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class="vector-toc-list"> </ul> </li> <li id="toc-Receptors" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Receptors</span> </div> </a> <button aria-controls="toc-Receptors-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 Receptors subsection</span> </button> <ul id="toc-Receptors-sublist" class="vector-toc-list"> <li id="toc-Activating_receptors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Activating_receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Activating receptors</span> </div> </a> <ul id="toc-Activating_receptors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inhibitory_receptors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inhibitory_receptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Inhibitory receptors</span> </div> </a> <ul id="toc-Inhibitory_receptors-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Function</span> </div> </a> <button aria-controls="toc-Function-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 Function subsection</span> </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Cytolytic_granule_mediated_cell_apoptosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cytolytic_granule_mediated_cell_apoptosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Cytolytic granule mediated cell apoptosis</span> </div> </a> <ul id="toc-Cytolytic_granule_mediated_cell_apoptosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Antibody-dependent_cell-mediated_cytotoxicity_(ADCC)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Antibody-dependent_cell-mediated_cytotoxicity_(ADCC)"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Antibody-dependent cell-mediated cytotoxicity (ADCC)</span> </div> </a> <ul id="toc-Antibody-dependent_cell-mediated_cytotoxicity_(ADCC)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cytokine-induced_NK_and_Cytotoxic_T_lymphocyte_(CTL)_activation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cytokine-induced_NK_and_Cytotoxic_T_lymphocyte_(CTL)_activation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Cytokine-induced NK and Cytotoxic T lymphocyte (CTL) activation</span> </div> </a> <ul id="toc-Cytokine-induced_NK_and_Cytotoxic_T_lymphocyte_(CTL)_activation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Missing_'self'_hypothesis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Missing_'self'_hypothesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Missing 'self' hypothesis</span> </div> </a> <ul id="toc-Missing_'self'_hypothesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tumor_cell_surveillance" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tumor_cell_surveillance"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Tumor cell surveillance</span> </div> </a> <ul id="toc-Tumor_cell_surveillance-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clearance_of_senescent_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Clearance_of_senescent_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.6</span> <span>Clearance of senescent cells</span> </div> </a> <ul id="toc-Clearance_of_senescent_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Adaptive_features_of_NK_cells—"memory-like",_"adaptive"_and_memory_NK_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Adaptive_features_of_NK_cells—"memory-like",_"adaptive"_and_memory_NK_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.7</span> <span>Adaptive features of NK cells—"memory-like", "adaptive" and memory NK cells</span> </div> </a> <ul id="toc-Adaptive_features_of_NK_cells—"memory-like",_"adaptive"_and_memory_NK_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NK_cell_function_in_pregnancy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#NK_cell_function_in_pregnancy"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.8</span> <span>NK cell function in pregnancy</span> </div> </a> <ul id="toc-NK_cell_function_in_pregnancy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NK_cell_evasion_by_tumor_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#NK_cell_evasion_by_tumor_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.9</span> <span>NK cell evasion by tumor cells</span> </div> </a> <ul id="toc-NK_cell_evasion_by_tumor_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Excessive_NK_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Excessive_NK_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.10</span> <span>Excessive NK cells</span> </div> </a> <ul id="toc-Excessive_NK_cells-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Applications" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Applications"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Applications</span> </div> </a> <button aria-controls="toc-Applications-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 Applications subsection</span> </button> <ul id="toc-Applications-sublist" class="vector-toc-list"> <li id="toc-Anticancer_therapy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anticancer_therapy"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Anticancer therapy</span> </div> </a> <ul id="toc-Anticancer_therapy-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-CAR-NK_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#CAR-NK_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>CAR-NK cells</span> </div> </a> <ul id="toc-CAR-NK_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NK-92_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#NK-92_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>NK-92 cells</span> </div> </a> <ul id="toc-NK-92_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NKG2D-Fc_fusion_protein" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#NKG2D-Fc_fusion_protein"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>NKG2D-Fc fusion protein</span> </div> </a> <ul id="toc-NKG2D-Fc_fusion_protein-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-TLR_ligands" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#TLR_ligands"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.5</span> <span>TLR ligands</span> </div> </a> <ul id="toc-TLR_ligands-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-New_findings" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#New_findings"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>New findings</span> </div> </a> <button aria-controls="toc-New_findings-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 New findings subsection</span> </button> <ul id="toc-New_findings-sublist" class="vector-toc-list"> <li id="toc-Innate_resistance_to_HIV" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Innate_resistance_to_HIV"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Innate resistance to HIV</span> </div> </a> <ul id="toc-Innate_resistance_to_HIV-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tissue-resident_NK_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tissue-resident_NK_cells"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Tissue-resident NK cells</span> </div> </a> <ul id="toc-Tissue-resident_NK_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Adaptive_NK_cells_against_leukemia_targets" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Adaptive_NK_cells_against_leukemia_targets"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Adaptive NK cells against leukemia targets</span> </div> </a> <ul id="toc-Adaptive_NK_cells_against_leukemia_targets-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">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-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">9</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">10</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Natural killer 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 43 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-43" 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">43 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AE%D9%84%D9%8A%D8%A9_%D9%81%D8%A7%D8%AA%D9%83%D8%A9_%D8%B7%D8%A8%D9%8A%D8%B9%D9%8A%D8%A9" title="خلية فاتكة طبيعية – Arabic" lang="ar" hreflang="ar" data-title="خلية فاتكة طبيعية" 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/T%C9%99bii_killer_h%C3%BCceyr%C9%99l%C9%99ri" title="Təbii killer hüceyrələri – Azerbaijani" lang="az" hreflang="az" data-title="Təbii killer hüceyrələri" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/NK-%D0%BA%D0%BB%D0%B5%D1%82%D0%BA%D0%B0" title="NK-клетка – Bulgarian" lang="bg" hreflang="bg" data-title="NK-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/%C4%86elije_ubice" title="Ćelije ubice – Bosnian" lang="bs" hreflang="bs" data-title="Ćelije ubice" 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_NK" title="Cèl·lula NK – Catalan" lang="ca" hreflang="ca" data-title="Cèl·lula NK" 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/NK_bu%C5%88ka" title="NK buňka – Czech" lang="cs" hreflang="cs" data-title="NK 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/Naturlig_dr%C3%A6bercelle" title="Naturlig dræbercelle – Danish" lang="da" hreflang="da" data-title="Naturlig dræbercelle" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/NK-Zelle" title="NK-Zelle – German" lang="de" hreflang="de" data-title="NK-Zelle" 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/Loomulikud_tappurrakud" title="Loomulikud tappurrakud – Estonian" lang="et" hreflang="et" data-title="Loomulikud tappurrakud" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/C%C3%A9lula_NK" title="Célula NK – Spanish" lang="es" hreflang="es" data-title="Célula NK" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/NK_zelula" title="NK zelula – Basque" lang="eu" hreflang="eu" data-title="NK zelula" 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_%DA%A9%D8%B4%D9%86%D8%AF%D9%87_%D8%B7%D8%A8%DB%8C%D8%B9%DB%8C" title="سلول کشنده طبیعی – Persian" lang="fa" hreflang="fa" data-title="سلول کشنده طبیعی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Lymphocyte_NK" title="Lymphocyte NK – French" lang="fr" hreflang="fr" data-title="Lymphocyte NK" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/C%C3%A9lula_NK" title="Célula NK – Galician" lang="gl" hreflang="gl" data-title="Célula NK" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%9E%90%EC%97%B0_%EC%82%B4%ED%95%B4_%EC%84%B8%ED%8F%AC" title="자연 살해 세포 – Korean" lang="ko" hreflang="ko" data-title="자연 살해 세포" 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/%D4%B2%D5%B6%D5%A1%D5%AF%D5%A1%D5%B6_%D6%84%D5%AB%D5%AC%D5%AC%D5%A5%D6%80%D5%B6%D5%A5%D6%80" title="Բնական քիլլերներ – Armenian" lang="hy" hreflang="hy" data-title="Բնական քիլլերներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Sel_pembunuh_alami" title="Sel pembunuh alami – Indonesian" lang="id" hreflang="id" data-title="Sel pembunuh alami" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Linfocita_NK" title="Linfocita NK – Italian" lang="it" hreflang="it" data-title="Linfocita NK" 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%94%D7%A8%D7%92_%D7%98%D7%91%D7%A2%D7%99" title="תא הרג טבעי – Hebrew" lang="he" hreflang="he" data-title="תא הרג טבעי" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/NK_l%C4%85stel%C4%97" title="NK ląstelė – Lithuanian" lang="lt" hreflang="lt" data-title="NK ląstelė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Sel_pemusnah_semula_jadi" title="Sel pemusnah semula jadi – Malay" lang="ms" hreflang="ms" data-title="Sel pemusnah semula jadi" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/NK-cel" title="NK-cel – Dutch" lang="nl" hreflang="nl" data-title="NK-cel" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%8A%E3%83%81%E3%83%A5%E3%83%A9%E3%83%AB%E3%82%AD%E3%83%A9%E3%83%BC%E7%B4%B0%E8%83%9E" title="ナチュラルキラー細胞 – Japanese" lang="ja" hreflang="ja" data-title="ナチュラルキラー細胞" 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/Naturlige_dreperceller" title="Naturlige dreperceller – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Naturlige dreperceller" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" 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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_exterminadora_natural" title="Célula exterminadora natural – Portuguese" lang="pt" hreflang="pt" data-title="Célula exterminadora natural" 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/Celul%C4%83_Natural_Killer" title="Celulă Natural Killer – Romanian" lang="ro" hreflang="ro" data-title="Celulă Natural Killer" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a 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.hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Not to be confused with <a href="/wiki/Natural_killer_T_cell" title="Natural killer T cell">Natural killer T cell</a>.</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:beige">Natural killer cell</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Human_Natural_Killer_Cell_(29120480442).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Human_Natural_Killer_Cell_%2829120480442%29.jpg/250px-Human_Natural_Killer_Cell_%2829120480442%29.jpg" decoding="async" width="250" height="188" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Human_Natural_Killer_Cell_%2829120480442%29.jpg/375px-Human_Natural_Killer_Cell_%2829120480442%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/36/Human_Natural_Killer_Cell_%2829120480442%29.jpg/500px-Human_Natural_Killer_Cell_%2829120480442%29.jpg 2x" data-file-width="2560" data-file-height="1920" /></a></span><div class="infobox-caption">Human natural killer cell, colorized scanning electron micrograph</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Organ_system" title="Organ system">System</a></th><td class="infobox-data"><a href="/wiki/Immune_system" title="Immune system">Immune system</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Function</th><td class="infobox-data">Cytotoxic <a href="/wiki/Lymphocyte" title="Lymphocyte">lymphocyte</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/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=D007694">D007694</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%2Ffma63147">63147</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_microanatomy" title="Anatomical terms of microanatomy">Anatomical terms of microanatomy</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q332181" class="extiw" title="d:Q332181">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p><b>Natural killer cells</b>, also known as <b>NK cells</b>, are a type of <a href="/wiki/Cytotoxic" class="mw-redirect" title="Cytotoxic">cytotoxic</a> <a href="/wiki/Lymphocyte" title="Lymphocyte">lymphocyte</a> critical to the <a href="/wiki/Innate_immune_system" title="Innate immune system">innate immune system</a>. They are a kind of large granular lymphocytes<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> (LGL), and belong to the rapidly expanding family of known <a href="/wiki/Innate_lymphoid_cell" title="Innate lymphoid cell">innate lymphoid cells</a> (ILC) and represent 5–20% of all circulating lymphocytes in humans.<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> The role of NK cells is analogous to that of <a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">cytotoxic T cells</a> in the vertebrate <a href="/wiki/Adaptive_immune_response" class="mw-redirect" title="Adaptive immune response">adaptive immune response</a>. NK cells provide rapid responses to <a href="/wiki/Virus" title="Virus">virus</a>-infected cells, stressed cells, tumor cells, and other intracellular pathogens based on signals from several activating and inhibitory receptors. Most <a href="/wiki/Immune_cells" class="mw-redirect" title="Immune cells">immune cells</a> detect the antigen presented on <a href="/wiki/MHC_class_I" title="MHC class I">major histocompatibility complex I</a> (MHC-I) on infected cell surfaces, but NK cells can recognize and kill stressed cells in the absence of <a href="/wiki/Antibodies" class="mw-redirect" title="Antibodies">antibodies</a> and MHC, allowing for a much faster immune reaction. They were named "natural killers" because of the notion that they do not require activation to kill cells that are missing "self" markers of <a href="/wiki/MHC_class_I" title="MHC class I">MHC class I</a>.<sup id="cite_ref-Vivier2011_4-0" class="reference"><a href="#cite_note-Vivier2011-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This role is especially important because harmful cells that are missing MHC I markers cannot be detected and destroyed by other immune cells, such as T lymphocyte cells. </p><p>NK cells can be identified by the presence of <a href="/wiki/Neural_cell_adhesion_molecule" title="Neural cell adhesion molecule">CD56</a> and the absence of <a href="/wiki/CD3_(immunology)" title="CD3 (immunology)">CD3</a> (CD56<sup>+</sup>, CD3<sup>−</sup>).<sup id="cite_ref-pmid32477340_5-0" class="reference"><a href="#cite_note-pmid32477340-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> NK cells differentiate from <a href="/wiki/Interleukin-7_receptor-%CE%B1" title="Interleukin-7 receptor-α">CD127<sup>+</sup></a> common <a href="/wiki/Innate_lymphoid_cell" title="Innate lymphoid cell">innate lymphoid</a> progenitor,<sup id="cite_ref-:7_6-0" class="reference"><a href="#cite_note-:7-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> which is downstream of the <a href="/wiki/Lymphopoiesis#The_old_model:_lymphoid_vs_myeloid" title="Lymphopoiesis">common lymphoid progenitor</a> from which <a href="/wiki/B_cell" title="B cell">B</a> and <a href="/wiki/T_cell" title="T cell">T lymphocytes</a> are also derived.<sup id="cite_ref-:7_6-1" class="reference"><a href="#cite_note-:7-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> NK cells are known to differentiate and mature in the <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a>, <a href="/wiki/Lymph_node" title="Lymph node">lymph nodes</a>, <a href="/wiki/Spleen" title="Spleen">spleen</a>, <a href="/wiki/Tonsil" title="Tonsil">tonsils</a>, and <a href="/wiki/Thymus" title="Thymus">thymus</a>, where they then enter into the circulation.<sup id="cite_ref-Lannello2008_8-0" class="reference"><a href="#cite_note-Lannello2008-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> NK cells differ from <a href="/wiki/Natural_killer_T_cell" title="Natural killer T cell">natural killer T cells</a> (NKTs) phenotypically, by origin and by respective effector functions; often, NKT cell activity promotes NK cell activity by secreting <a href="/wiki/IFN%CE%B3" class="mw-redirect" title="IFNγ">interferon gamma</a>. In contrast to NKT cells, NK cells do not express <a href="/wiki/T-cell_antigen_receptors" class="mw-redirect" title="T-cell antigen receptors">T-cell antigen receptors</a> (TCR) or pan T marker <a href="/wiki/CD3_(immunology)" title="CD3 (immunology)">CD3</a> or surface <a href="/wiki/Immunoglobulin" class="mw-redirect" title="Immunoglobulin">immunoglobulins</a> (Ig) <a href="/wiki/B_cell_receptor" class="mw-redirect" title="B cell receptor">B cell receptors</a>, but they usually express the surface markers <a href="/wiki/CD16" title="CD16">CD16</a> (FcγRIII) and <a href="/wiki/CD57" class="mw-redirect" title="CD57">CD57</a> in humans, NK1.1 or NK1.2 in <a href="/wiki/C57BL/6" title="C57BL/6">C57BL/6</a> <a href="/wiki/Mouse" title="Mouse">mice</a>. The <a href="/wiki/NKp46" class="mw-redirect" title="NKp46">NKp46</a> cell surface marker constitutes, at the moment, another NK cell marker of preference being expressed in both humans, several strains of mice (including <a href="/wiki/BALB/c" title="BALB/c">BALB/c mice</a>) and in three common monkey species.<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><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>Outside of <a href="/wiki/Innate_immunity" class="mw-redirect" title="Innate immunity">innate immunity</a>, both activating and inhibitory NK cell receptors play important functional roles in self tolerance and the sustaining of NK cell activity. NK cells also play a role in the <a href="/wiki/Adaptive_immune_response" class="mw-redirect" title="Adaptive immune response">adaptive immune response</a>:<sup id="cite_ref-Arina2007_11-0" class="reference"><a href="#cite_note-Arina2007-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> numerous experiments have demonstrated their ability to readily adjust to the immediate environment and formulate antigen-specific <a href="/wiki/Immunological_memory" title="Immunological memory">immunological memory</a>, fundamental for responding to secondary infections with the same antigen.<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> The role of NK cells in both the innate and adaptive immune responses is becoming increasingly important in research using NK cell activity as a potential <a href="/wiki/Cancer_therapy" class="mw-redirect" title="Cancer therapy">cancer therapy</a> and HIV therapy.<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><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> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Early_history">Early history</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=1" title="Edit section: Early history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In early experiments on cell-mediated cytotoxicity against tumor target cells, both in cancer patients and animal models, investigators consistently observed what was termed a "natural" reactivity; that is, a certain population of cells seemed to be able to destroy tumor cells without having been previously sensitized to them. The first published study to assert that untreated lymphoid cells were able to confer a natural immunity to tumors was performed by Dr. Henry Smith at the University of Leeds School of Medicine in 1966,<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> leading to the conclusion that the "phenomenon appear[ed] to be an expression of defense mechanisms to tumor growth present in normal mice." Other researchers had also made similar observations, but as these discoveries were inconsistent with the established model at the time, many initially considered these observations to be artifacts.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>By 1973, 'natural killing' activity was established across a wide variety of species, and the existence of a separate lineage of cells possessing this ability was postulated. The discovery that a unique type of lymphocyte was responsible for "natural" or spontaneous cytotoxicity was made in the early 1970s by doctoral student Rolf Kiessling and postdoctoral fellow Hugh Pross, in the mouse,<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> and by Hugh Pross and doctoral student Mikael Jondal in the human.<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> The mouse and human work was carried out under the supervision of professors <a href="/wiki/Eva_Klein" title="Eva Klein">Eva Klein</a> and Hans Wigzell, respectively, of the Karolinska Institute, Stockholm. Kiessling's research involved the well-characterized ability of T lymphocytes to attack tumor cells which they had been previously immunized against. Pross and Jondal were studying cell-mediated cytotoxicity in normal human blood and the effect of the removal of various receptor-bearing cells on this cytotoxicity. Later that same year, <a href="/wiki/Ronald_B._Herberman,_M.D." class="mw-redirect" title="Ronald B. Herberman, M.D.">Ronald Herberman</a> published similar data with respect to the unique nature of the mouse effector cell.<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> The human data were confirmed, for the most part, by West <i>et al.</i><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> using similar techniques and the same erythroleukemic target cell line, <a href="/wiki/K562_cells" title="K562 cells">K562</a>. K562 is highly sensitive to lysis by human NK cells and, over the decades, the K562 <sup>51</sup>chromium-release assay has become the most commonly used assay to detect human NK functional activity.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Its almost universal use has meant that experimental data can be compared easily by different laboratories around the world. </p><p>Using discontinuous density centrifugation, and later <a href="/wiki/Monoclonal_antibodies" class="mw-redirect" title="Monoclonal antibodies">monoclonal antibodies</a>, natural killing ability was mapped to the subset of large, granular lymphocytes known today as NK cells. The demonstration that density gradient-isolated large granular lymphocytes were responsible for human NK activity, made by Timonen and Saksela in 1980,<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> was the first time that NK cells had been visualized microscopically, and was a major breakthrough in the field. </p> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=2" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>NK cells can be classified as CD56<sup>bright</sup> or CD56<sup>dim</sup>.<sup id="cite_ref-pmid32545516_24-0" class="reference"><a href="#cite_note-pmid32545516-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid32762681_25-0" class="reference"><a href="#cite_note-pmid32762681-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid32477340_5-1" class="reference"><a href="#cite_note-pmid32477340-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> CD56<sup>bright</sup> NK cells are similar to <a href="/wiki/T_helper_cell" title="T helper cell">T helper cells</a> in exerting their influence by releasing <a href="/wiki/Cytokine" title="Cytokine">cytokines</a>.<sup id="cite_ref-pmid32762681_25-1" class="reference"><a href="#cite_note-pmid32762681-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> CD56<sup>bright</sup> NK cells constitute the majority of NK cells, being found in bone marrow, secondary lymphoid tissue, liver, and skin.<sup id="cite_ref-pmid32477340_5-2" class="reference"><a href="#cite_note-pmid32477340-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> CD56<sup>bright</sup> NK cells are characterized by their preferential killing of highly proliferative cells,<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> and thus might have an immunoregulatory role. CD56<sup>dim</sup> NK cells are primarily found in the <a href="/wiki/Venous_blood" title="Venous blood">peripheral blood</a>,<sup id="cite_ref-pmid32477340_5-3" class="reference"><a href="#cite_note-pmid32477340-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> and are characterized by their cell killing ability.<sup id="cite_ref-pmid32762681_25-2" class="reference"><a href="#cite_note-pmid32762681-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> CD56<sup>dim</sup> NK cells are always <a href="/wiki/CD16" title="CD16">CD16</a> positive (CD16 is the key mediator of <a href="/wiki/Antibody-dependent_cellular_cytotoxicity" title="Antibody-dependent cellular cytotoxicity">antibody-dependent cellular cytotoxicity</a>, or ADCC).<sup id="cite_ref-pmid32762681_25-3" class="reference"><a href="#cite_note-pmid32762681-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> CD56<sup>bright</sup> can transition into CD56<sup>dim</sup> by acquiring CD16.<sup id="cite_ref-pmid32477340_5-4" class="reference"><a href="#cite_note-pmid32477340-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>NK cells can eliminate virus-infected cells via CD16-mediated ADCC.<sup id="cite_ref-pmid32655581_27-0" class="reference"><a href="#cite_note-pmid32655581-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> All <a href="/wiki/Coronavirus_disease_2019" class="mw-redirect" title="Coronavirus disease 2019">coronavirus disease 2019</a> (COVID-19) patients show depleted CD56<sup>bright</sup> NK cells, but CD56<sup>dim</sup> is only depleted in patients with severe COVID-19.<sup id="cite_ref-pmid32655581_27-1" class="reference"><a href="#cite_note-pmid32655581-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Receptors">Receptors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=3" title="Edit section: Receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:HLA-Cw4.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/HLA-Cw4.png/150px-HLA-Cw4.png" decoding="async" width="150" height="151" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/HLA-Cw4.png/225px-HLA-Cw4.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/13/HLA-Cw4.png/300px-HLA-Cw4.png 2x" data-file-width="548" data-file-height="550" /></a><figcaption>The HLA ligand for KIR</figcaption></figure> <p>NK cell receptors can also be differentiated based on function. Natural <a href="/wiki/Cytotoxicity" title="Cytotoxicity">cytotoxicity</a> receptors directly induce <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> (cell death) after binding to <a href="/wiki/Fas_ligand" title="Fas ligand">Fas ligand</a> that directly indicate infection of a cell. The MHC-independent receptors (described above) use an alternate pathway to induce apoptosis in infected cells. Natural killer cell activation is determined by the balance of inhibitory and activating receptor stimulation. For example, if the inhibitory receptor signaling is more prominent, then NK cell activity will be inhibited; similarly, if the activating signal is dominant, then NK cell activation will result.<sup id="cite_ref-Terunuma2008_28-0" class="reference"><a href="#cite_note-Terunuma2008-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:PDB_1hq8_EBI.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/PDB_1hq8_EBI.jpg/150px-PDB_1hq8_EBI.jpg" decoding="async" width="150" height="113" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/PDB_1hq8_EBI.jpg/225px-PDB_1hq8_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/PDB_1hq8_EBI.jpg/300px-PDB_1hq8_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a><figcaption>Protein structure of NKG2D</figcaption></figure> <p>NK cell receptor types (with inhibitory, as well as some activating members) are differentiated by structure, with a few examples to follow: </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:PDB_1hkf_EBI.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/PDB_1hkf_EBI.jpg/150px-PDB_1hkf_EBI.jpg" decoding="async" width="150" height="113" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/PDB_1hkf_EBI.jpg/225px-PDB_1hkf_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/PDB_1hkf_EBI.jpg/300px-PDB_1hkf_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a><figcaption>Protein structure of NKp44</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Activating_receptors">Activating receptors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=4" title="Edit section: Activating receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><b><a href="/wiki/Ly49" title="Ly49">Ly49</a> </b>(homodimers), relatively ancient, C-type <a href="/wiki/Lectin" title="Lectin">lectin</a> family receptors, are of multigenic presence in mice, while humans have only one <a href="/wiki/Pseudogene" title="Pseudogene">pseudogenic</a> Ly49, the receptor for classical (polymorphic) <a href="/wiki/MHC_I" class="mw-redirect" title="MHC I">MHC I</a> molecules.</li> <li><b>NCR</b> (natural cytotoxicity receptors), type 1 transmembrane proteins of the immunoglobulin superfamily, upon stimulation mediate NK killing and release of <a href="/wiki/IFN%CE%B3" class="mw-redirect" title="IFNγ">IFNγ</a>. They bind viral ligands such as hemagglutinins and hemagglutinin neuraminidases, some bacterial ligands and cellular ligands related to tumour growth such as <a href="/wiki/PCNA" class="mw-redirect" title="PCNA">PCNA</a>.</li> <li><b><a href="/wiki/CD16" title="CD16">CD16</a> (FcγIIIA)</b> plays a role in <a href="/wiki/Antibody-dependent_cell-mediated_cytotoxicity" class="mw-redirect" title="Antibody-dependent cell-mediated cytotoxicity">antibody-dependent cell-mediated cytotoxicity</a>; in particular, they bind <a href="/wiki/Immunoglobulin_G" title="Immunoglobulin G">immunoglobulin G</a>.</li> <li><b><a href="/wiki/Toll-like_receptor" title="Toll-like receptor">TLR</a></b> – Toll-like receptors are receptors that belong in the group of <a href="/wiki/Pattern_recognition_receptor" title="Pattern recognition receptor">pattern recognition receptors (PRR)</a> which are typical for the cells of <a href="/wiki/Innate_immunity" class="mw-redirect" title="Innate immunity">innate immunity</a> but are expressed also on NK cells. They recognize <a href="/wiki/Pathogen-associated_molecular_pattern" title="Pathogen-associated molecular pattern">PAMPs</a> (pathogen-associated molecular patterns) and <a href="/wiki/Damage-associated_molecular_pattern" title="Damage-associated molecular pattern">DAMPs</a> (damage-associated molecular patterns) as their ligands. These receptors are crucial for the induction of the <a href="/wiki/Immune_response" title="Immune response">immune response</a>. TLR induction amplifies the immune response by promoting the production of inflammatory <a href="/wiki/Cytokine" title="Cytokine">cytokines</a> and <a href="/wiki/Chemokine" title="Chemokine">chemokines</a> and ultimately leads to the activation of NK cell effector functions.<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> So NK cells directly reacts to the presence of <a href="/wiki/Pathogen" title="Pathogen">pathogens</a> in its surroundings. Apart from <a href="/wiki/Toll-like_receptor_10" title="Toll-like receptor 10">TLR-10</a> NK cells express all of the human TLR although in various levels. NK cells express high levels of <a href="/wiki/Toll-like_receptor_1" title="Toll-like receptor 1">TLR-1</a>, moderate levels of <a href="/wiki/Toll-like_receptor_2" title="Toll-like receptor 2">TLR-2</a>, <a href="/wiki/Toll-like_receptor_3" title="Toll-like receptor 3">TLR-3</a>, <a href="/wiki/Toll-like_receptor_5" title="Toll-like receptor 5">TLR-5</a> and <a href="/wiki/Toll-like_receptor_6" title="Toll-like receptor 6">TLR-6</a>, low levels of <a href="/wiki/Toll-like_receptor_4" title="Toll-like receptor 4">TLR-4</a>, <a href="/wiki/Toll-like_receptor_8" title="Toll-like receptor 8">TLR-8</a> and TLR-9 and very low levels of <a href="/wiki/Toll-like_receptor_7" title="Toll-like receptor 7">TLR-7</a>.<sup id="cite_ref-:4_30-0" class="reference"><a href="#cite_note-:4-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> TLR receptors are constitutionally expressed independently of their state of activation and they cooperate with cytokines and chemokines on the activation of the natural killer cells.<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> These receptors are expressed extracellularly on the cell surface or endosomally inside the <a href="/wiki/Endosome" title="Endosome">endosomes</a>. Apart from TLR-3 and TLR-4, all TLR signal through adaptor protein <a href="/wiki/MYD88" title="MYD88">MyD88</a> which ultimately leads mainly to the activation of <a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-κB</a>. TLR-3 signals through the adaptor protein <a href="/wiki/TRIF" class="mw-redirect" title="TRIF">TRIF</a> and TLR-4 can switch between signaling through MyD88 and TRIF respectively. Induction of different TLR leads to distinct activation of NK cell functions.<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></li></ul> <div class="mw-heading mw-heading3"><h3 id="Inhibitory_receptors">Inhibitory receptors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=5" title="Edit section: Inhibitory receptors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Killer-cell_immunoglobulin-like_receptor" title="Killer-cell immunoglobulin-like receptor">Killer-cell immunoglobulin-like receptors</a> (KIRs) belong to a multigene family of more recently <a href="/wiki/Evolution" title="Evolution">evolved</a> Ig-like extracellular domain receptors; they are present in nonhuman primates, and are the main receptors for both classical MHC I (<a href="/wiki/HLA-A" title="HLA-A">HLA-A</a>, <a href="/wiki/HLA-B" title="HLA-B">HLA-B</a>, <a href="/wiki/HLA-C" title="HLA-C">HLA-C</a>) and nonclassical Mamu-G (<a href="/wiki/HLA-G" title="HLA-G">HLA-G</a>) in primates. Some KIRs are specific for certain HLA subtypes. Most KIRs are inhibitory and dominant. Regular cells express MHC class 1, so are recognised by KIR receptors and NK cell killing is inhibited.<sup id="cite_ref-Lannello2008_8-1" class="reference"><a href="#cite_note-Lannello2008-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li> <li><b><a href="/wiki/CD94/NKG2" title="CD94/NKG2">CD94/NKG2</a></b> (heterodimers), a C-type lectin family receptor, is conserved in both rodents and primates and identifies nonclassical (also nonpolymorphic) MHC I molecules such as <a href="/wiki/HLA-E" title="HLA-E">HLA-E</a>. Expression of HLA-E at the cell surface is dependent on the presence of nonamer peptide epitope derived from the signal sequence of classical MHC class I molecules, which is generated by the sequential action of <a href="/wiki/Signal_peptide_peptidase" title="Signal peptide peptidase">signal peptide peptidase</a> and the <a href="/wiki/Proteasome" title="Proteasome">proteasome</a>. Though indirect, this is a way to survey the levels of classical (polymorphic) HLA molecules.</li> <li><b>ILT</b> or <b>LIR</b> (immunoglobulin-like receptor) – are recently discovered members of the Ig receptor family.</li> <li><b>Ly49</b> (homodimers) have both activating and inhibitory isoforms. They are highly polymorphic on the population level; though they are structurally unrelated to KIRs, they are the functional homologues of KIRs in mice, including the expression pattern. Ly49s are receptor for classical (polymorphic) MHC I molecules.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=6" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Cytolytic_granule_mediated_cell_apoptosis">Cytolytic granule mediated cell apoptosis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=7" title="Edit section: Cytolytic granule mediated cell apoptosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>NK cells are <a href="/wiki/Cytotoxic" class="mw-redirect" title="Cytotoxic">cytotoxic</a>; small <a href="/wiki/Granulocytes" class="mw-redirect" title="Granulocytes">granules</a> in their <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a> contain proteins such as <a href="/wiki/Perforin" class="mw-redirect" title="Perforin">perforin</a> and <a href="/wiki/Protease" title="Protease">proteases</a> known as <a href="/wiki/Granzyme" title="Granzyme">granzymes</a>. Upon release in close proximity to a cell slated for killing, perforin forms pores in the <a href="/wiki/Cell_membrane" title="Cell membrane">cell membrane</a> of the target cell, creating an aqueous channel through which the granzymes and associated molecules can enter, inducing either <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> or osmotic cell lysis. The distinction between apoptosis and cell <a href="/wiki/Lysis" title="Lysis">lysis</a> is important in <a href="/wiki/Immunology" title="Immunology">immunology</a>: lysing a virus-infected cell could potentially release the <a href="/wiki/Virion" class="mw-redirect" title="Virion">virions</a>, whereas apoptosis leads to destruction of the virus inside. <a href="/wiki/Alpha_defensin" title="Alpha defensin">α-defensins</a>, antimicrobial molecules, are also secreted by NK cells, and directly kill bacteria by disrupting their cell walls in a manner analogous to that of <a href="/wiki/Neutrophil" title="Neutrophil">neutrophils</a>.<sup id="cite_ref-Lannello2008_8-2" class="reference"><a href="#cite_note-Lannello2008-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Antibody-dependent_cell-mediated_cytotoxicity_(ADCC)"><span id="Antibody-dependent_cell-mediated_cytotoxicity_.28ADCC.29"></span>Antibody-dependent cell-mediated cytotoxicity (ADCC)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=8" title="Edit section: Antibody-dependent cell-mediated cytotoxicity (ADCC)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Infected cells are routinely <a href="/wiki/Opsonized" class="mw-redirect" title="Opsonized">opsonized</a> with antibodies for detection by immune cells. Antibodies that bind to antigens can be recognised by FcγRIII (<a href="/wiki/CD16" title="CD16">CD16</a>) receptors expressed on NK cells, resulting in NK activation, release of cytolytic granules and consequent cell <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>. This is a major killing mechanism of some <a href="/wiki/Monoclonal_antibodies" class="mw-redirect" title="Monoclonal antibodies">monoclonal antibodies</a> like <a href="/wiki/Rituximab" title="Rituximab">rituximab (Rituxan)</a>, <a href="/wiki/Ofatumumab" title="Ofatumumab"> ofatumumab (Azzera)</a>, and others. The contribution of antibody-dependent cell-mediated cytotoxicity to tumor cell killing can be measured with a specific test that uses <a href="/wiki/NK-92" title="NK-92">NK-92</a>, an immortal line of NK-like cells licensed to <a rel="nofollow" class="external text" href="https://nantkwest.com">NantKwest, Inc.</a>: the response of <a rel="nofollow" class="external text" href="https://nantkwest.com">NK-92 cells</a> that have been transfected with a high-affinity <a href="/wiki/Fc_receptor" title="Fc receptor">Fc receptor</a> are compared to that of the "wild type" NK-92 which does not express the Fc receptor.<sup id="cite_ref-Smyth2002_33-0" class="reference"><a href="#cite_note-Smyth2002-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cytokine-induced_NK_and_Cytotoxic_T_lymphocyte_(CTL)_activation"><span id="Cytokine-induced_NK_and_Cytotoxic_T_lymphocyte_.28CTL.29_activation"></span>Cytokine-induced NK and Cytotoxic T lymphocyte (CTL) activation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=9" title="Edit section: Cytokine-induced NK and Cytotoxic T lymphocyte (CTL) activation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a> play a crucial role in NK cell activation. As these are <a href="/wiki/Stress_(biology)" title="Stress (biology)">stress</a> molecules released by cells upon viral infection, they serve to signal to the NK cell the presence of <a href="/wiki/Virus" title="Virus">viral pathogens</a> in the affected area. Cytokines involved in NK activation include <a href="/wiki/Interleukin_12" title="Interleukin 12">IL-12</a>, <a href="/wiki/Interleukin_15" title="Interleukin 15">IL-15</a>, <a href="/wiki/Interleukin_18" title="Interleukin 18">IL-18</a>, <a href="/wiki/Interleukin_2" title="Interleukin 2">IL-2</a>, and <a href="/wiki/CCL5" title="CCL5">CCL5</a>. NK cells are activated in response to interferons or macrophage-derived cytokines. They serve to contain viral infections while the adaptive immune response generates antigen-specific <a href="/wiki/Cytotoxic_T_cells" class="mw-redirect" title="Cytotoxic T cells">cytotoxic T cells</a> that can clear the infection. NK cells work to control viral infections by secreting <a href="/wiki/IFN%CE%B3" class="mw-redirect" title="IFNγ">IFNγ</a> and <a href="/wiki/TNF%CE%B1" class="mw-redirect" title="TNFα">TNFα</a>. IFNγ activates macrophages for phagocytosis and lysis, and TNFα acts to promote direct NK tumor cell killing. Patients deficient in NK cells prove to be highly susceptible to early phases of herpes virus infection. [Citation needed] </p> <div class="mw-heading mw-heading3"><h3 id="Missing_'self'_hypothesis"><span id="Missing_.27self.27_hypothesis"></span>Missing 'self' hypothesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=10" title="Edit section: Missing 'self' hypothesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Missing_self.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Missing_self.svg/400px-Missing_self.svg.png" decoding="async" width="400" height="336" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Missing_self.svg/600px-Missing_self.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b7/Missing_self.svg/800px-Missing_self.svg.png 2x" data-file-width="1504" data-file-height="1264" /></a><figcaption> Schematic diagram indicating the complementary activities of <a href="/wiki/Cytotoxic_T_cells" class="mw-redirect" title="Cytotoxic T cells">cytotoxic T cells</a> and NK cells</figcaption></figure> <p>For NK cells to defend the body against <a href="/wiki/Viruses" class="mw-redirect" title="Viruses">viruses</a> and other <a href="/wiki/Pathogens" class="mw-redirect" title="Pathogens">pathogens</a>, they require mechanisms that enable the determination of whether a cell is infected or not. The exact mechanisms remain the subject of current investigation, but recognition of an "altered self" state is thought to be involved. To control their cytotoxic activity, NK cells possess two types of surface <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptors</a>: activating receptors and inhibitory receptors, including <a href="/wiki/Killer-cell_immunoglobulin-like_receptors" class="mw-redirect" title="Killer-cell immunoglobulin-like receptors">killer-cell immunoglobulin-like receptors</a>. Most of these receptors are not unique to NK cells and can be present in some <a href="/wiki/T_cell" title="T cell">T cell</a> subsets, as well. </p><p>The inhibitory receptors recognize <a href="/wiki/MHC_class_I" title="MHC class I">MHC class I</a> <a href="/wiki/Allele" title="Allele">alleles</a>, which could explain why NK cells preferentially kill cells that possess low levels of MHC class I molecules. This mode of NK cell target interaction is known as "missing-self recognition", a term coined by <a href="/wiki/Klas_K%C3%A4rre" title="Klas Kärre">Klas Kärre</a> and co-workers in the late 90s. MHC class I molecules are the main mechanism by which cells display viral or tumor antigens to cytotoxic T cells. A common evolutionary adaptation to this is seen in both intracellular <a href="/wiki/Microbes" class="mw-redirect" title="Microbes">microbes</a> and tumors: the chronic down-regulation of MHC I molecules, which makes affected cells invisible to T cells, allowing them to evade T cell-mediated immunity. NK cells apparently evolved as an evolutionary response to this adaptation (the loss of the MHC eliminates CD4/CD8 action, so another immune cell evolved to fulfill the function).<sup id="cite_ref-Lodoen_34-0" class="reference"><a href="#cite_note-Lodoen-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tumor_cell_surveillance">Tumor cell surveillance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=11" title="Edit section: Tumor cell surveillance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Natural killer cells often lack antigen-specific cell surface receptors, so are part of innate immunity, <i>i.e.</i> able to react immediately with no prior exposure to the pathogen. In both mice and humans, NKs can be seen to play a role in tumor immunosurveillance by directly inducing the death of tumor cells (NKs act as cytolytic effector lymphocytes), even in the absence of surface adhesion molecules and antigenic peptides. This role of NK cells is critical to immune success particularly because T cells are unable to recognize pathogens in the absence of surface antigens.<sup id="cite_ref-Vivier2011_4-1" class="reference"><a href="#cite_note-Vivier2011-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Tumor cell detection results in activation of NK cells and consequent cytokine production and release. </p><p>If tumor cells do not cause inflammation, they will also be regarded as self and will not induce a T cell response. A number of cytokines are produced by NKs, including tumor necrosis factor α (<a href="/wiki/TNF%CE%B1" class="mw-redirect" title="TNFα">TNFα</a>), <a href="/wiki/IFN%CE%B3" class="mw-redirect" title="IFNγ">IFNγ</a>, and <a href="/wiki/Interleukin" title="Interleukin">interleukin</a> (<a href="/wiki/Interleukin_10" title="Interleukin 10">IL-10</a>). TNFα and IL-10 act as proinflammatory and immunosuppressors, respectively. The activation of NK cells and subsequent production of cytolytic effector cells impacts <a href="/wiki/Macrophages" class="mw-redirect" title="Macrophages">macrophages</a>, <a href="/wiki/Dendritic_cells" class="mw-redirect" title="Dendritic cells">dendritic cells</a>, and <a href="/wiki/Neutrophils" class="mw-redirect" title="Neutrophils">neutrophils</a>, which subsequently enables antigen-specific T and B cell responses. Instead of acting via antigen-specific receptors, lysis of tumor cells by NK cells is mediated by alternative receptors, including <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a>, NKp44, NKp46, NKp30, and DNAM.<sup id="cite_ref-Terunuma2008_28-1" class="reference"><a href="#cite_note-Terunuma2008-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a> is a <a href="/wiki/Disulfide_bond" class="mw-redirect" title="Disulfide bond">disulfide</a>-linked <a href="/wiki/Homodimer" class="mw-redirect" title="Homodimer">homodimer</a> which recognizes a number of ligands, including ULBP and <a href="/wiki/MHC_class_I_polypeptide-related_sequence_A" class="mw-redirect" title="MHC class I polypeptide-related sequence A">MICA</a>, which are typically expressed on tumor cells. The role of dendritic cell—NK cell interface in immunobiology have been studied and defined as critical for the comprehension of the complex immune system.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="removed citation to predatory publisher content (December 2019)">citation needed</span></a></i>]</sup> </p><p>NK cells, along with <a href="/wiki/Macrophage" title="Macrophage">macrophages</a> and several other cell types, express the Fc receptor (FcR) molecule (FC-gamma-RIII = CD16), an activating biochemical <a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">receptor</a> that binds the <a href="/wiki/Fragment_crystallizable_region" title="Fragment crystallizable region">Fc</a> portion of IgG class <a href="/wiki/Antibody" title="Antibody">antibodies</a>. This allows NK cells to target cells against which there has been a <a href="/wiki/Humoral_immunity" title="Humoral immunity">humoral response</a> and to <a href="/wiki/Lysis" title="Lysis">lyse</a> cells through antibody-dependant cytotoxicity (ADCC). This response depends on the affinity of the Fc receptor expressed on NK cells, which can have high, intermediate, and low affinity for the Fc portion of the antibody. This affinity is determined by the amino acid in position 158 of the protein, which can be phenylalanine (F allele) or valine (V allele). Individuals with high-affinity FcRgammRIII (158 V/V allele) respond better to antibody therapy. This has been shown for lymphoma patients who received the antibody Rituxan. Patients who express the 158 V/V allele had a better antitumor response. Only 15–25% of the population expresses the 158 V/V allele. To determine the ADCC contribution of monoclonal antibodies, NK-92 cells (a "pure" NK cell line) has been transfected with the gene for the high-affinity FcR. </p> <div class="mw-heading mw-heading3"><h3 id="Clearance_of_senescent_cells">Clearance of senescent cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=12" title="Edit section: Clearance of senescent cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Natural killer cells (NK cells) and <a href="/wiki/Macrophage" title="Macrophage">macrophages</a> play a major role in clearance of <a href="/wiki/Cellular_senescence" title="Cellular senescence">senescent cells</a>.<sup id="cite_ref-pmid30811627_35-0" class="reference"><a href="#cite_note-pmid30811627-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Natural killer cells directly kill senescent cells, and produce <a href="/wiki/Cytokine" title="Cytokine">cytokines</a> which activate macrophages which remove senescent cells.<sup id="cite_ref-pmid30811627_35-1" class="reference"><a href="#cite_note-pmid30811627-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Natural killer cells can use <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a> receptors to detect senescent cells, and kill those cells using <a href="/wiki/Perforin" class="mw-redirect" title="Perforin">perforin</a> pore-forming <a href="/wiki/Cytolysis" title="Cytolysis">cytolytic</a> protein.<sup id="cite_ref-pmid31088710_36-0" class="reference"><a href="#cite_note-pmid31088710-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">CD8+ cytotoxic T-lymphocytes</a> also use NKG2D receptors to detect senescent cells, and promote killing similar to NK cells.<sup id="cite_ref-pmid31088710_36-1" class="reference"><a href="#cite_note-pmid31088710-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> For example, in patients with Parkinson's disease, levels of Natural killer cells are elevated as they degrade alpha-synuclein aggregates, destroy senescent neurons, and attenuate the neuroinflammation by leukocytes in the central nervous system.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Adaptive_features_of_NK_cells—"memory-like",_"adaptive"_and_memory_NK_cells"><span id="Adaptive_features_of_NK_cells.E2.80.94.22memory-like.22.2C_.22adaptive.22_and_memory_NK_cells"></span>Adaptive features of NK cells—"memory-like", "adaptive" and memory NK cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=13" title="Edit section: Adaptive features of NK cells—"memory-like", "adaptive" and memory NK 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/Adaptive_NK_cells" class="mw-redirect" title="Adaptive NK cells">Adaptive NK cells</a></div> <p>The ability to generate memory cells following a primary infection and the consequent rapid immune activation and response to succeeding infections by the same antigen is fundamental to the role that T and B cells play in the adaptive immune response. For many years, NK cells have been considered to be a part of the innate immune system. However, recently increasing evidence suggests that NK cells can display several features that are usually attributed to adaptive immune cells (e.g. T cell responses) such as dynamic expansion and contraction of subsets, increased longevity and a form of immunological memory, characterized by a more potent response upon secondary challenge with the same antigen.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> In mice, the majority of research was carried out with murine cytomegalovirus (MCMV) and in models of hapten-hypersensitivity reactions. Especially, in the MCMV model, protective memory functions of MCMV-induced NK cells were discovered<sup id="cite_ref-SunBeilke2009_40-0" class="reference"><a href="#cite_note-SunBeilke2009-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> and direct recognition of the MCMV-ligand m157 by the receptor Ly49 was demonstrated to be crucial for the generation of adaptive NK cell responses.<sup id="cite_ref-SunBeilke2009_40-1" class="reference"><a href="#cite_note-SunBeilke2009-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> In humans, most studies have focused on the expansion of an NK cell subset carrying the activating receptor <a href="/wiki/NKG2C" class="mw-redirect" title="NKG2C">NKG2C</a> (<a href="/wiki/KLRC2" title="KLRC2">KLRC2</a>). Such expansions were observed primarily in response to <a href="/wiki/Human_cytomegalovirus" class="mw-redirect" title="Human cytomegalovirus">human cytomegalovirus</a> (HCMV),<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> but also in other infections including <a href="/wiki/Hantavirus" class="mw-redirect" title="Hantavirus">Hantavirus</a>, <a href="/wiki/Chikungunya_virus" class="mw-redirect" title="Chikungunya virus">Chikungunya virus</a>, <a href="/wiki/HIV" title="HIV">HIV</a>, or viral <a href="/wiki/Hepatitis" title="Hepatitis">hepatitis</a>. However, whether these virus infections trigger the expansion of adaptive NKG2C+ NK cells or whether other infections result in re-activation of latent HCMV (as suggested for hepatitis <sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup>), remains a field of study. Notably, recent research suggests that <a href="/wiki/Adaptive_NK_cells" class="mw-redirect" title="Adaptive NK cells">adaptive NK cells</a> can use the activating receptor <a href="/wiki/NKG2C" class="mw-redirect" title="NKG2C">NKG2C</a> (<a href="/wiki/KLRC2" title="KLRC2">KLRC2</a>) to directly bind to <a href="/wiki/Human_cytomegalovirus" class="mw-redirect" title="Human cytomegalovirus">human cytomegalovirus</a>-derived peptide antigens and respond to peptide recognition with activation, expansion, and differentiation,<sup id="cite_ref-HammerRuckert2018_43-0" class="reference"><a href="#cite_note-HammerRuckert2018-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> a mechanism of responding to virus infections that was previously only known for <a href="/wiki/T_cells" class="mw-redirect" title="T cells">T cells</a> of the <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a>. </p> <div class="mw-heading mw-heading3"><h3 id="NK_cell_function_in_pregnancy">NK cell function in pregnancy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=14" title="Edit section: NK cell function in pregnancy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As the majority of pregnancies involve two parents who are not tissue-matched, successful <a href="/wiki/Pregnancy" title="Pregnancy">pregnancy</a> requires the mother's immune system to be <a href="/wiki/Immune_tolerance_in_pregnancy" title="Immune tolerance in pregnancy">suppressed</a>. NK cells are thought to be an important cell type in this process.<sup id="cite_ref-pmid20061017_44-0" class="reference"><a href="#cite_note-pmid20061017-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> These cells are known as "<a href="/wiki/Uterine_Natural_Killer_Cells" class="mw-redirect" title="Uterine Natural Killer Cells">uterine NK cells</a>" (uNK cells) and they differ from peripheral NK cells. They are in the <a href="/wiki/CD56" class="mw-redirect" title="CD56">CD56<sup>bright</sup></a> NK cell subset, potent at cytokine secretion, but with low cytotoxic ability and relatively similar to peripheral CD56<sup>bright</sup> NK cells, with a slightly different receptor profile.<sup id="cite_ref-pmid20061017_44-1" class="reference"><a href="#cite_note-pmid20061017-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> These uNK cells are the most abundant <a href="/wiki/Leukocyte" class="mw-redirect" title="Leukocyte">leukocytes</a> present <i>in utero</i> in early pregnancy, representing about 70% of leukocytes here, but from where they originate remains controversial.<sup id="cite_ref-pmid19876837_45-0" class="reference"><a href="#cite_note-pmid19876837-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>These NK cells have the ability to elicit cell cytotoxicity <i>in vitro</i>, but at a lower level than peripheral NK cells, despite containing <a href="/wiki/Perforin" class="mw-redirect" title="Perforin">perforin</a>.<sup id="cite_ref-PMCid1266146_46-0" class="reference"><a href="#cite_note-PMCid1266146-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Lack of cytotoxicity <i>in vivo</i> may be due to the presence of ligands for their inhibitory receptors. <a href="/wiki/Trophoblast" title="Trophoblast">Trophoblast</a> cells downregulate <a href="/wiki/HLA-A" title="HLA-A">HLA-A</a> and <a href="/wiki/HLA-B" title="HLA-B">HLA-B</a> to defend against <a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">cytotoxic T cell</a>-mediated death. This would normally trigger NK cells by missing self recognition; however, these cells survive. The selective retention of <a href="/wiki/HLA-E" title="HLA-E">HLA-E</a> (which is a ligand for NK cell inhibitory receptor <a href="/wiki/NKG2" title="NKG2">NKG2A</a>) and <a href="/wiki/HLA-G" title="HLA-G">HLA-G</a> (which is a ligand for NK cell inhibitory receptor <a href="/wiki/KIR2DL4" title="KIR2DL4">KIR2DL4</a>) by the trophoblast is thought to defend it against NK cell-mediated death.<sup id="cite_ref-pmid20061017_44-2" class="reference"><a href="#cite_note-pmid20061017-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p><p>Uterine NK cells have shown no significant difference in women with <a href="/wiki/Recurrent_miscarriage" title="Recurrent miscarriage">recurrent miscarriage</a> compared with controls. However, higher peripheral NK cell percentages occur in women with recurrent miscarriages than in control groups.<sup id="cite_ref-SeshadriSunkara2013_47-0" class="reference"><a href="#cite_note-SeshadriSunkara2013-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p>NK cells secrete a high level of cytokines which help mediate their function. NK cells interact with <a href="/wiki/HLA-C" title="HLA-C">HLA-C</a> to produce cytokines necessary for trophoblastic proliferation. Some important cytokines they secrete include <a href="/wiki/TNF-%CE%B1" class="mw-redirect" title="TNF-α">TNF-α</a>, <a href="/wiki/Interleukin_10" title="Interleukin 10">IL-10</a>, <a href="/wiki/IFN-%CE%B3" class="mw-redirect" title="IFN-γ">IFN-γ</a>, <a href="/wiki/GM-CSF" class="mw-redirect" title="GM-CSF">GM-CSF</a> and <a href="/wiki/TGF-%CE%B2" class="mw-redirect" title="TGF-β">TGF-β</a>, among others.<sup id="cite_ref-pmid20061017_44-3" class="reference"><a href="#cite_note-pmid20061017-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> For example, IFN-γ dilates and thins the walls of maternal <a href="/wiki/Spiral_arteries" class="mw-redirect" title="Spiral arteries">spiral arteries</a> to enhance blood flow to the implantation site.<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> </p> <div class="mw-heading mw-heading3"><h3 id="NK_cell_evasion_by_tumor_cells">NK cell evasion by tumor cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=15" title="Edit section: NK cell evasion by tumor cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By shedding decoy <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a> soluble ligands, tumor cells may avoid immune responses. These soluble NKG2D ligands bind to NK cell NKG2D receptors, activating a false NK response and consequently creating competition for the receptor site.<sup id="cite_ref-Vivier2011_4-2" class="reference"><a href="#cite_note-Vivier2011-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This method of evasion occurs in <a href="/wiki/Prostate_cancer" title="Prostate cancer">prostate cancer</a>. In addition, prostate cancer tumors can evade CD8 cell recognition due to their ability to downregulate expression of MHC class 1 molecules. This example of immune evasion actually highlights NK cells' importance in tumor surveillance and response, as CD8 cells can consequently only act on tumor cells in response to NK-initiated cytokine production (adaptive immune response).<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> </p> <div class="mw-heading mw-heading3"><h3 id="Excessive_NK_cells">Excessive NK cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=16" title="Edit section: Excessive NK cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Experimental treatments with NK cells have resulted in excessive cytokine production, and even <a href="/wiki/Septic_shock" title="Septic shock">septic shock</a>. Depletion of the inflammatory cytokine <a href="/wiki/Interferon_gamma" title="Interferon gamma">interferon gamma</a> reversed the effect.<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. (November 2020)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=17" title="Edit section: Applications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Anticancer_therapy">Anticancer therapy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=18" title="Edit section: Anticancer therapy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Tumor-infiltrating NK cells have been reported to play a critical role in promoting drug-induced cell death in human triple-negative breast cancer.<sup id="cite_ref-pmid33077554_50-0" class="reference"><a href="#cite_note-pmid33077554-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Since NK cells recognize target cells when they express nonself HLA antigens (but not self), autologous (patients' own) NK cell infusions have not shown any antitumor effects. Instead, investigators are working on using allogeneic cells from peripheral blood, which requires that all T cells be removed before infusion into the patients to remove the risk of <a href="/wiki/Graft-versus-host_disease" title="Graft-versus-host disease">graft versus host disease</a>, which can be fatal. This can be achieved using an immunomagnetic column (CliniMACS). In addition, because of the limited number of NK cells in blood (only 10% of lymphocytes are NK cells), their number needs to be expanded in culture. This can take a few weeks and the yield is donor-dependent. </p> <div class="mw-heading mw-heading3"><h3 id="CAR-NK_cells">CAR-NK cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=19" title="Edit section: CAR-NK cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Chimeric_antigen_receptor" class="mw-redirect" title="Chimeric antigen receptor">Chimeric antigen receptors</a> (CARs) are genetically modified receptors targeting cell surface <a href="/wiki/Antigen" title="Antigen">antigens</a> that provide a valuable approach to enhance effector cell efficacy. CARs induce high-affinity binding of effector cells carrying these <a href="/w/index.php?title=Receptors_(biochemistry)&action=edit&redlink=1" class="new" title="Receptors (biochemistry) (page does not exist)">receptors</a> to cells expressing the target antigen, thereby lowering the threshold for cellular activation and inducing effector functions.<sup id="cite_ref-:5_51-0" class="reference"><a href="#cite_note-:5-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/CAR_T_cell" title="CAR T cell">CAR T cells</a> are now a fairly well-known <a href="/wiki/Cell_therapy" title="Cell therapy">cell therapy</a>. However, wider use is limited by several fundamental problems: The high cost of CAR T cell therapy, which is due to the need to generate specific CAR T cells for each patient; the necessity to use only autologous T cells, due to the high risk of <a href="/wiki/GvHD" class="mw-redirect" title="GvHD">GvHD</a> if allogeneic T cells are used; the inability to reinfuse CAR T cells if the patient relapses or low CAR T cell survival is observed; CAR T therapy also has a high toxicity, mainly due to <a href="/wiki/Interferon_type_I" title="Interferon type I">IFN-γ</a> production and subsequent induction of CRS (<a href="/wiki/Cytokine_release_syndrome" title="Cytokine release syndrome">cytokine release syndrome</a>) and/or <a href="/wiki/Neurotoxicity" title="Neurotoxicity">neurotoxicity</a>.<sup id="cite_ref-:6_52-0" class="reference"><a href="#cite_note-:6-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>The use of CAR NK cells is not limited by the need to generate patient-specific cells, and at the same time, GvHD is not caused by NK cells, thus obviating the need for autologous cells.<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> Toxic effects of CAR T therapy, such as CSR, have not been observed with the use of CAR NK cells. Thus, NK cells are considered an interesting "off-the-shelf" product option. Compared to CAR T cells, CAR NK cells retain unchanged expression of NK cell activating receptors. Thus, NK cells recognize and kill tumor cells even if, due to a tumor-escape strategy on tumor cells, ligand expression for the CAR receptor is downregulated.<sup id="cite_ref-:6_52-1" class="reference"><a href="#cite_note-:6-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>NK cells derived from umbilical cord blood have been used to generate CAR.CD19 NK cells. These cells are capable of self-producing the cytokine <a href="/wiki/Interleukin_15" title="Interleukin 15">IL-15</a>, thereby enhancing autocrine/paracrine expression and persistence <a href="/wiki/In_vivo" title="In vivo">in vivo</a>. Administration of these modified NK cells is not associated with the development of CSR, neurotoxicity, or GvHD.<sup id="cite_ref-:5_51-1" class="reference"><a href="#cite_note-:5-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>The FT596 product is the first "Off-the-Shelf", universal, and allogenic CAR NK cellular product derived from <a href="/wiki/Induced_pluripotent_stem_cell" title="Induced pluripotent stem cell">iPSCs</a> to be authorized for use in clinical studies in the USA.<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> It consists of an anti-CD19 CAR optimized for NK cells with a transmembrane domain for the <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a> activation receptor, a 2B4 costimulatory domain and a CD3ζ signaling domain. Two additional key components were added: 1) a high-affinity, non-cleavable Fc receptor CD16 (hnCD16) that enables tumor targeting and enhanced antibody-dependent cell cytotoxicity without negative regulation, combined with 2) a therapeutic monoclonal antibody targeting tumor cells and an IL-15/IL-15 receptor fusion protein (IL-15RF) promoting cytokine-independent persistence.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="NK-92_cells">NK-92 cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=20" title="Edit section: NK-92 cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A more efficient way to obtain high numbers of NK cells is to expand <a href="/wiki/NK-92" title="NK-92">NK-92 cells</a>, an NK cell line with all the characteristics of highly active blood Natural Killer (NK) cells but with much broader and higher cytotoxicity. NK-92 cells grow continuously in culture and can be expanded to clinical-grade numbers in bags or bioreactors.<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> Clinical studies have shown NK-92 cells to be safe and to exhibit anti-tumor activity in patients with lung or pancreatic cancer, melanoma, and lymphoma.<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><sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> When NK-92 cells originate from a patient with lymphoma, they must be irradiated prior to infusion.<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><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> Efforts, however, are being made to engineer the cells to eliminate the need for irradiation. The irradiated cells maintain full cytotoxicity. NK-92 are allogeneic (from a donor different from the recipient), but in clinical studies have not been shown to elicit significant host reaction.<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><sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>Unmodified NK-92 cells lack CD-16, making them unable to perform antibody-dependent cellular cytotoxicity (ADCC); however, the cells have been engineered to express a high affinity Fc-receptor (CD16A, 158V) genetically linked to IL-2 that is bound to the endoplasmic reticulum (ER).<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup><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> These high affinity NK-92 cells can perform ADCC and have greatly expanded therapeutic utility.<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><sup id="cite_ref-ReferenceA_66-0" class="reference"><a href="#cite_note-ReferenceA-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><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> </p><p>NK-92 cells have also been engineered to expressed chimeric antigen receptors (CARs), in an approach similar to that used for T cells. An example of this is an NK-92 derived cell engineered with both a CD16 and an anti-PD-L1 CAR; currently in clinical development for oncology indications.<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> A clinical grade NK-92 variant that expresses a CAR for HER2 (ErbB2) has been generated<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> and is in a clinical study in patients with HER2 positive <a href="/wiki/Glioblastoma" title="Glioblastoma">glioblastoma</a>.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Several other clinical grade clones have been generated expressing the CARs for PD-L1, CD19, HER-2, and EGFR.<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-ReferenceA_66-1" class="reference"><a href="#cite_note-ReferenceA-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> PD-L1 targeted high affinity NK cells have been given to a number of patients with solid tumors in a phase I/II study, which is underway.<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> </p> <div class="mw-heading mw-heading3"><h3 id="NKG2D-Fc_fusion_protein">NKG2D-Fc fusion protein</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=21" title="Edit section: NKG2D-Fc fusion protein"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In a study at Boston Children's Hospital, in coordination with <a href="/wiki/Dana%E2%80%93Farber_Cancer_Institute" title="Dana–Farber Cancer Institute">Dana–Farber Cancer Institute</a>, in which immunocompromised mice had contracted <a href="/wiki/Lymphomas" class="mw-redirect" title="Lymphomas">lymphomas</a> from <a href="/wiki/Epstein-Barr_virus" class="mw-redirect" title="Epstein-Barr virus">EBV</a> infection, an NK-activating receptor called <a href="/wiki/NKG2D" title="NKG2D">NKG2D</a> was fused with a stimulatory <a href="/wiki/Fc_region" class="mw-redirect" title="Fc region">Fc</a> portion of the EBV antibody. The NKG2D-Fc fusion proved capable of reducing tumor growth and prolonging survival of the recipients. In a transplantation model of LMP1-fueled lymphomas, the NKG2D-Fc fusion proved capable of reducing tumor growth and prolonging survival of the recipients. </p><p>In Hodgkin lymphoma, in which the malignant Hodgkin Reed-Sternberg cells are typically HLA class I deficient, immune evasion is in part mediated by skewing towards an exhausted PD-1hi NK cell phenotype, and re-activation of these NK cells appears to be one mechanism of action induced by checkpoint-blockade.<sup id="cite_ref-pmid29449276_76-0" class="reference"><a href="#cite_note-pmid29449276-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="TLR_ligands">TLR ligands</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=22" title="Edit section: TLR ligands"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Signaling through <a href="/wiki/Toll-like_receptor" title="Toll-like receptor">TLR</a> can effectively activate NK cell effector functions <i>in vitro</i> and <i>in vivo</i>. TLR ligands are then potentially able to enhance NK cell effector functions during NK cell anti-tumor <a href="/wiki/Immunotherapy" title="Immunotherapy">immunotherapy</a>.<sup id="cite_ref-:4_30-1" class="reference"><a href="#cite_note-:4-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Trastuzumab" title="Trastuzumab">Trastuzumab</a> is a <a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">monoclonal</a> anti-HER2 <a href="/wiki/Antibody" title="Antibody">antibody</a> that is used as a treatment of the <a href="/wiki/HER2" title="HER2">HER2+</a> <a href="/wiki/Breast_cancer" title="Breast cancer">breast cancer</a>.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> NK cells are an important part of the therapeutical effect of trastzumab as NK cells recognize the antibody coated cancer cells which induces <a href="/wiki/Antibody-dependent_cellular_cytotoxicity" title="Antibody-dependent cellular cytotoxicity">ADCC</a> (antibody-dependent cellular cytotoxicity) reaction. TLR ligand is used in addition to trastuzumab as a means to enhance its effect. The <a href="/wiki/Polysaccharide-K" title="Polysaccharide-K">polysaccharide krestin</a>, which is extracted from <i><a href="/wiki/Trametes_versicolor" title="Trametes versicolor">Trametes versicolor</a></i>, is a potent ligand of <a href="/wiki/Toll-like_receptor_2" title="Toll-like receptor 2">TLR-2</a> and so activates NK cells, induces the production of <a href="/wiki/IFNG" class="mw-redirect" title="IFNG">IFNg</a> and enhances the <a href="/wiki/Antibody-dependent_cellular_cytotoxicity" title="Antibody-dependent cellular cytotoxicity">ADCC</a> caused by recognition of trastuzumab-coated cells.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p><p>Stimulation of <a href="/wiki/Toll-like_receptor_7" title="Toll-like receptor 7">TLR-7</a> induces the expression of <a href="/wiki/Interferon_type_I" title="Interferon type I">IFN type I</a> and other pro-inflammatory cytokines like <a href="/wiki/IL-1B" class="mw-redirect" title="IL-1B">IL-1b</a>, <a href="/wiki/Interleukin_6" title="Interleukin 6">IL-6</a> and <a href="/wiki/Interleukin_12" title="Interleukin 12">IL-12</a>. Mice suffering with <a href="/w/index.php?title=NK_cell-sensitive_lymphoma_RMA-S&action=edit&redlink=1" class="new" title="NK cell-sensitive lymphoma RMA-S (page does not exist)">NK cell-sensitive lymphoma RMA-S</a> were treated with SC1 molecule. SC1 is novel small-molecule TLR-7 agonist and its repeated administration reportedly activated NK cells in TLR-7- and IFN type I- dependent manner thus reversing the NK cell <a href="/wiki/Anergy" class="mw-redirect" title="Anergy">anergy</a> which ultimately lead to <a href="/wiki/Lysis" title="Lysis">lysis</a> of the tumor.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/VTX-2337" class="mw-redirect" title="VTX-2337">VTX-2337</a> is a selective <a href="/wiki/Toll-like_receptor_8" title="Toll-like receptor 8">TLR-8</a> agonist and together with <a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">monoclonal antibody</a> <a href="/wiki/Cetuximab" title="Cetuximab">cetuximab</a> it was used as a potential therapy for the treatment of recurrent or metastatic <a href="/wiki/SCCHN" class="mw-redirect" title="SCCHN">SCCHN</a>. Results have shown that the NK cells had become more reactive to the treatment with <a href="/wiki/Cetuximab" title="Cetuximab">cetuximab</a> antibody upon pretreatment with VTX-2337. This indicates that the stimulation of TLR-8 and subsequent activation of <a href="/wiki/Inflammasome" title="Inflammasome">inflammasome</a> enhances the <a href="/wiki/Fc_receptor" title="Fc receptor">CD-16</a> mediated <a href="/wiki/Antibody-dependent_cellular_cytotoxicity" title="Antibody-dependent cellular cytotoxicity">ADCC</a> reaction in patients treated with <a href="/wiki/Cetuximab" title="Cetuximab">cetuximab</a> antibody.<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>NK cells play a role in controlling <a href="/wiki/HIV-1" class="mw-redirect" title="HIV-1">HIV-1</a> infection. TLR are potent enhancers of innate antiviral immunity and potentially can reverse HIV-1 latency. Incubation of <a href="/wiki/Peripheral_blood_mononuclear_cell" title="Peripheral blood mononuclear cell">peripheral blood mononuclear cells</a> with novel potent <a href="/wiki/Toll-like_receptor_9" title="Toll-like receptor 9">TLR-9</a> ligand <a href="/w/index.php?title=MGN1703&action=edit&redlink=1" class="new" title="MGN1703 (page does not exist)">MGN1703</a> have resulted in enhancement of NK cell effector functions, thus significantly inhibiting the spread of HIV-1 in culture of <a href="/wiki/Autologous" class="mw-redirect" title="Autologous">autologous</a> <a href="/wiki/Cd4%2B_t_cells" class="mw-redirect" title="Cd4+ t cells">CD4+ T-cells</a>. The stimulation of TLR-9 in NK cells induced a strong antiviral innate immune response, an increase in HIV-1 <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">transcription</a> (indicating the reverse in latency of the virus) and it also boosted the NK cell-mediated suppression of HIV-1 infections in autologous CD4+ T cells.<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> <div class="mw-heading mw-heading2"><h2 id="New_findings">New findings</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=23" title="Edit section: New findings"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Innate_resistance_to_HIV">Innate resistance to HIV</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=24" title="Edit section: Innate resistance to HIV"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Recent research suggests specific KIR-MHC class I gene interactions might control innate genetic resistance to certain viral infections, including <a href="/wiki/HIV" title="HIV">HIV</a> and its consequent development of <a href="/wiki/AIDS" class="mw-redirect" title="AIDS">AIDS</a>.<sup id="cite_ref-Lannello2008_8-3" class="reference"><a href="#cite_note-Lannello2008-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Certain HLA allotypes have been found to determine the progression of HIV to AIDS; an example is the <a href="/wiki/HLA-B57" title="HLA-B57">HLA-B57</a> and <a href="/wiki/HLA-B27" title="HLA-B27">HLA-B27</a> alleles, which have been found to delay progression from HIV to AIDS. This is evident because patients expressing these HLA alleles are observed to have lower viral loads and a more gradual decline in <a href="/wiki/T_helper_cell" title="T helper cell">CD4<sup>+</sup> T</a> cells numbers. Despite considerable research and data collected measuring the genetic correlation of HLA alleles and KIR allotypes, a firm conclusion has not yet been drawn as to what combination provides decreased HIV and AIDS susceptibility. </p><p>NK cells can impose immune pressure on HIV, which had previously been described only for T cells and antibodies.<sup id="cite_ref-Alter_et_al._82-0" class="reference"><a href="#cite_note-Alter_et_al.-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> HIV mutates to avoid NK cell detection.<sup id="cite_ref-Alter_et_al._82-1" class="reference"><a href="#cite_note-Alter_et_al.-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tissue-resident_NK_cells">Tissue-resident NK cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=25" title="Edit section: Tissue-resident NK cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most of our current knowledge is derived from investigations of mouse splenic and human peripheral blood NK cells. However, in recent years tissue-resident NK cell populations have been described.<sup id="cite_ref-:0_83-0" class="reference"><a href="#cite_note-:0-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_84-0" class="reference"><a href="#cite_note-:1-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> These tissue-resident NK cells share transcriptional similarity to tissue-resident memory T cells described previously. However, tissue-resident NK cells are not necessarily of the memory phenotype, and in fact, the majority of the tissue-resident NK cells are functionally immature.<sup id="cite_ref-Dogra2020_85-0" class="reference"><a href="#cite_note-Dogra2020-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> These specialized NK-cell subsets can play a role in organ homeostasis. For example, NK cells are enriched in the human liver with a specific phenotype and take part in the control of liver fibrosis.<sup id="cite_ref-:2_86-0" class="reference"><a href="#cite_note-:2-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:3_87-0" class="reference"><a href="#cite_note-:3-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> Tissue-resident NK cells have also been identified in sites like bone marrow, spleen and more recently, in lung, intestines and lymph nodes. In these sites, tissue-resident NK cells may act as reservoir for maintaining immature NK cells in humans throughout life.<sup id="cite_ref-Dogra2020_85-1" class="reference"><a href="#cite_note-Dogra2020-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Adaptive_NK_cells_against_leukemia_targets">Adaptive NK cells against leukemia targets</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=26" title="Edit section: Adaptive NK cells against leukemia targets"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Natural killer cells are being investigated as an emerging treatment for patients with acute myeloid leukemia (AML), and cytokine-induced memory-like NK cells have shown promise with their enhanced antileukemia functionality.<sup id="cite_ref-Romee2016_88-0" class="reference"><a href="#cite_note-Romee2016-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> It has been shown that this kind of NK cell has enhanced interferon-γ production and cytotoxicity against leukemia cell lines and primary AML blasts in patients.<sup id="cite_ref-Romee2016_88-1" class="reference"><a href="#cite_note-Romee2016-88"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> During a phase 1 clinical trial, five out of nine patients exhibited clinical responses to the treatment, and four patients experienced a complete remission, which suggests that these NK cells have major potential as a successful translational immunotherapy approach for patients with AML in the future.<sup id="cite_ref-Romee2016_88-2" class="reference"><a href="#cite_note-Romee2016-88"><span class="cite-bracket">[</span>88<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=Natural_killer_cell&action=edit&section=27" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Active_hexose_correlated_compound" class="mw-redirect" title="Active hexose correlated compound">Active hexose correlated compound</a></li> <li><a href="/wiki/Granzyme" title="Granzyme">Granzymes</a></li> <li><a href="/wiki/Hematopoiesis" class="mw-redirect" title="Hematopoiesis">Hematopoiesis</a></li> <li><a href="/wiki/Immune_system" title="Immune system">Immune system</a></li> <li><a href="/wiki/Interleukin" title="Interleukin">Interleukin</a></li> <li><a href="/wiki/Lymphatic_system" title="Lymphatic system">Lymphatic system</a></li> <li><a href="/wiki/List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">List of distinct cell types in the adult human body</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=28" 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-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output 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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="CITEREFOshimi2017" class="citation journal cs1">Oshimi, Kazuo (2017). <a rel="nofollow" class="external text" href="https://www.jstage.jst.go.jp/article/internalmedicine/56/14/56_56.8881/_article">"Clinical Features, Pathogenesis, and Treatment of 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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=Natural_killer_cell&action=edit&section=29" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPerera_Molligoda_Arachchige2021" class="citation journal cs1">Perera Molligoda Arachchige AS (April 2021). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8054151">"Human NK cells: From development to effector functions"</a>. <i>Innate Immunity</i>. <b>27</b> (3): 212–229. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1177%2F17534259211001512">10.1177/17534259211001512</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8054151">8054151</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/33761782">33761782</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Innate+Immunity&rft.atitle=Human+NK+cells%3A+From+development+to+effector+functions&rft.volume=27&rft.issue=3&rft.pages=212-229&rft.date=2021-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8054151%23id-name%3DPMC&rft_id=info%3Apmid%2F33761782&rft_id=info%3Adoi%2F10.1177%2F17534259211001512&rft.aulast=Perera+Molligoda+Arachchige&rft.aufirst=AS&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8054151&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANatural+killer+cell" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAbbasLichtman2003" class="citation book cs1">Abbas AK, Lichtman A (2003). <i>Cellular and Molecular Immunology</i>. Saunders.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Cellular+and+Molecular+Immunology&rft.pub=Saunders&rft.date=2003&rft.aulast=Abbas&rft.aufirst=AK&rft.au=Lichtman%2C+A&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANatural+killer+cell" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSompayrac2003" class="citation book cs1">Sompayrac L (2003). <i>How the Immune System Works</i> (2nd ed.). Blackwell Publishing.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=How+the+Immune+System+Works&rft.edition=2nd&rft.pub=Blackwell+Publishing&rft.date=2003&rft.aulast=Sompayrac&rft.aufirst=L&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANatural+killer+cell" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJaneway_JrTraversWalportShlomchik2001" class="citation book cs1">Janeway Jr CA, Travers P, Walport M, Shlomchik MJ (2001). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK10757/"><i>Immunobiology: The Immune System In Health And Disease</i></a> (5th ed.). Garland Science. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-8153-3642-X" title="Special:BookSources/0-8153-3642-X"><bdi>0-8153-3642-X</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Immunobiology%3A+The+Immune+System+In+Health+And+Disease&rft.edition=5th&rft.pub=Garland+Science&rft.date=2001&rft.isbn=0-8153-3642-X&rft.aulast=Janeway+Jr&rft.aufirst=CA&rft.au=Travers%2C+P&rft.au=Walport%2C+M&rft.au=Shlomchik%2C+MJ&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2FNBK10757%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANatural+killer+cell" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKindtGoldsbyOsborne" class="citation book cs1">Kindt TJ, Goldsby RA, Osborne BA. <i>Kuby Immunology</i> (6th ed.). New York: W.H. Freeman and Company.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Kuby+Immunology&rft.place=New+York&rft.edition=6th&rft.pub=W.H.+Freeman+and+Company&rft.aulast=Kindt&rft.aufirst=TJ&rft.au=Goldsby%2C+RA&rft.au=Osborne%2C+BA&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANatural+killer+cell" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Natural_killer_cell&action=edit&section=30" 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 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href="/wiki/Plasma_cell" title="Plasma cell">Plasma</a></li> <li><a href="/wiki/Memory_B_cell" title="Memory B cell">Memory</a></li> <li><a href="/wiki/Follicular_B_cell" title="Follicular B cell">Follicular</a></li> <li><a href="/wiki/Marginal_zone_B-cell" class="mw-redirect" title="Marginal zone B-cell">Marginal zone</a></li> <li><a href="/wiki/Naive_B_cell" title="Naive B cell">Naïve</a></li> <li><a href="/wiki/B_cell#Development" title="B cell">Pre-B</a></li> <li><a href="/wiki/Regulatory_B_cell" title="Regulatory B cell">B<sub>reg</sub> cell</a> <ul><li><a href="/wiki/B10_cell" title="B10 cell">B10 cell</a></li></ul></li> <li><a href="/wiki/Transitional_B_cell" title="Transitional B cell">Transitional B cell</a></li> <li><a href="/w/index.php?title=Lymphoplasmacytoid_cell&action=edit&redlink=1" class="new" title="Lymphoplasmacytoid cell (page does not exist)">Lymphoplasmacytoid cell</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/T_cell" title="T cell">T cells</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Thymocyte" title="Thymocyte">Thymocyte</a></li> <li>αβ (<a href="/wiki/Cytotoxic_T_cell" title="Cytotoxic T cell">Cytotoxic CD8<sup>+</sup></a></li> <li><a href="/wiki/T_helper_cell" title="T helper cell">Helper CD4<sup>+</sup></a> / <a href="/wiki/Follicular_B_helper_T_cells" title="Follicular B helper T cells">T<sub>FH</sub></a> / <a href="/wiki/T_helper_3_cell" title="T helper 3 cell">T<sub>h</sub>3</a> / <a href="/wiki/T_helper_17_cell" title="T helper 17 cell">T<sub>h</sub>17</a> / <a href="/wiki/Regulatory_T_cell" title="Regulatory T cell">Regulatory</a>)</li> <li><a href="/wiki/Naive_T_cell" title="Naive T cell">Naïve</a></li> <li><a href="/wiki/Memory_T_cell" title="Memory T cell">Memory T cell</a> <ul><li><a href="/wiki/Central_memory_T_cell" class="mw-redirect" title="Central memory T cell">T<sub>CM</sub></a></li> <li><a href="/wiki/Effector_memory_T_cell" class="mw-redirect" title="Effector memory T cell">T<sub>EM</sub></a></li> <li><a href="/wiki/Tissue-resident_memory_T_cell" title="Tissue-resident memory T cell">T<sub>RM</sub></a></li> <li><a href="/wiki/Virtual_memory_T_cell" title="Virtual memory T cell">T<sub>VM</sub></a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Innate-like_T_cell" class="mw-redirect" title="Innate-like T cell">Innate-like T cells</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Natural_killer_T_cell" title="Natural killer T cell">Natural killer T cell</a></li> <li><a href="/wiki/Gamma_delta_T_cell" title="Gamma delta T cell">γδ</a></li> <li><a href="/wiki/Mucosal_associated_invariant_T_cell" class="mw-redirect" title="Mucosal associated invariant T cell">Mucosal associated invariant T cell</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Innate_lymphoid_cell" title="Innate lymphoid cell">Innate lymphoid cells</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="NK_cells" scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">NK cells</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cytokine-induced_killer_cell" title="Cytokine-induced killer cell">Cytokine-induced killer cell</a></li> <li><a href="/wiki/Lymphokine-activated_killer_cell" title="Lymphokine-activated killer cell">Lymphokine-activated killer cell</a></li> <li><a href="/wiki/Null_cell" title="Null cell">Null cell</a></li> <li><a href="/wiki/Adaptive_NK_cells" class="mw-redirect" title="Adaptive NK cells">Adaptive NK cell</a></li> <li><a href="/wiki/Uterine_natural_killer_cells" title="Uterine natural killer cells">Uterine natural killer cells</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Type_1_innate_lymphoid_cell" class="mw-redirect" title="Type 1 innate lymphoid cell">Type 1 innate lymphoid cells</a></li> <li><a href="/wiki/Type_2_innate_lymphoid_cell" class="mw-redirect" title="Type 2 innate lymphoid cell">Type 2 innate lymphoid cells</a> <ul><li><a href="/wiki/Nuocyte" title="Nuocyte">Nuocytes</a></li></ul></li> <li><a href="/wiki/Type_3_innate_lymphoid_cells" title="Type 3 innate lymphoid cells">Type 3 innate lymphoid cells</a></li> <li><a href="/wiki/LTi_cells" class="mw-redirect" title="LTi cells">LTi cells</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lymphopoiesis" title="Lymphopoiesis">Lymphopoiesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hematopoietic_stem_cell" title="Hematopoietic stem cell">Hematopoietic stem cell</a></li> <li><a href="/wiki/Lymphoblast" title="Lymphoblast">Lymphoblast</a></li> <li><a href="/wiki/Prolymphocyte" title="Prolymphocyte">Prolymphocyte</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Lymphocytic_adaptive_immune_system_and_complement" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Lymphocytic_immune_system" title="Template:Lymphocytic immune system"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Lymphocytic_immune_system" title="Template talk:Lymphocytic immune system"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Lymphocytic_immune_system" title="Special:EditPage/Template:Lymphocytic immune system"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Lymphocytic_adaptive_immune_system_and_complement" style="font-size:114%;margin:0 4em"><a href="/wiki/Lymphocyte" title="Lymphocyte">Lymphocytic</a> <a href="/wiki/Adaptive_immune_system" title="Adaptive immune system">adaptive immune system</a> and <a href="/wiki/Complement_system" title="Complement system">complement</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lymphoid</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Antigens</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antigen" title="Antigen">Antigen</a> <ul><li><a href="/wiki/Superantigen" title="Superantigen">Superantigen</a></li> <li><a href="/wiki/Allergen" title="Allergen">Allergen</a></li> <li><a href="/wiki/Antigenic_variation" title="Antigenic variation">Antigenic variation</a></li></ul></li> <li><a href="/wiki/Hapten" title="Hapten">Hapten</a></li></ul> <ul><li><a href="/wiki/Epitope" title="Epitope">Epitope</a> <ul><li><a href="/wiki/Linear_epitope" title="Linear epitope">Linear</a></li> <li><a href="/wiki/Conformational_epitope" title="Conformational epitope">Conformational</a></li></ul></li> <li><a href="/wiki/Mimotope" title="Mimotope">Mimotope</a></li></ul> <ul><li><a href="/wiki/Antigen_presentation" title="Antigen presentation">Antigen presentation</a>/<a href="/wiki/Antigen-presenting_cell" title="Antigen-presenting cell">professional APCs</a>: <a href="/wiki/Dendritic_cell" title="Dendritic cell">Dendritic cell</a></li> <li><a href="/wiki/Macrophage" title="Macrophage">Macrophage</a></li> <li><a href="/wiki/B_cell" title="B cell">B cell</a></li> <li><a href="/wiki/Immunogen" title="Immunogen">Immunogen</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Antibodies</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibody" title="Antibody">Antibody</a> <ul><li><a href="/wiki/Monoclonal_antibody" title="Monoclonal antibody">Monoclonal antibodies</a></li> <li><a href="/wiki/Polyclonal_antibodies" title="Polyclonal antibodies">Polyclonal antibodies</a></li> <li><a href="/wiki/Autoantibody" title="Autoantibody">Autoantibody</a></li> <li><a href="/wiki/Microantibody" title="Microantibody">Microantibody</a></li></ul></li> <li><a href="/wiki/Polyclonal_B_cell_response" title="Polyclonal B cell response">Polyclonal B cell response</a></li> <li><a href="/wiki/Allotype_(immunology)" title="Allotype (immunology)">Allotype</a></li> <li><a href="/wiki/Isotype_(immunology)" title="Isotype (immunology)">Isotype</a></li> <li><a href="/wiki/Idiotype" title="Idiotype">Idiotype</a></li></ul> <ul><li><a href="/wiki/Immune_complex" title="Immune complex">Immune complex</a></li> <li><a href="/wiki/Paratope" title="Paratope">Paratope</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Immunity vs.<br /> tolerance</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Action: <a href="/wiki/Immunity_(medical)" class="mw-redirect" title="Immunity (medical)">Immunity</a></li> <li><a href="/wiki/Autoimmunity" title="Autoimmunity">Autoimmunity</a></li> <li><a href="/wiki/Alloimmunity" title="Alloimmunity">Alloimmunity</a></li> <li><a href="/wiki/Allergy" title="Allergy">Allergy</a></li> <li><a href="/wiki/Hypersensitivity" title="Hypersensitivity">Hypersensitivity</a></li> <li><a href="/wiki/Inflammation" title="Inflammation">Inflammation</a></li> <li><a href="/wiki/Cross-reactivity" title="Cross-reactivity">Cross-reactivity</a></li> <li><a href="/wiki/Co-stimulation" title="Co-stimulation">Co-stimulation</a></li></ul> <ul><li>Inaction: <a href="/wiki/Immune_tolerance" title="Immune tolerance">Tolerance</a> <ul><li><a href="/wiki/Central_tolerance" title="Central tolerance">Central</a></li> <li><a href="/wiki/Peripheral_tolerance" title="Peripheral tolerance">Peripheral</a></li> <li><a href="/wiki/Clonal_anergy" title="Clonal anergy">Clonal anergy</a></li> <li><a href="/wiki/Clonal_deletion" title="Clonal deletion">Clonal deletion</a></li> <li><a href="/wiki/Immune_tolerance_in_pregnancy" title="Immune tolerance in pregnancy">Tolerance in pregnancy</a></li></ul></li> <li><a href="/wiki/Immunodeficiency" title="Immunodeficiency">Immunodeficiency</a></li> <li><a href="/wiki/Immune_privilege" title="Immune privilege">Immune privilege</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affinity_maturation" title="Affinity maturation">Affinity maturation</a> <ul><li><a href="/wiki/Somatic_hypermutation" title="Somatic hypermutation">Somatic hypermutation</a></li> <li><a href="/wiki/Clonal_selection" title="Clonal selection">Clonal selection</a></li></ul></li> <li><a href="/wiki/V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a></li> <li><a href="/wiki/Junctional_diversity" title="Junctional diversity">Junctional diversity</a></li> <li><a href="/wiki/Immunoglobulin_class_switching" title="Immunoglobulin class switching">Immunoglobulin class switching</a></li> <li><a href="/wiki/Major_histocompatibility_complex" title="Major histocompatibility complex">MHC</a>/<a href="/wiki/Human_leukocyte_antigen" title="Human leukocyte antigen">HLA</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lymphocyte" title="Lymphocyte">Lymphocytes</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell-mediated_immunity" title="Cell-mediated immunity">Cellular</a> <ul><li><a href="/wiki/T_cell" title="T cell">T cell</a></li></ul></li> <li><a href="/wiki/Humoral_immunity" title="Humoral immunity">Humoral</a> <ul><li><a href="/wiki/B_cell" title="B cell">B cell</a></li></ul></li> <li><a class="mw-selflink selflink">NK cell</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Substances</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a></li> <li><a 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