective tissue. The engineered organizer cells were also fitted with a chemical switch that enabled the researchers to turn the delivery of cellular instructions on and off, as well as a ‘suicide switch’ for eliminating the cells when needed. SOs carry spatial and biochemical information, allowing considerable discretion in organoid formation.<sup id="cite_ref-149" class="reference"><a href="#cite_note-149"><span class="cite-bracket">[</span>149<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Risks">Risks</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=24" title="Edit section: Risks"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Hepatotoxicity" title="Hepatotoxicity">Hepatotoxicity</a> and drug-induced liver injury account for a substantial number of failures of new drugs in development and market withdrawal, highlighting the need for screening assays such as stem cell-derived hepatocyte-like cells, that are capable of detecting toxicity early in the <a href="/wiki/Drug_development" title="Drug development">drug development</a> process.<sup id="cite_ref-stem2012_150-0" class="reference"><a href="#cite_note-stem2012-150"><span class="cite-bracket">[</span>150<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dormancy">Dormancy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=25" title="Edit section: Dormancy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In August 2021, researchers in the <a href="/wiki/Princess_Margaret_Cancer_Centre" title="Princess Margaret Cancer Centre">Princess Margaret Cancer Centre</a> at the <a href="/wiki/University_Health_Network" title="University Health Network">University Health Network</a> published their discovery of a dormancy mechanism in key stem cells which could help develop cancer treatments in the future.<sup id="cite_ref-151" class="reference"><a href="#cite_note-151"><span class="cite-bracket">[</span>151<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=26" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 30em;"> <ul><li><a href="/wiki/Cell_bank" title="Cell bank">Cell bank</a></li> <li><a href="/wiki/Human_genome" title="Human genome">Human genome</a></li> <li><a href="/wiki/Meristem" title="Meristem">Meristem</a></li> <li><a href="/wiki/Mesenchymal_stem_cell" title="Mesenchymal stem cell">Mesenchymal stem cell</a></li> <li><a href="/wiki/Ovarian_stem_cell" title="Ovarian stem cell">Ovarian stem cell</a></li> <li><a href="/wiki/Partial_cloning" title="Partial cloning">Partial cloning</a></li> <li><a href="/wiki/Plant_stem_cell" title="Plant stem cell">Plant stem cell</a></li> <li><a href="/wiki/Stem_cell_controversy" title="Stem cell controversy">Stem cell controversy</a></li> <li><a href="/wiki/Stem_cell_marker" title="Stem cell marker">Stem cell marker</a></li> <li><a href="/wiki/Stem_cell_laws_and_policy_in_Iran" title="Stem cell laws and policy in Iran">Stem cell laws and policy in Iran</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=27" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-:7-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-:7_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:7_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFAtalaLanza2012" class="citation book cs1">Atala A, Lanza R (2012). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=wm-K_dKpjBAC&pg=RA1-PA451"><i>Handbook of Stem Cells</i></a>. Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-385943-3" title="Special:BookSources/978-0-12-385943-3"><bdi>978-0-12-385943-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Handbook+of+Stem+Cells&rft.pub=Academic+Press&rft.date=2012&rft.isbn=978-0-12-385943-3&rft.aulast=Atala&rft.aufirst=Anthony&rft.au=Lanza%2C+Robert&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dwm-K_dKpjBAC%26pg%3DRA1-PA451&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStem+cell" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeckerMcCullochTill1963" class="citation journal cs1">Becker AJ, McCulloch EA, Till JE (February 1963). "Cytological demonstration of the clonal nature of spleen colonies derived from transplanted mouse marrow cells". <i>Nature</i>. <b>197</b> (4866): <span class="nowrap">452–</span>454. <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/1963Natur.197..452B">1963Natur.197..452B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F197452a0">10.1038/197452a0</a>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://hdl.handle.net/1807%2F2779">1807/2779</a></span>. <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/0028-0836">0028-0836</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/13970094">13970094</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:11106827">11106827</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Cytological+demonstration+of+the+clonal+nature+of+spleen+colonies+derived+from+transplanted+mouse+marrow+cells&rft.volume=197&rft.issue=4866&rft.pages=%3Cspan+class%3D%22nowrap%22%3E452-%3C%2Fspan%3E454&rft.date=1963-02&rft_id=info%3Ahdl%2F1807%2F2779&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A11106827%23id-name%3DS2CID&rft_id=info%3Abibcode%2F1963Natur.197..452B&rft.issn=0028-0836&rft_id=info%3Adoi%2F10.1038%2F197452a0&rft_id=info%3Apmid%2F13970094&rft.aulast=Becker&rft.aufirst=AJ&rft.au=McCulloch%2C+EA&rft.au=Till%2C+JE&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStem+cell" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSiminovitchMcCullochTill1963" class="citation journal cs1">Siminovitch L, McCulloch EA, Till JE (December 1963). 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"Stem Cell-Based Toxicity Screening: Recent Advances in Hepatocyte Generation". <i>Pharmaceutical Medicine</i>. <b>26</b> (2): <span class="nowrap">85–</span>89. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF03256896">10.1007/BF03256896</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:15893493">15893493</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Pharmaceutical+Medicine&rft.atitle=Stem+Cell-Based+Toxicity+Screening%3A+Recent+Advances+in+Hepatocyte+Generation&rft.volume=26&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E85-%3C%2Fspan%3E89&rft.date=2012-04&rft_id=info%3Adoi%2F10.1007%2FBF03256896&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A15893493%23id-name%3DS2CID&rft.aulast=Greenhough&rft.aufirst=Sebastian&rft.au=Hay%2C+David+C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStem+cell" class="Z3988"></span></span> </li> <li id="cite_note-151"><span class="mw-cite-backlink"><b><a href="#cite_ref-151">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGarcía-PratKaufmannSchneiterVoisin2021" class="citation journal cs1">García-Prat, Laura; Kaufmann, Kerstin B.; Schneiter, Florin; Voisin, Veronique; Murison, Alex; Chen, Jocelyn; Chan-Seng-Yue, Michelle; Gan, Olga I.; McLeod, Jessica L.; Smith, Sabrina A.; Shoong, Michelle C.; Parris, Darrien; Pan, Kristele; Zeng, Andy G.X.; Krivdova, Gabriela; Gupta, Kinam; Takayanagi, Shin-Ichiro; Wagenblast, Elvin; Wang, Weijia; Lupien, Mathieu; Schroeder, Timm; Xie, Stephanie Z.; Dick, John E. (August 2021). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.stem.2021.07.003">"TFEB-mediated endolysosomal activity controls human hematopoietic stem cell fate"</a>. <i>Cell Stem Cell</i>. <b>28</b> (10): 1838–1850.e10. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.stem.2021.07.003">10.1016/j.stem.2021.07.003</a></span>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://hdl.handle.net/20.500.11850%2F510219">20.500.11850/510219</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34343492">34343492</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:236915618">236915618</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cell+Stem+Cell&rft.atitle=TFEB-mediated+endolysosomal+activity+controls+human+hematopoietic+stem+cell+fate&rft.volume=28&rft.issue=10&rft.pages=1838-1850.e10&rft.date=2021-08&rft_id=info%3Ahdl%2F20.500.11850%2F510219&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A236915618%23id-name%3DS2CID&rft_id=info%3Apmid%2F34343492&rft_id=info%3Adoi%2F10.1016%2Fj.stem.2021.07.003&rft.aulast=Garc%C3%ADa-Prat&rft.aufirst=Laura&rft.au=Kaufmann%2C+Kerstin+B.&rft.au=Schneiter%2C+Florin&rft.au=Voisin%2C+Veronique&rft.au=Murison%2C+Alex&rft.au=Chen%2C+Jocelyn&rft.au=Chan-Seng-Yue%2C+Michelle&rft.au=Gan%2C+Olga+I.&rft.au=McLeod%2C+Jessica+L.&rft.au=Smith%2C+Sabrina+A.&rft.au=Shoong%2C+Michelle+C.&rft.au=Parris%2C+Darrien&rft.au=Pan%2C+Kristele&rft.au=Zeng%2C+Andy+G.X.&rft.au=Krivdova%2C+Gabriela&rft.au=Gupta%2C+Kinam&rft.au=Takayanagi%2C+Shin-Ichiro&rft.au=Wagenblast%2C+Elvin&rft.au=Wang%2C+Weijia&rft.au=Lupien%2C+Mathieu&rft.au=Schroeder%2C+Timm&rft.au=Xie%2C+Stephanie+Z.&rft.au=Dick%2C+John+E.&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.stem.2021.07.003&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStem+cell" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=28" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFManzoTorreno-PinaMassignanLapeyre2015" class="citation journal cs1">Manzo, Carlo; Torreno-Pina, Juan A.; Massignan, Pietro; Lapeyre, Gerald J.; Lewenstein, Maciej; Garcia Parajo, Maria F. (25 February 2015). "Weak Ergodicity Breaking of Receptor Motion in Living Cells Stemming from Random Diffusivity". <i>Physical Review X</i>. <b>5</b> (1): 011021. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1407.2552">1407.2552</a></span>. <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/2015PhRvX...5a1021M">2015PhRvX...5a1021M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1103%2FPhysRevX.5.011021">10.1103/PhysRevX.5.011021</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:73582473">73582473</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Physical+Review+X&rft.atitle=Weak+Ergodicity+Breaking+of+Receptor+Motion+in+Living+Cells+Stemming+from+Random+Diffusivity&rft.volume=5&rft.issue=1&rft.pages=011021&rft.date=2015-02-25&rft_id=info%3Aarxiv%2F1407.2552&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A73582473%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1103%2FPhysRevX.5.011021&rft_id=info%3Abibcode%2F2015PhRvX...5a1021M&rft.aulast=Manzo&rft.aufirst=Carlo&rft.au=Torreno-Pina%2C+Juan+A.&rft.au=Massignan%2C+Pietro&rft.au=Lapeyre%2C+Gerald+J.&rft.au=Lewenstein%2C+Maciej&rft.au=Garcia+Parajo%2C+Maria+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStem+cell" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Stem_cell&action=edit&section=29" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></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 .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Commons-logo.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; 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title="Endothelial stem cell">Endothelial stem cell</a></li> <li><a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">Hematopoietic stem cell</a></li> <li><a href="/wiki/Mesenchymal_stem_cell" title="Mesenchymal stem cell">Mesenchymal stem cell</a></li> <li><a href="/wiki/Neural_stem_cell" title="Neural stem cell">Neural stem cell</a></li></ul></li> <li><i>Unipotent</i> <ul><li><a href="/wiki/Precursor_cell" title="Precursor cell">Precursor cell</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related articles</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/Cellular_differentiation" title="Cellular differentiation">Cellular differentiation</a></li> <li><a href="/wiki/Stem_cell_therapy" class="mw-redirect" title="Stem cell therapy">Stem cell therapy</a></li> <li><a href="/wiki/Stem_cell_controversy" title="Stem cell controversy">Stem cell controversy</a></li> <li><a href="/wiki/Stem_cell_line" class="mw-redirect" title="Stem cell line">Stem cell line</a></li> <li><a href="/wiki/Stem_cell_laws" title="Stem cell laws">Stem cell laws</a></li> <li><a href="/wiki/Stem_cell_laws_and_policy_in_the_United_States" title="Stem cell laws and policy in the United States">Stem cell laws and policy in the United States</a></li> <li><a href="/wiki/Epigenetics_in_stem_cell_differentiation" class="mw-redirect" title="Epigenetics in stem cell differentiation">Epigenetics in stem cell differentiation</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" 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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:Breakthrough_of_the_Year" title="Template:Breakthrough of the Year"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Breakthrough_of_the_Year" title="Template talk:Breakthrough of the Year"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Breakthrough_of_the_Year" title="Special:EditPage/Template:Breakthrough of the Year"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Science_Breakthroughs_of_the_Year112" style="font-size:114%;margin:0 4em"><a href="/wiki/Breakthrough_of_the_Year" title="Breakthrough of the Year"><i>Science</i> Breakthroughs of the Year</a></div></th></tr><tr><th scope="row" class="navbox-group" style="text-align:center;;width:1%"><i><a href="/wiki/Science_(journal)" title="Science (journal)">Science</a></i><br />journal</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><span class="nowrap">1996: <a href="/wiki/HIV" title="HIV">HIV</a> understanding</span></li> <li><span class="nowrap">1997: <a href="/wiki/Dolly_(sheep)" title="Dolly (sheep)">Dolly the sheep</a></span></li> <li><span class="nowrap">1998: <a href="/wiki/Accelerating_expansion_of_the_universe" title="Accelerating expansion of the universe">Accelerating universe</a></span></li> <li><span class="nowrap">1999: <a class="mw-selflink selflink">Stem cell</a></span></li> <li><span class="nowrap">2000: <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">Whole genome sequencing</a></span></li> <li><span class="nowrap">2001: <a href="/wiki/Nanocircuitry" title="Nanocircuitry">Nanocircuits</a> or <a href="/wiki/Molecular_scale_electronics" title="Molecular scale electronics">Molecular circuit</a></span></li> <li><span class="nowrap">2002: <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a></span></li> <li><span class="nowrap">2003: <a href="/wiki/Dark_energy" title="Dark energy">Dark energy</a></span></li> <li><span class="nowrap">2004: <a href="/wiki/Spirit_(rover)" title="Spirit (rover)"><i>Spirit</i> rover</a></span></li> <li><span class="nowrap">2005: <a href="/wiki/Evolution" title="Evolution">Evolution</a> in action</span></li> <li><span class="nowrap">2006: <a href="/wiki/Poincar%C3%A9_conjecture" title="Poincaré conjecture">Poincaré conjecture</a> proof</span></li> <li><span class="nowrap">2007: <a href="/wiki/Human_genetic_variation" title="Human genetic variation">Human genetic variation</a></span></li> <li><span class="nowrap">2008: <a href="/wiki/Induced_pluripotent_stem_cell" title="Induced pluripotent stem cell">Cellular reprogramming</a></span></li> <li><span class="nowrap">2009: <i><a href="/wiki/Ardipithecus_ramidus" title="Ardipithecus ramidus">Ardipithecus ramidus</a></i></span></li> <li><span class="nowrap">2010: First <a href="/wiki/Quantum_machine" title="Quantum machine">quantum machine</a></span></li> <li><span class="nowrap">2011: <a href="/wiki/HPTN_052" title="HPTN 052">HPTN 052</a> clinical trial</span></li> <li><span class="nowrap">2012: <a href="/wiki/Higgs_boson" title="Higgs boson">Higgs boson</a> discovery</span></li> <li><span class="nowrap">2013: <a href="/wiki/Cancer_immunotherapy" title="Cancer immunotherapy">Cancer immunotherapy</a></span></li> <li><span class="nowrap">2014: <a href="/wiki/Rosetta_(spacecraft)" title="Rosetta (spacecraft)"><i>Rosetta</i> comet mission</a></span></li> <li><span class="nowrap">2015: <a href="/wiki/CRISPR_gene_editing" title="CRISPR gene editing">CRISPR genome-editing method</a></span></li> <li><span class="nowrap">2016: <a href="/wiki/First_observation_of_gravitational_waves" title="First observation of gravitational waves">First observation</a> of <a href="/wiki/Gravitational_wave" title="Gravitational wave">gravitational waves</a></span></li> <li><span class="nowrap">2017: <a href="/wiki/GW170817" title="GW170817">GW170817</a> (<a href="/wiki/Neutron_star_merger" title="Neutron star merger">neutron star merger</a>)</span></li> <li><span class="nowrap">2018: <a href="/wiki/Single-cell_sequencing" title="Single-cell sequencing">Single-cell sequencing</a></span></li> <li><span class="nowrap">2019: A <a href="/wiki/Black_hole" title="Black hole">black hole</a> <a href="/wiki/Messier_87" title="Messier 87">made visible</a></span></li> <li><span class="nowrap">2020: <a href="/wiki/COVID-19_vaccine" title="COVID-19 vaccine">COVID-19 vaccines</a> developed at record speed</span></li> <li><span class="nowrap">2021: <a href="/wiki/AlphaFold" title="AlphaFold">AI</a> brings <a href="/wiki/Protein_structure_prediction" title="Protein structure prediction">protein structures</a> to all</span></li> <li><span class="nowrap">2022: <a href="/wiki/James_Webb_Space_Telescope" title="James Webb Space Telescope">James Webb Space Telescope</a> debut</span></li> <li><span class="nowrap">2023: <a href="/wiki/GLP-1_receptor_agonist" title="GLP-1 receptor agonist">GLP-1 drugs</a></span></li> <li><span class="nowrap">2024: <a href="/wiki/Lenacapavir" title="Lenacapavir">Lenacapavir</a></span></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Biotechnology389" 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="3"><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:Biotechnology" title="Template:Biotechnology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biotechnology" title="Template talk:Biotechnology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Biotechnology" title="Special:EditPage/Template:Biotechnology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Biotechnology389" style="font-size:114%;margin:0 4em"><a href="/wiki/Biotechnology" title="Biotechnology">Biotechnology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">History</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/History_of_biotechnology" title="History of biotechnology">History of biotechnology</a></li> <li><a href="/wiki/Timeline_of_biotechnology" title="Timeline of biotechnology">Timeline of biotechnology</a></li> <li><a href="/wiki/Competitions_and_prizes_in_biotechnology" title="Competitions and prizes in biotechnology">Competitions and prizes in biotechnology</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="8" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:DNA_replication_split.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/50px-DNA_replication_split.svg.png" decoding="async" width="50" height="101" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/75px-DNA_replication_split.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/DNA_replication_split.svg/100px-DNA_replication_split.svg.png 2x" data-file-width="315" data-file-height="635" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Branches</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/Colors_of_biotechnology" class="mw-redirect" title="Colors of biotechnology">Colors of biotechnology</a></li> <li><a href="/wiki/Industrial_biotechnology" class="mw-redirect" title="Industrial biotechnology">Industrial biotechnology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Biological concepts</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/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Cell_(biology)" title="Cell (biology)">Cell</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a>/<a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Fermentation" title="Fermentation">Fermentation</a></li> <li><a href="/wiki/Gene" title="Gene">Gene</a></li> <li><a href="/wiki/Plasmid" title="Plasmid">Plasmid</a></li> <li><a href="/wiki/Protein" title="Protein">Protein</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General concepts</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/Biotechnology_industrial_park" class="mw-redirect" title="Biotechnology industrial park">Biotechnology industrial park</a></li> <li><a href="/wiki/Biotechnology_products" class="mw-redirect" title="Biotechnology products">Biotechnology products</a></li> <li><a href="/wiki/Biotechnology_law" class="mw-redirect" title="Biotechnology law">Biotechnology law</a></li> <li><a href="/wiki/Green_Revolution" title="Green Revolution">Green Revolution</a></li> <li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li> <li><a href="/wiki/Pharmaceutical_company" class="mw-redirect" title="Pharmaceutical company">Pharmaceutical company</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basic techniques<br /> and tools</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Biology field</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/Bioreactor" title="Bioreactor">Bioreactor</a></li> <li><a href="/wiki/Cell_culture" title="Cell culture">Cell culture</a></li> <li><a href="/wiki/Cultured_meat" title="Cultured meat">Cultured meat</a></li> <li><a href="/wiki/Flow_cytometry" title="Flow cytometry">Flow cytometry</a></li> <li><a href="/wiki/Hybridoma_technology" title="Hybridoma technology">Hybridoma technology</a></li> <li><a href="/wiki/High-performance_liquid_chromatography" title="High-performance liquid chromatography">HPLC</a></li> <li><a href="/wiki/Nuclear_magnetic_resonance" title="Nuclear magnetic resonance">NMR</a></li> <li><a href="/wiki/Spectroscopy" title="Spectroscopy">Spectroscopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Chemical field</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/Centrifugation" title="Centrifugation">Centrifugation</a></li> <li><a href="/wiki/Continuous_stirred-tank_reactor" title="Continuous stirred-tank reactor">CSTR</a></li> <li><a href="/wiki/Crystallization" title="Crystallization">Crystallization</a></li> <li><a href="/wiki/Chromatography" title="Chromatography">Chromatography</a></li> <li><a href="/wiki/Kidney_dialysis" title="Kidney dialysis">Kidney dialysis</a></li> <li><a href="/wiki/Electrophoresis" title="Electrophoresis">Electrophoresis</a></li> <li><a href="/wiki/Extraction_(chemistry)" title="Extraction (chemistry)">Extraction</a></li> <li><a href="/wiki/Fed-batch_culture" title="Fed-batch culture">Fed Batch</a></li> <li><a href="/wiki/Filtration" title="Filtration">Filtration</a></li> <li><a href="/wiki/Plug_flow_reactor_model" title="Plug flow reactor model">PFR</a></li> <li><a href="/wiki/Sedimentation" title="Sedimentation">Sedimentation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Applications</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/Animal_cell_culture" class="mw-redirect" title="Animal cell culture">Animal cell culture</a></li> <li><a href="/wiki/Biofabrication" title="Biofabrication">Biofabrication</a></li> <li><a href="/wiki/Bioinformatics" title="Bioinformatics">Bioinformatics</a></li> <li><a href="/wiki/Biosynthesis" title="Biosynthesis">Biosynthesis</a></li> <li><a href="/wiki/Bionic_architecture" title="Bionic architecture">Bionic architecture</a></li> <li><a href="/wiki/Cell_immunity" class="mw-redirect" title="Cell immunity">Cell immunity</a></li> <li><a href="/wiki/Cloning" title="Cloning">Cloning</a> <ul><li><a href="/wiki/Reproductive_cloning" class="mw-redirect" title="Reproductive cloning">Reproductive cloning</a></li> <li><a href="/wiki/Therapeutic_cloning" class="mw-redirect" title="Therapeutic cloning">Therapeutic cloning</a></li></ul></li> <li><a href="/wiki/Embryology" title="Embryology">Embryology</a></li> <li><a href="/wiki/Environmental_biotechnology" title="Environmental biotechnology">Environmental biotechnology</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a> <ul><li><a href="/wiki/Genetically_modified_organism" title="Genetically modified organism">Genetically modified organism</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular genetics</a></li></ul></li> <li><a href="/wiki/Gene_therapy" title="Gene therapy">Gene therapy</a></li> <li><a href="/wiki/Microbial_biodegradation" title="Microbial biodegradation">Microbial biodegradation</a></li> <li><a href="/wiki/Omics" title="Omics">Omics</a></li> <li><a href="/wiki/Pharmacogenomics" title="Pharmacogenomics">Pharmacogenomics</a></li> <li><a href="/wiki/Stem_cells" class="mw-redirect" title="Stem cells">Stem cells</a></li> <li><a href="/wiki/Telomere" title="Telomere">Telomere</a></li> <li><a href="/wiki/Tissue_culture" title="Tissue culture">Tissue culture</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Interdisciplinary <br />fields</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/Biobased_economy" class="mw-redirect" title="Biobased economy">Bioeconomy</a></li> <li><a href="/wiki/Bioelectronics" title="Bioelectronics">Bioelectronics</a></li> <li><a href="/wiki/Biological_engineering" title="Biological engineering">Bioengineering</a></li> <li><a href="/wiki/Biology" title="Biology">Biology</a></li> <li><a href="/wiki/Biopharmaceutical" title="Biopharmaceutical">Biopharmacology</a></li> <li><a href="/wiki/Biomedical_engineering" title="Biomedical engineering">Biomedical engineering</a></li> <li><a href="/wiki/Biomedicine" title="Biomedicine">Biomedicine</a></li> <li><a href="/wiki/Biomimetics" title="Biomimetics">Biomimetics</a></li> <li><a href="/wiki/Biochemistry" title="Biochemistry">Biochemicals</a></li> <li><a href="/wiki/Biorobotics" title="Biorobotics">Biorobotics</a></li> <li><a href="/wiki/Chemical_engineering" title="Chemical engineering">Chemical engineering</a></li> <li><a href="/wiki/Microbiology" title="Microbiology">Microbiology</a></li> <li><a href="/wiki/Mining" title="Mining">Mining</a></li> <li><a href="/wiki/Molecular_biology" title="Molecular biology">Molecular biology</a></li> <li><a href="/wiki/Nanobiotechnology" title="Nanobiotechnology">Nanobiotechnology</a></li> <li><a href="/wiki/Virology" title="Virology">Virology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/Index_of_biotechnology_articles" title="Index of biotechnology articles">Index of biotechnology articles</a></li> <li><a href="/wiki/List_of_biotechnology_articles" class="mw-redirect" title="List of biotechnology articles">List of biotechnology articles</a></li> <li><a href="/wiki/List_of_largest_biomedical_companies_by_market_capitalization" title="List of largest biomedical companies by market capitalization">List of largest biomedical companies by market capitalization</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Biotechnology" title="Category:Biotechnology">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Biotechnology" class="extiw" title="commons:Category:Biotechnology">Commons</a></b></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link 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href="https://id.loc.gov/authorities/sh85127923">United States</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Cellules souches"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb11964831j">France</a></span></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="Cellules souches"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb11964831j">BnF data</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/01143628">Japan</a></span></li><li><span class="uid"><span class="rt-commentedText tooltip tooltip-dotted" title="kmenové buňky"><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph325855&CON_LNG=ENG">Czech Republic</a></span></span></li><li><span class="uid"><a rel="nofollow" class="external text" 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