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and landmarks subsection </button> <ul id="toc-Nuclear_structures_and_landmarks-sublist" class="vector-toc-list"> <li id="toc-Nuclear_envelope_and_pores" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nuclear_envelope_and_pores"> <div class="vector-toc-text"> 2.1 Nuclear envelope and pores </div> </a> <ul id="toc-Nuclear_envelope_and_pores-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nuclear_lamina" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nuclear_lamina"> <div class="vector-toc-text"> 2.2 Nuclear lamina </div> </a> <ul id="toc-Nuclear_lamina-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nucleolus" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nucleolus"> <div class="vector-toc-text"> 2.3 Nucleolus </div> </a> <ul id="toc-Nucleolus-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Splicing_speckles" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Splicing_speckles"> <div class="vector-toc-text"> 2.4 Splicing speckles </div> </a> <ul id="toc-Splicing_speckles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cajal_bodies_and_gems" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cajal_bodies_and_gems"> <div class="vector-toc-text"> 2.5 Cajal bodies and gems </div> </a> <ul id="toc-Cajal_bodies_and_gems-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_nuclear_bodies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Other_nuclear_bodies"> <div class="vector-toc-text"> 2.6 Other nuclear bodies </div> </a> <ul id="toc-Other_nuclear_bodies-sublist" class="vector-toc-list"> <li id="toc-PIKA_and_PTF_domains" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#PIKA_and_PTF_domains"> <div class="vector-toc-text"> 2.6.1 PIKA and PTF domains </div> </a> <ul id="toc-PIKA_and_PTF_domains-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-PML-nuclear_bodies" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#PML-nuclear_bodies"> <div class="vector-toc-text"> 2.6.2 PML-nuclear bodies </div> </a> <ul id="toc-PML-nuclear_bodies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Paraspeckles" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Paraspeckles"> <div class="vector-toc-text"> 2.6.3 Paraspeckles </div> </a> <ul id="toc-Paraspeckles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Perichromatin_fibrils" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Perichromatin_fibrils"> <div class="vector-toc-text"> 2.6.4 Perichromatin fibrils </div> </a> <ul id="toc-Perichromatin_fibrils-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clastosomes" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Clastosomes"> <div class="vector-toc-text"> 2.6.5 Clastosomes </div> </a> <ul id="toc-Clastosomes-sublist" class="vector-toc-list"> </ul> </li> </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"> 3 Function </div> </a> <button aria-controls="toc-Function-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Function subsection </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Cell_compartmentalization" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cell_compartmentalization"> <div class="vector-toc-text"> 3.1 Cell compartmentalization </div> </a> <ul id="toc-Cell_compartmentalization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Replication" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Replication"> <div class="vector-toc-text"> 3.2 Replication </div> </a> <ul id="toc-Replication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gene_expression" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Gene_expression"> <div class="vector-toc-text"> 3.3 Gene expression </div> </a> <ul id="toc-Gene_expression-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Processing_of_pre-mRNA" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Processing_of_pre-mRNA"> <div class="vector-toc-text"> 3.4 Processing of pre-mRNA </div> </a> <ul id="toc-Processing_of_pre-mRNA-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Dynamics_and_regulation" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Dynamics_and_regulation"> <div class="vector-toc-text"> 4 Dynamics and regulation </div> </a> <button aria-controls="toc-Dynamics_and_regulation-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Dynamics and regulation subsection </button> <ul id="toc-Dynamics_and_regulation-sublist" class="vector-toc-list"> <li id="toc-Nuclear_transport" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nuclear_transport"> <div class="vector-toc-text"> 4.1 Nuclear transport </div> </a> <ul id="toc-Nuclear_transport-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Assembly_and_disassembly" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Assembly_and_disassembly"> <div class="vector-toc-text"> 4.2 Assembly and disassembly </div> </a> <ul id="toc-Assembly_and_disassembly-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Disease-related_dynamics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Disease-related_dynamics"> <div class="vector-toc-text"> 4.3 Disease-related dynamics </div> </a> <ul id="toc-Disease-related_dynamics-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Nuclei_per_cell" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Nuclei_per_cell"> <div class="vector-toc-text"> 5 Nuclei per cell </div> </a> <button aria-controls="toc-Nuclei_per_cell-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Nuclei per cell subsection </button> <ul id="toc-Nuclei_per_cell-sublist" class="vector-toc-list"> <li id="toc-Anucleated_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anucleated_cells"> <div class="vector-toc-text"> 5.1 Anucleated cells </div> </a> <ul id="toc-Anucleated_cells-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Multinucleated_cells" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Multinucleated_cells"> <div class="vector-toc-text"> 5.2 Multinucleated cells </div> </a> <ul id="toc-Multinucleated_cells-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> 6 Evolution </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> 7 History </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </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"> 8 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 9 References </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"> 10 Further reading </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"> 11 External links </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" > Toggle the table of contents </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Cell nucleus</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 100 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-100" aria-hidden="true" > 100 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Selkern" title="Selkern – Afrikaans" lang="af" hreflang="af" data-title="Selkern" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-als mw-list-item"><a href="https://als.wikipedia.org/wiki/Zellkern" title="Zellkern – Alemannic" lang="gsw" hreflang="gsw" data-title="Zellkern" data-language-autonym="Alemannisch" data-language-local-name="Alemannic" class="interlanguage-link-target">Alemannisch</a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D9%86%D9%88%D8%A7%D8%A9_(%D8%AE%D9%84%D9%8A%D8%A9)" title="نواة (خلية) – Arabic" lang="ar" hreflang="ar" data-title="نواة (خلية)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Nuclio_celular" title="Nuclio celular – Aragonese" lang="an" hreflang="an" data-title="Nuclio celular" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Nucleu_celular" title="Nucleu celular – Asturian" lang="ast" hreflang="ast" data-title="Nucleu celular" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/H%C3%BCceyr%C9%99_n%C3%BCv%C9%99si" title="Hüceyrə nüvəsi – Azerbaijani" lang="az" hreflang="az" data-title="Hüceyrə nüvəsi" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%95%E0%A7%8B%E0%A6%B7_%E0%A6%A8%E0%A6%BF%E0%A6%89%E0%A6%95%E0%A7%8D%E0%A6%B2%E0%A6%BF%E0%A6%AF%E0%A6%BC%E0%A6%BE%E0%A6%B8" title="কোষ নিউক্লিয়াস – Bangla" lang="bn" hreflang="bn" data-title="কোষ নিউক্লিয়াস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/S%C3%A8-pau-hu%CC%8Dt" title="Sè-pau-hu̍t – Minnan" lang="nan" hreflang="nan" data-title="Sè-pau-hu̍t" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D0%9A%D2%AF%D2%99%D3%99%D0%BD%D3%99%D0%BA_%D1%8F%D0%B4%D1%80%D0%BE%D2%BB%D1%8B" title="Күҙәнәк ядроһы – Bashkir" lang="ba" hreflang="ba" data-title="Күҙәнәк ядроһы" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target">Башҡортса</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B5%D1%82%D0%B0%D1%87%D0%BD%D0%B0%D0%B5_%D1%8F%D0%B4%D1%80%D0%BE" title="Клетачнае ядро – Belarusian" lang="be" hreflang="be" data-title="Клетачнае ядро" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%AF%D0%B4%D1%80%D0%BE_%D0%B2%D1%83%D0%B7%D1%8B" title="Ядро вузы – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Ядро вузы" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target">Беларуская (тарашкевіца)</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B5%D1%82%D1%8A%D1%87%D0%BD%D0%BE_%D1%8F%D0%B4%D1%80%D0%BE" title="Клетъчно ядро – Bulgarian" lang="bg" hreflang="bg" data-title="Клетъчно ядро" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Jedro_(biologija)" title="Jedro (biologija) – Bosnian" lang="bs" hreflang="bs" data-title="Jedro (biologija)" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-ca badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ca.wikipedia.org/wiki/Nucli_cel%C2%B7lular" title="Nucli cel·lular – Catalan" lang="ca" hreflang="ca" data-title="Nucli cel·lular" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%9F%D0%B0%D0%B9%D1%80%C4%83%D0%BC_%D1%82%C4%95%D1%88%C4%95" title="Пайрăм тĕшĕ – Chuvash" lang="cv" hreflang="cv" data-title="Пайрăм тĕшĕ" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target">Чӑвашла</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Bun%C4%9B%C4%8Dn%C3%A9_j%C3%A1dro" title="Buněčné jádro – Czech" lang="cs" hreflang="cs" data-title="Buněčné jádro" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Cnewyllyn_cell" title="Cnewyllyn cell – Welsh" lang="cy" hreflang="cy" data-title="Cnewyllyn cell" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Cellekerne" title="Cellekerne – Danish" lang="da" hreflang="da" data-title="Cellekerne" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-se mw-list-item"><a href="https://se.wikipedia.org/wiki/Seallav%C3%A1immus" title="Seallaváimmus – Northern Sami" lang="se" hreflang="se" data-title="Seallaváimmus" data-language-autonym="Davvisámegiella" data-language-local-name="Northern Sami" class="interlanguage-link-target">Davvisámegiella</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Zellkern" title="Zellkern – German" lang="de" hreflang="de" data-title="Zellkern" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Rakutuum" title="Rakutuum – Estonian" lang="et" hreflang="et" data-title="Rakutuum" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CF%85%CF%84%CF%84%CE%B1%CF%81%CE%B9%CE%BA%CF%8C%CF%82_%CF%80%CF%85%CF%81%CE%AE%CE%BD%CE%B1%CF%82" title="Κυτταρικός πυρήνας – Greek" lang="el" hreflang="el" data-title="Κυτταρικός πυρήνας" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/N%C3%BAcleo_celular" title="Núcleo celular – Spanish" lang="es" hreflang="es" data-title="Núcleo celular" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/%C4%88elkerno" title="Ĉelkerno – Esperanto" lang="eo" hreflang="eo" data-title="Ĉelkerno" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Zelula_nukleo" title="Zelula nukleo – Basque" lang="eu" hreflang="eu" data-title="Zelula nukleo" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%87%D8%B3%D8%AA%D9%87_%D8%B3%D9%84%D9%88%D9%84" title="هسته سلول – Persian" lang="fa" hreflang="fa" data-title="هسته سلول" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Noyau_(biologie)" title="Noyau (biologie) – French" lang="fr" hreflang="fr" data-title="Noyau (biologie)" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/N%C3%BAicl%C3%A9as_cille" title="Núicléas cille – Irish" lang="ga" hreflang="ga" data-title="Núicléas cille" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gv mw-list-item"><a href="https://gv.wikipedia.org/wiki/%C3%87heshvean_killag" title="Çheshvean killag – Manx" lang="gv" hreflang="gv" data-title="Çheshvean killag" data-language-autonym="Gaelg" data-language-local-name="Manx" class="interlanguage-link-target">Gaelg</a></li><li class="interlanguage-link interwiki-gl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://gl.wikipedia.org/wiki/N%C3%BAcleo_celular" title="Núcleo celular – Galician" lang="gl" hreflang="gl" data-title="Núcleo celular" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-inh mw-list-item"><a href="https://inh.wikipedia.org/wiki/%D0%9E%D0%B0%D0%B3%D0%B8%D0%BB%D0%B3%D0%B0_%D1%82%D3%80%D1%83%D0%BC" title="Оагилга тӀум – Ingush" lang="inh" hreflang="inh" data-title="Оагилга тӀум" data-language-autonym="ГӀалгӀай" data-language-local-name="Ingush" class="interlanguage-link-target">ГӀалгӀай</a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/Se-p%C3%A2u-fu%CC%8Dt" title="Se-pâu-fu̍t – Hakka Chinese" lang="hak" hreflang="hak" data-title="Se-pâu-fu̍t" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target">客家語 / Hak-kâ-ngî</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%84%B8%ED%8F%AC%ED%95%B5" title="세포핵 – Korean" lang="ko" hreflang="ko" data-title="세포핵" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D5%BB%D5%BB%D5%A1%D5%AF%D5%B8%D6%80%D5%AB%D5%A6" title="Բջջակորիզ – Armenian" lang="hy" hreflang="hy" data-title="Բջջակորիզ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%95%E0%A5%8B%E0%A4%B6%E0%A4%BF%E0%A4%95%E0%A4%BE_%E0%A4%95%E0%A5%87%E0%A4%A8%E0%A5%8D%E0%A4%A6%E0%A5%8D%E0%A4%B0%E0%A4%95" title="कोशिका केन्द्रक – Hindi" lang="hi" hreflang="hi" data-title="कोशिका केन्द्रक" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Stani%C4%8Dna_jezgra" title="Stanična jezgra – Croatian" lang="hr" hreflang="hr" data-title="Stanična jezgra" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Inti_sel" title="Inti sel – Indonesian" lang="id" hreflang="id" data-title="Inti sel" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Nucleo_(biologia)" title="Nucleo (biologia) – Interlingua" lang="ia" hreflang="ia" data-title="Nucleo (biologia)" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target">Interlingua</a></li><li class="interlanguage-link interwiki-zu mw-list-item"><a href="https://zu.wikipedia.org/wiki/INdeni_yengqamuzana" title="INdeni yengqamuzana – Zulu" lang="zu" hreflang="zu" data-title="INdeni yengqamuzana" data-language-autonym="IsiZulu" data-language-local-name="Zulu" class="interlanguage-link-target">IsiZulu</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Frumukjarni" title="Frumukjarni – Icelandic" lang="is" hreflang="is" data-title="Frumukjarni" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Nucleo_cellulare" title="Nucleo cellulare – Italian" lang="it" hreflang="it" data-title="Nucleo cellulare" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%92%D7%A8%D7%A2%D7%99%D7%9F_%D7%94%D7%AA%D7%90" title="גרעין התא – Hebrew" lang="he" hreflang="he" data-title="גרעין התא" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Inti_sel" title="Inti sel – Javanese" lang="jv" hreflang="jv" data-title="Inti sel" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%9C%E0%B3%80%E0%B2%B5%E0%B2%95%E0%B2%A3%E0%B2%A6_%E0%B2%AC%E0%B3%80%E0%B2%9C%E0%B2%95%E0%B2%A3" title="ಜೀವಕಣದ ಬೀಜಕಣ – Kannada" lang="kn" hreflang="kn" data-title="ಜೀವಕಣದ ಬೀಜಕಣ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A3%E1%83%AF%E1%83%A0%E1%83%94%E1%83%93%E1%83%98%E1%83%A1_%E1%83%91%E1%83%98%E1%83%A0%E1%83%97%E1%83%95%E1%83%98" title="უჯრედის ბირთვი – Georgian" lang="ka" hreflang="ka" data-title="უჯრედის ბირთვი" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%96%D0%B0%D1%81%D1%83%D1%88%D0%B0_%D1%8F%D0%B4%D1%80%D0%BE%D1%81%D1%8B" title="Жасуша ядросы – Kazakh" lang="kk" hreflang="kk" data-title="Жасуша ядросы" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Kiini_cha_seli" title="Kiini cha seli – Swahili" lang="sw" hreflang="sw" data-title="Kiini cha seli" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target">Kiswahili</a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Nwayo_(selil)" title="Nwayo (selil) – Haitian Creole" lang="ht" hreflang="ht" data-title="Nwayo (selil)" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target">Kreyòl ayisyen</a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Navik" title="Navik – Kurdish" lang="ku" hreflang="ku" data-title="Navik" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target">Kurdî</a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%AF%D0%B4%D1%80%D0%BE_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F)" title="Ядро (биология) – Kyrgyz" lang="ky" hreflang="ky" data-title="Ядро (биология)" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target">Кыргызча</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Nucleus_cellulae" title="Nucleus cellulae – Latin" lang="la" hreflang="la" data-title="Nucleus cellulae" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/%C5%A0%C5%ABnas_kodols" title="Šūnas kodols – Latvian" lang="lv" hreflang="lv" data-title="Šūnas kodols" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Zellk%C3%A4r" title="Zellkär – Luxembourgish" lang="lb" hreflang="lb" data-title="Zellkär" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target">Lëtzebuergesch</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/L%C4%85stel%C4%97s_branduolys" title="Ląstelės branduolys – Lithuanian" lang="lt" hreflang="lt" data-title="Ląstelės branduolys" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-li mw-list-item"><a href="https://li.wikipedia.org/wiki/Celkern" title="Celkern – Limburgish" lang="li" hreflang="li" data-title="Celkern" data-language-autonym="Limburgs" data-language-local-name="Limburgish" class="interlanguage-link-target">Limburgs</a></li><li class="interlanguage-link interwiki-hu badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://hu.wikipedia.org/wiki/Sejtmag" title="Sejtmag – Hungarian" lang="hu" hreflang="hu" data-title="Sejtmag" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B5%D1%82%D0%BE%D1%87%D0%BD%D0%BE_%D1%98%D0%B0%D0%B4%D1%80%D0%BE" title="Клеточно јадро – Macedonian" lang="mk" hreflang="mk" data-title="Клеточно јадро" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%95%E0%B5%8B%E0%B4%B6%E0%B4%AE%E0%B5%BC%E0%B4%AE%E0%B5%8D%E0%B4%AE%E0%B4%82" title="കോശമർമ്മം – Malayalam" lang="ml" hreflang="ml" data-title="കോശമർമ്മം" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-xmf mw-list-item"><a href="https://xmf.wikipedia.org/wiki/%E1%83%97%E1%83%9D%E1%83%9A%E1%83%98%E1%83%A8_%E1%83%92%E1%83%95%E1%83%98%E1%83%A0%E1%83%96%E1%83%98%E1%83%9A%E1%83%98" title="თოლიშ გვირზილი – Mingrelian" lang="xmf" hreflang="xmf" data-title="თოლიშ გვირზილი" data-language-autonym="მარგალური" data-language-local-name="Mingrelian" class="interlanguage-link-target">მარგალური</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Nukleus_sel" title="Nukleus sel – Malay" lang="ms" hreflang="ms" data-title="Nukleus sel" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%AD%D1%81%D0%B8%D0%B9%D0%BD_%D0%B1%D3%A9%D3%A9%D0%BC" title="Эсийн бөөм – Mongolian" lang="mn" hreflang="mn" data-title="Эсийн бөөм" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target">Монгол</a></li><li class="interlanguage-link interwiki-nl badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://nl.wikipedia.org/wiki/Celkern" title="Celkern – Dutch" lang="nl" hreflang="nl" data-title="Celkern" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B4%B0%E8%83%9E%E6%A0%B8" title="細胞核 – Japanese" lang="ja" hreflang="ja" data-title="細胞核" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Cellekjerne" title="Cellekjerne – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Cellekjerne" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Sellekjerne" title="Sellekjerne – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Sellekjerne" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Nucl%C3%A8u_(biologia)" title="Nuclèu (biologia) – Occitan" lang="oc" hreflang="oc" data-title="Nuclèu (biologia)" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-om mw-list-item"><a href="https://om.wikipedia.org/wiki/Wirmaata" title="Wirmaata – Oromo" lang="om" hreflang="om" data-title="Wirmaata" data-language-autonym="Oromoo" data-language-local-name="Oromo" class="interlanguage-link-target">Oromoo</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Hujayra_yadrosi" title="Hujayra yadrosi – Uzbek" lang="uz" hreflang="uz" data-title="Hujayra yadrosi" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D9%86%DB%8C%D9%88%DA%A9%D9%84%DB%8C%D8%B3_(%D9%88%D9%84%DA%AF%D9%86)" title="نیوکلیس (ولگن) – Western Punjabi" lang="pnb" hreflang="pnb" data-title="نیوکلیس (ولگن)" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target">پنجابی</a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/Zellkarn" title="Zellkarn – Low German" lang="nds" hreflang="nds" data-title="Zellkarn" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target">Plattdüütsch</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/J%C4%85dro_kom%C3%B3rkowe" title="Jądro komórkowe – Polish" lang="pl" hreflang="pl" data-title="Jądro komórkowe" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/N%C3%BAcleo_celular" title="Núcleo celular – Portuguese" lang="pt" hreflang="pt" data-title="Núcleo celular" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Nucleu_celular" title="Nucleu celular – Romanian" lang="ro" hreflang="ro" data-title="Nucleu celular" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-rue mw-list-item"><a href="https://rue.wikipedia.org/wiki/%D0%91%D1%83%D0%BD%D0%BA%D0%BE%D0%B2%D0%B5_%D1%8F%D0%B4%D1%80%D0%BE" title="Бункове ядро – Rusyn" lang="rue" hreflang="rue" data-title="Бункове ядро" data-language-autonym="Русиньскый" data-language-local-name="Rusyn" class="interlanguage-link-target">Русиньскый</a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B5%D1%82%D0%BE%D1%87%D0%BD%D0%BE%D0%B5_%D1%8F%D0%B4%D1%80%D0%BE" title="Клеточное ядро – Russian" lang="ru" hreflang="ru" data-title="Клеточное ядро" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-sco mw-list-item"><a href="https://sco.wikipedia.org/wiki/Cell_nucleus" title="Cell nucleus – Scots" lang="sco" hreflang="sco" data-title="Cell nucleus" data-language-autonym="Scots" data-language-local-name="Scots" class="interlanguage-link-target">Scots</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cell_nucleus" title="Cell nucleus – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cell nucleus" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Bunkov%C3%A9_jadro" title="Bunkové jadro – Slovak" lang="sk" hreflang="sk" data-title="Bunkové jadro" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Celi%C4%8Dno_jedro" title="Celično jedro – Slovenian" lang="sl" hreflang="sl" data-title="Celično jedro" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%86%D8%A7%D9%88%DA%A9%DB%95%D8%AE%D8%A7%D9%86%DB%95" title="ناوکەخانە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="ناوکەخانە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%88%D0%B5%D0%B4%D1%80%D0%BE" title="Једро – Serbian" lang="sr" hreflang="sr" data-title="Једро" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Jezgra_(stanica)" title="Jezgra (stanica) – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Jezgra (stanica)" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Inti_s%C3%A9l" title="Inti sél – Sundanese" lang="su" hreflang="su" data-title="Inti sél" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target">Sunda</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Tuma" title="Tuma – Finnish" lang="fi" hreflang="fi" data-title="Tuma" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Cellk%C3%A4rna" title="Cellkärna – Swedish" lang="sv" hreflang="sv" data-title="Cellkärna" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Nukleyus_ng_selula" title="Nukleyus ng selula – Tagalog" lang="tl" hreflang="tl" data-title="Nukleyus ng selula" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%89%E0%AE%AF%E0%AE%BF%E0%AE%B0%E0%AE%A3%E0%AF%81%E0%AE%95%E0%AF%8D_%E0%AE%95%E0%AE%B0%E0%AF%81" title="உயிரணுக் கரு – Tamil" lang="ta" hreflang="ta" data-title="உயிரணுக் கரு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-tt 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id="mw-content-subtitle">(Redirected from <a href="/w/index.php?title=Nucleated&redirect=no" class="mw-redirect" title="Nucleated">Nucleated</a>)</div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Eukaryotic membrane-bounded organelle containing DNA</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:HeLa_cells_stained_with_Hoechst_33258.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/HeLa_cells_stained_with_Hoechst_33258.jpg/300px-HeLa_cells_stained_with_Hoechst_33258.jpg" decoding="async" width="300" height="229" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/HeLa_cells_stained_with_Hoechst_33258.jpg/450px-HeLa_cells_stained_with_Hoechst_33258.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/6/6c/HeLa_cells_stained_with_Hoechst_33258.jpg 2x" data-file-width="459" data-file-height="350" /></a><figcaption><a href="/wiki/HeLa" title="HeLa">HeLa cells</a> stained for <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a> with the blue <a href="/wiki/Fluorescent" class="mw-redirect" title="Fluorescent">fluorescent</a> <a href="/wiki/Hoechst_stain" title="Hoechst stain">Hoechst dye</a>. The central and rightmost cells are in <a href="/wiki/Interphase" title="Interphase">interphase</a>, thus their entire nuclei are labeled. On the left, a cell is going through <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> and its DNA has condensed.</figcaption></figure> <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" style="width: 300px; padding: 0; border-collapse:collapse; background:transparent;"><tbody><tr><th colspan="2" class="infobox-above" style="background: rgba(0,0,0,0.1); padding: 5px; width: 290px;"><a href="/wiki/Cell_biology" title="Cell biology">Cell biology</a></th></tr><tr><td colspan="2" class="infobox-subheader" style="font-weight:bold; font-size:125%; background-color: #0073CF; color: #FFF;"><a href="/wiki/Animal_cell" class="mw-redirect" title="Animal cell">Animal cell diagram</a></td></tr><tr><td colspan="2" class="infobox-image" style="padding: 0;"><a href="/wiki/File:Animal_Cell.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/300px-Animal_Cell.svg.png" decoding="async" width="300" height="173" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/450px-Animal_Cell.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/11/Animal_Cell.svg/600px-Animal_Cell.svg.png 2x" data-file-width="1405" data-file-height="811" /></a><div class="infobox-caption" style="text-align:left; padding: 10px; background: rgba(0,0,0,0.02);">Components of a typical animal cell: <ol><li><a href="/wiki/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a class="mw-selflink selflink">Nucleus</a></li> <li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a> (dots as part of 5)</li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a></li> <li><a href="/wiki/Rough_endoplasmic_reticulum" class="mw-redirect" title="Rough endoplasmic reticulum">Rough endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> (or, Golgi body)</li> <li><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</a></li> <li><a href="/wiki/Smooth_endoplasmic_reticulum" class="mw-redirect" title="Smooth endoplasmic reticulum">Smooth endoplasmic reticulum</a></li> <li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a> (fluid that contains <a href="/wiki/Organelle" title="Organelle">organelles</a>; with which, comprises <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>)</li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Cell_membrane" title="Cell membrane">Cell membrane</a></li></ol></div> </td></tr></tbody></table> The cell nucleus (from <a href="/wiki/Latin_language" class="mw-redirect" title="Latin language">Latin</a> <a href="https://en.wiktionary.org/wiki/nucleus#Latin" class="extiw" title="wikt:nucleus">nucleus</a> or <a href="https://en.wiktionary.org/wiki/nuculeus#Latin" class="extiw" title="wikt:nuculeus">nuculeus</a> 'kernel, seed'; <abbr title="plural form">pl.</abbr>: nuclei) is a <a href="/wiki/Membrane-bound_organelle" class="mw-redirect" title="Membrane-bound organelle">membrane-bound organelle</a> found in <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> <a href="/wiki/Cell_(biology)" title="Cell (biology)">cells</a>. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian <a href="/wiki/Red_blood_cell" title="Red blood cell">red blood cells</a>, have <a href="#Anucleated_cells">no nuclei</a>, and a few others including <a href="/wiki/Osteoclast" title="Osteoclast">osteoclasts</a> have <a href="/wiki/Multinucleate" title="Multinucleate">many</a>. The main structures making up the nucleus are the <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a>, a double membrane that encloses the entire organelle and isolates its contents from the cellular <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>; and the <a href="/wiki/Nuclear_matrix" title="Nuclear matrix">nuclear matrix</a>, a network within the nucleus that adds mechanical support. The cell nucleus contains nearly all of the cell's <a href="/wiki/Genome" title="Genome">genome</a>. <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">Nuclear DNA</a> is often organized into multiple <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> – long strands of <a href="/wiki/DNA" title="DNA">DNA</a> dotted with various <a href="/wiki/Protein" title="Protein">proteins</a>, such as <a href="/wiki/Histone" title="Histone">histones</a>, that protect and organize the DNA. The <a href="/wiki/Gene" title="Gene">genes</a> within these chromosomes are <a href="/wiki/Nuclear_organization" title="Nuclear organization">structured</a> in such a way to promote cell function. The nucleus maintains the integrity of genes and controls the activities of the cell by regulating <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a>. Because the nuclear envelope is impermeable to <a href="/wiki/Macromolecule" title="Macromolecule">large molecules</a>, <a href="/wiki/Nuclear_pore" title="Nuclear pore">nuclear pores</a> are required to regulate <a href="/wiki/Nuclear_transport" title="Nuclear transport">nuclear transport</a> of molecules across the envelope. The pores cross both nuclear membranes, providing a <a href="/wiki/Ion_channel" title="Ion channel">channel</a> through which larger molecules must be <a href="/wiki/Active_transport" title="Active transport">actively transported</a> by carrier proteins while allowing free movement of small molecules and <a href="/wiki/Ion" title="Ion">ions</a>. Movement of large molecules such as proteins and <a href="/wiki/RNA" title="RNA">RNA</a> through the pores is required for both gene expression and the maintenance of chromosomes. Although the interior of the nucleus does not contain any membrane-bound subcompartments, a number of <a href="/wiki/Nuclear_bodies" title="Nuclear bodies">nuclear bodies</a> exist, made up of unique proteins, RNA molecules, and particular parts of the chromosomes. The best-known of these is the <a href="/wiki/Nucleolus" title="Nucleolus">nucleolus</a>, involved in the assembly of <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Chromosomes">Chromosomes</h2></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Chromosome" title="Chromosome">Chromosome</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Nuclear_organization" title="Nuclear organization">Nuclear organization</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:MouseChromosomeTerritoriesBMC_Cell_Biol6-44Fig2e.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/MouseChromosomeTerritoriesBMC_Cell_Biol6-44Fig2e.jpg/200px-MouseChromosomeTerritoriesBMC_Cell_Biol6-44Fig2e.jpg" decoding="async" width="200" height="200" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/f/f4/MouseChromosomeTerritoriesBMC_Cell_Biol6-44Fig2e.jpg 1.5x" data-file-width="235" data-file-height="235" /></a><figcaption>A mouse <a href="/wiki/Fibroblast" title="Fibroblast">fibroblast</a> nucleus in which <a href="/wiki/DNA" title="DNA">DNA</a> is stained blue. The distinct chromosome territories of chromosome 2 (red) and chromosome 9 (green) are stained with <a href="/wiki/Fluorescent_in_situ_hybridization" class="mw-redirect" title="Fluorescent in situ hybridization">fluorescent in situ hybridization</a>.</figcaption></figure> The cell nucleus contains the majority of the cell's genetic material in the form of multiple linear DNA molecules organized into structures called <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>. Each human cell contains roughly two meters of DNA.<a href="#cite_note-Lodish-1">[1]</a>: 405  During most of the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a> these are organized in a DNA-protein complex known as <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>, and during cell division the chromatin can be seen to form the well-defined chromosomes familiar from a <a href="/wiki/Karyotype" title="Karyotype">karyotype</a>. A small fraction of the cell's genes are located instead in the <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a>.<a href="#cite_note-Lodish-1">[1]</a>: 438  There are two types of chromatin. <a href="/wiki/Euchromatin" title="Euchromatin">Euchromatin</a> is the less compact DNA form, and contains genes that are frequently <a href="/wiki/Gene_expression" title="Gene expression">expressed</a> by the cell.<a href="#cite_note-Ehrenhofer-2">[2]</a> The other type, <a href="/wiki/Heterochromatin" title="Heterochromatin">heterochromatin</a>, is the more compact form, and contains DNA that is infrequently transcribed. This structure is further categorized into <a href="/wiki/Facultative_heterochromatin#Facultative_heterochromatin" class="mw-redirect" title="Facultative heterochromatin">facultative heterochromatin</a>, consisting of genes that are organized as heterochromatin only in certain cell types or at certain stages of development, and <a href="/wiki/Constitutive_heterochromatin" title="Constitutive heterochromatin">constitutive heterochromatin</a> that consists of chromosome structural components such as <a href="/wiki/Telomere" title="Telomere">telomeres</a> and <a href="/wiki/Centromere" title="Centromere">centromeres</a>.<a href="#cite_note-Grigoryev-3">[3]</a> During interphase the chromatin organizes itself into discrete individual patches,<a href="#cite_note-Schardin-4">[4]</a> called <a href="/wiki/Chromosome_territories" title="Chromosome territories">chromosome territories</a>.<a href="#cite_note-Lamond-5">[5]</a> Active genes, which are generally found in the euchromatic region of the chromosome, tend to be located towards the chromosome's territory boundary.<a href="#cite_note-Kurz-6">[6]</a> Antibodies to certain types of chromatin organization, in particular, <a href="/wiki/Nucleosome" title="Nucleosome">nucleosomes</a>, have been associated with a number of <a href="/wiki/Autoimmune_disease" title="Autoimmune disease">autoimmune diseases</a>, such as <a href="/wiki/Systemic_lupus_erythematosus" class="mw-redirect" title="Systemic lupus erythematosus">systemic lupus erythematosus</a>.<a href="#cite_note-Rothfield-7">[7]</a> These are known as <a href="/wiki/Anti-nuclear_antibody" class="mw-redirect" title="Anti-nuclear antibody">anti-nuclear antibodies</a> (ANA) and have also been observed in concert with <a href="/wiki/Multiple_sclerosis" title="Multiple sclerosis">multiple sclerosis</a> as part of general immune system dysfunction.<a href="#cite_note-Barned-8">[8]</a> <div class="mw-heading mw-heading2"><h2 id="Nuclear_structures_and_landmarks">Nuclear structures and landmarks</h2></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Nuclear_equivalence" title="Nuclear equivalence">Nuclear equivalence</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Diagram_human_cell_nucleus.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Diagram_human_cell_nucleus.svg/280px-Diagram_human_cell_nucleus.svg.png" decoding="async" width="280" height="229" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/38/Diagram_human_cell_nucleus.svg/420px-Diagram_human_cell_nucleus.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/38/Diagram_human_cell_nucleus.svg/560px-Diagram_human_cell_nucleus.svg.png 2x" data-file-width="462" data-file-height="378" /></a><figcaption>Diagram of the nucleus showing the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a>-studded <a href="/wiki/Outer_nuclear_membrane" class="mw-redirect" title="Outer nuclear membrane">outer nuclear membrane</a>, <a href="/wiki/Nuclear_pores" class="mw-redirect" title="Nuclear pores">nuclear pores</a>, <a href="/wiki/DNA" title="DNA">DNA</a> (complexed as <a href="/wiki/Chromatin" title="Chromatin">chromatin</a>), and the <a href="/wiki/Nucleolus" title="Nucleolus">nucleolus</a>.</figcaption></figure> The nucleus contains nearly all of the cell's <a href="/wiki/DNA" title="DNA">DNA</a>, surrounded by a network of fibrous <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a> called the <a href="/wiki/Nuclear_matrix" title="Nuclear matrix">nuclear matrix</a>, and is enveloped in a double membrane called the <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a>. The nuclear envelope separates the fluid inside the nucleus, called the <a href="/wiki/Nucleoplasm" title="Nucleoplasm">nucleoplasm</a>, from the rest of the cell. The size of the nucleus is correlated to the size of the cell, and this <a href="/wiki/NC_ratio" class="mw-redirect" title="NC ratio">ratio</a> is reported across a range of cell types and species.<a href="#cite_note-Kume-9">[9]</a> In eukaryotes the nucleus in many cells typically occupies 10% of the cell volume.<a href="#cite_note-Alberts2015-10">[10]</a>: 178  The nucleus is the largest <a href="/wiki/Organelle" title="Organelle">organelle</a> in animal cells.<a href="#cite_note-Lodish_2016-11">[11]</a>: 12  In human cells, the diameter of the nucleus is approximately six <a href="/wiki/Micrometre" title="Micrometre">micrometres</a> (μm).<a href="#cite_note-Alberts2015-10">[10]</a>: 179  <div class="mw-heading mw-heading3"><h3 id="Nuclear_envelope_and_pores">Nuclear envelope and pores</h3></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">Nuclear envelope</a> and <a href="/wiki/Nuclear_pore" title="Nuclear pore">Nuclear pore</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:NuclearPore_crop.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/NuclearPore_crop.svg/250px-NuclearPore_crop.svg.png" decoding="async" width="250" height="208" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/NuclearPore_crop.svg/375px-NuclearPore_crop.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c8/NuclearPore_crop.svg/500px-NuclearPore_crop.svg.png 2x" data-file-width="726" data-file-height="604" /></a><figcaption>A cross section of a <a href="/wiki/Nuclear_pore" title="Nuclear pore">nuclear pore</a> on the surface of the <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a> (1). Other diagram labels show (2) the outer ring, (3) spokes, (4) basket, and (5) filaments.</figcaption></figure> The <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a> consists of two <a href="/wiki/Cell_membrane" title="Cell membrane">membranes</a>, an <a href="/wiki/Inner_nuclear_membrane" class="mw-redirect" title="Inner nuclear membrane">inner</a> and an <a href="/wiki/Outer_nuclear_membrane" class="mw-redirect" title="Outer nuclear membrane">outer nuclear membrane</a>, perforated by <a href="/wiki/Nuclear_pore" title="Nuclear pore">nuclear pores</a>.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  Together, these membranes serve to separate the cell's genetic material from the rest of the cell contents, and allow the nucleus to maintain an environment distinct from the rest of the cell. Despite their close apposition around much of the nucleus, the two membranes differ substantially in shape and contents. The inner membrane surrounds the nuclear content, providing its defining edge.<a href="#cite_note-Lodish_2016-11">[11]</a>: 14  Embedded within the inner membrane, various proteins bind the intermediate filaments that give the nucleus its structure.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  The outer membrane encloses the inner membrane, and is continuous with the adjacent <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a> membrane.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  As part of the endoplasmic reticulum membrane, the outer nuclear membrane is studded with <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a> that are actively translating proteins across membrane.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  The space between the two membranes is called the perinuclear space, and is continuous with the endoplasmic reticulum <a href="/wiki/Lumen_(anatomy)" title="Lumen (anatomy)">lumen</a>.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  In a mammalian nuclear envelope there are between 3000 and 4000 <a href="/wiki/Nuclear_pore_complex" class="mw-redirect" title="Nuclear pore complex">nuclear pore complexes</a> (NPCs) perforating the envelope.<a href="#cite_note-Alberts2015-10">[10]</a>: 650  Each NPC contains an eightfold-symmetric ring-shaped structure at a position where the inner and outer membranes fuse.<a href="#cite_note-Shulga-12">[12]</a> The number of NPCs can vary considerably across cell types; small <a href="/wiki/Glial_cell" class="mw-redirect" title="Glial cell">glial cells</a> only have about a few hundred, with large <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cells</a> having around 20,000.<a href="#cite_note-Alberts2015-10">[10]</a>: 650  The NPC provides selective transport of molecules between the <a href="/wiki/Nucleoplasm" title="Nucleoplasm">nucleoplasm</a> and the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a>.<a href="#cite_note-Alberts2019-13">[13]</a> The nuclear pore complex is composed of approximately thirty different proteins known as <a href="/wiki/Nucleoporin" title="Nucleoporin">nucleoporins</a>.<a href="#cite_note-Alberts2015-10">[10]</a>: 649  The pores are about 60–80 million <a href="/wiki/Atomic_mass_unit" class="mw-redirect" title="Atomic mass unit">daltons</a> in <a href="/wiki/Molecular_weight" class="mw-redirect" title="Molecular weight">molecular weight</a> and consist of around 50 (in <a href="/wiki/Yeast" title="Yeast">yeast</a>) to several hundred proteins (in <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a>).<a href="#cite_note-Lodish_2016-11">[11]</a>: 622–4  The pores are 100 nm in total diameter; however, the gap through which molecules freely diffuse is only about 9 nm wide, due to the presence of regulatory systems within the center of the pore. This size selectively allows the passage of small water-soluble molecules while preventing larger molecules, such as <a href="/wiki/Nucleic_acid" title="Nucleic acid">nucleic acids</a> and larger proteins, from inappropriately entering or exiting the nucleus. These large molecules must be actively transported into the nucleus instead. Attached to the ring is a structure called the nuclear basket that extends into the nucleoplasm, and a series of filamentous extensions that reach into the cytoplasm. Both structures serve to mediate binding to nuclear transport proteins.<a href="#cite_note-Lodish-1">[1]</a>: 509–10  Most proteins, ribosomal subunits, and some RNAs are transported through the pore complexes in a process mediated by a family of transport factors known as <a href="/wiki/Karyopherin" title="Karyopherin">karyopherins</a>. Those karyopherins that mediate movement into the nucleus are also called importins, whereas those that mediate movement out of the nucleus are called exportins. Most karyopherins interact directly with their cargo, although some use <a href="/wiki/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">adaptor proteins</a>.<a href="#cite_note-Pemberton-14">[14]</a> <a href="/wiki/Steroid_hormone" title="Steroid hormone">Steroid hormones</a> such as <a href="/wiki/Cortisol" title="Cortisol">cortisol</a> and <a href="/wiki/Aldosterone" title="Aldosterone">aldosterone</a>, as well as other small lipid-soluble molecules involved in intercellular <a href="/wiki/Cell_signaling" title="Cell signaling">signaling</a>, can diffuse through the cell membrane and into the cytoplasm, where they bind <a href="/wiki/Nuclear_receptor" title="Nuclear receptor">nuclear receptor</a> proteins that are trafficked into the nucleus. There they serve as <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> when bound to their <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligand</a>; in the absence of a ligand, many such receptors function as <a href="/wiki/Histone_deacetylase" title="Histone deacetylase">histone deacetylases</a> that repress gene expression.<a href="#cite_note-Lodish-1">[1]</a>: 488  <div class="mw-heading mw-heading3"><h3 id="Nuclear_lamina">Nuclear lamina</h3></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/Nuclear_lamina" title="Nuclear lamina">Nuclear lamina</a></div> In animal cells, two networks of <a href="/wiki/Intermediate_filaments" class="mw-redirect" title="Intermediate filaments">intermediate filaments</a> provide the nucleus with mechanical support: The <a href="/wiki/Nuclear_lamina" title="Nuclear lamina">nuclear lamina</a> forms an organized meshwork on the internal face of the envelope, while less organized support is provided on the cytosolic face of the envelope. Both systems provide structural support for the nuclear envelope and anchoring sites for chromosomes and nuclear pores.<a href="#cite_note-MBoC-15">[15]</a> The nuclear lamina is composed mostly of <a href="/wiki/Lamin" title="Lamin">lamin</a> proteins. Like all proteins, lamins are synthesized in the cytoplasm and later transported to the nucleus interior, where they are assembled before being incorporated into the existing network of nuclear lamina.<a href="#cite_note-Sturrman-16">[16]</a><a href="#cite_note-Goldman-17">[17]</a> Lamins found on the cytosolic face of the membrane, such as <a href="/wiki/Emerin" title="Emerin">emerin</a> and <a href="/wiki/Nesprin" title="Nesprin">nesprin</a>, bind to the cytoskeleton to provide structural support. Lamins are also found inside the nucleoplasm where they form another regular structure, known as the nucleoplasmic veil,<a href="#cite_note-RGoldman-18">[18]</a><a href="#cite_note-Broers_2004-19">[19]</a> that is visible using <a href="/wiki/Fluorescence_microscopy" class="mw-redirect" title="Fluorescence microscopy">fluorescence microscopy</a>. The actual function of the veil is not clear, although it is excluded from the nucleolus and is present during <a href="/wiki/Interphase" title="Interphase">interphase</a>.<a href="#cite_note-Moir-20">[20]</a> Lamin structures that make up the veil, such as <a href="/wiki/LEM_domain-containing_protein_3" title="LEM domain-containing protein 3">LEM3</a>, bind chromatin and disrupting their structure inhibits transcription of protein-coding genes.<a href="#cite_note-Spann-21">[21]</a> Like the components of other intermediate filaments, the lamin <a href="/wiki/Monomer" title="Monomer">monomer</a> contains an <a href="/wiki/Alpha-helix" class="mw-redirect" title="Alpha-helix">alpha-helical</a> domain used by two monomers to coil around each other, forming a <a href="/wiki/Protein_dimer" title="Protein dimer">dimer</a> structure called a <a href="/wiki/Coiled_coil" title="Coiled coil">coiled coil</a>. Two of these dimer structures then join side by side, in an <a href="/wiki/Antiparallel_(biochemistry)" title="Antiparallel (biochemistry)">antiparallel</a> arrangement, to form a <a href="/wiki/Tetramer_protein" class="mw-redirect" title="Tetramer protein">tetramer</a> called a protofilament. Eight of these protofilaments form a lateral arrangement that is twisted to form a ropelike filament. These filaments can be assembled or disassembled in a dynamic manner, meaning that changes in the length of the filament depend on the competing rates of filament addition and removal.<a href="#cite_note-MBoC-15">[15]</a> Mutations in lamin genes leading to defects in filament assembly cause a group of rare genetic disorders known as <a href="/wiki/Laminopathies" class="mw-redirect" title="Laminopathies">laminopathies</a>. The most notable laminopathy is the family of diseases known as <a href="/wiki/Progeria" title="Progeria">progeria</a>, which causes the appearance of premature <a href="/wiki/Aging" class="mw-redirect" title="Aging">aging</a> in those with the condition. The exact mechanism by which the associated <a href="/wiki/Biochemistry" title="Biochemistry">biochemical</a> changes give rise to the aged <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> is not well understood.<a href="#cite_note-Mounkes-22">[22]</a> <div class="mw-heading mw-heading3"><h3 id="Nucleolus">Nucleolus</h3></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/Nucleolus" title="Nucleolus">Nucleolus</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Nuclear_bodies" title="Nuclear bodies">Nuclear bodies</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Micrograph_of_a_cell_nucleus.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/5/57/Micrograph_of_a_cell_nucleus.png" decoding="async" width="200" height="199" class="mw-file-element" data-file-width="200" data-file-height="199" /></a><figcaption>An <a href="/wiki/Electron_micrograph" class="mw-redirect" title="Electron micrograph">electron micrograph</a> of a cell nucleus, showing the darkly stained <a href="/wiki/Nucleolus" title="Nucleolus">nucleolus</a></figcaption></figure> The <a href="/wiki/Nucleolus" title="Nucleolus">nucleolus</a> is the largest of the discrete densely stained, membraneless structures known as <a href="/wiki/Nuclear_bodies" title="Nuclear bodies">nuclear bodies</a> found in the nucleus. It forms around <a href="/wiki/Tandem_repeat" title="Tandem repeat">tandem repeats</a> of <a href="/wiki/Ribosomal_DNA" title="Ribosomal DNA">rDNA</a>, DNA coding for <a href="/wiki/Ribosomal_RNA" title="Ribosomal RNA">ribosomal RNA</a> (rRNA). These regions are called <a href="/wiki/Nucleolar_organizer_regions" class="mw-redirect" title="Nucleolar organizer regions">nucleolar organizer regions</a> (NOR). The main roles of the nucleolus are to synthesize rRNA and <a href="/wiki/Nucleolus#Function_and_ribosome_assembly" title="Nucleolus">assemble ribosomes</a>. The structural cohesion of the nucleolus depends on its activity, as ribosomal assembly in the nucleolus results in the transient association of nucleolar components, facilitating further ribosomal assembly, and hence further association. This model is supported by observations that inactivation of rDNA results in intermingling of nucleolar structures.<a href="#cite_note-Hernandez-Verdun-23">[23]</a> In the first step of ribosome assembly, a protein called <a href="/wiki/RNA_polymerase_I" title="RNA polymerase I">RNA polymerase I</a> transcribes rDNA, which forms a large pre-rRNA precursor. This is cleaved into two <a href="/wiki/LSU_rRNA" title="LSU rRNA">large rRNA subunits</a> – <a href="/wiki/5.8S_ribosomal_RNA" title="5.8S ribosomal RNA">5.8S</a>, and <a href="/wiki/28S_ribosomal_RNA" title="28S ribosomal RNA">28S</a>, and a <a href="/wiki/SSU_rRNA" title="SSU rRNA">small rRNA subunit</a> <a href="/wiki/18S_ribosomal_RNA" title="18S ribosomal RNA">18S</a>.<a href="#cite_note-Alberts2015-10">[10]</a>: 328 <a href="#cite_note-Lamond-Sleeman-24">[24]</a> The transcription, post-transcriptional processing, and assembly of rRNA occurs in the nucleolus, aided by <a href="/wiki/Small_nucleolar_RNA" title="Small nucleolar RNA">small nucleolar RNA</a> (snoRNA) molecules, some of which are derived from spliced <a href="/wiki/Intron" title="Intron">introns</a> from <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNAs</a> encoding genes related to ribosomal function. The assembled ribosomal subunits are the largest structures passed through the <a href="/wiki/Nuclear_pore" title="Nuclear pore">nuclear pores</a>.<a href="#cite_note-Lodish-1">[1]</a>: 526  When observed under the <a href="/wiki/Electron_microscope" title="Electron microscope">electron microscope</a>, the nucleolus can be seen to consist of three distinguishable regions: the innermost fibrillar centers (FCs), surrounded by the dense fibrillar component (DFC) (that contains <a href="/wiki/Fibrillarin" title="Fibrillarin">fibrillarin</a> and <a href="/wiki/Nucleolin" title="Nucleolin">nucleolin</a>), which in turn is bordered by the granular component (GC) (that contains the protein <a href="/wiki/Nucleophosmin" class="mw-redirect" title="Nucleophosmin">nucleophosmin</a>). Transcription of the rDNA occurs either in the FC or at the FC-DFC boundary, and, therefore, when rDNA transcription in the cell is increased, more FCs are detected. Most of the cleavage and modification of rRNAs occurs in the DFC, while the latter steps involving protein assembly onto the ribosomal subunits occur in the GC.<a href="#cite_note-Lamond-Sleeman-24">[24]</a> <div class="mw-heading mw-heading3"><h3 id="Splicing_speckles"> Splicing speckles</h3></div> Speckles are subnuclear structures that are enriched in pre-messenger RNA splicing factors and are located in the interchromatin regions of the nucleoplasm of mammalian cells.<a href="#cite_note-25">[25]</a> At the fluorescence-microscope level they appear as irregular, punctate structures, which vary in size and shape, and when examined by electron microscopy they are seen as clusters of <a href="/wiki/Interchromatin_granules" class="mw-redirect" title="Interchromatin granules">interchromatin granules</a>. Speckles are dynamic structures, and both their protein and RNA-protein components can cycle continuously between speckles and other nuclear locations, including active transcription sites. Speckles can work with <a href="/wiki/P53" title="P53">p53</a> as enhancers of gene activity to directly enhance the activity of certain genes. Moreover, speckle-associating and non-associating p53 gene targets are functionally distinct.<a href="#cite_note-26">[26]</a> Studies on the composition, structure and behaviour of speckles have provided a model for understanding the functional compartmentalization of the nucleus and the organization of the gene-expression machinery<a href="#cite_note-ReferenceA-27">[27]</a> splicing <a href="/wiki/SnRNP" title="SnRNP">snRNPs</a><a href="#cite_note-28">[28]</a><a href="#cite_note-29">[29]</a> and other splicing proteins necessary for pre-mRNA processing.<a href="#cite_note-ReferenceA-27">[27]</a> Because of a cell's changing requirements, the composition and location of these bodies changes according to mRNA transcription and regulation via <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylation</a> of specific proteins.<a href="#cite_note-Handwerger-30">[30]</a> The splicing speckles are also known as nuclear speckles (nuclear specks), splicing factor compartments (SF compartments), interchromatin granule clusters (IGCs), and <a href="/wiki/Snurposome" title="Snurposome">B snurposomes</a>.<a href="#cite_note-31">[31]</a> B snurposomes are found in the amphibian oocyte nuclei and in <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a> embryos. B snurposomes appear alone or attached to the Cajal bodies in the electron micrographs of the amphibian nuclei.<a href="#cite_note-32">[32]</a> While nuclear speckles were originally thought to be storage sites for the splicing factors,<a href="#cite_note-Matera2007_NatureMolCellBio-33">[33]</a> a more recent study demonstrated that organizing genes and pre-mRNA substrates near speckles increases the kinetic efficiency of pre-mRNA splicing, ultimately boosting protein levels by modulation of splicing.<a href="#cite_note-34">[34]</a> <div class="mw-heading mw-heading3"><h3 id="Cajal_bodies_and_gems">Cajal bodies and gems</h3></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cajal-Body-Overview.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Cajal-Body-Overview.svg/220px-Cajal-Body-Overview.svg.png" decoding="async" width="220" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Cajal-Body-Overview.svg/330px-Cajal-Body-Overview.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Cajal-Body-Overview.svg/440px-Cajal-Body-Overview.svg.png 2x" data-file-width="1134" data-file-height="720" /></a><figcaption>Cajal body</figcaption></figure> A nucleus typically contains between one and ten compact structures called <a href="/wiki/Cajal_body" title="Cajal body">Cajal bodies</a> or coiled bodies (CB), whose diameter measures between 0.2 μm and 2.0 μm depending on the cell type and species.<a href="#cite_note-Cioce-35">[35]</a> When seen under an electron microscope, they resemble balls of tangled thread<a href="#cite_note-Pollard-36">[36]</a> and are dense foci of distribution for the protein <a href="/wiki/Coilin" title="Coilin">coilin</a>.<a href="#cite_note-MateraFrey-37">[37]</a> CBs are involved in a number of different roles relating to RNA processing, specifically <a href="/wiki/SnoRNA" class="mw-redirect" title="SnoRNA">small nucleolar RNA</a> (snoRNA) and <a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">small nuclear RNA</a> (snRNA) maturation, and histone mRNA modification.<a href="#cite_note-Cioce-35">[35]</a> Similar to Cajal bodies are Gemini of Cajal bodies, or gems, whose name is derived from the <a href="/wiki/Gemini_(constellation)" title="Gemini (constellation)">Gemini constellation</a> in reference to their close "twin" relationship with CBs. Gems are similar in size and shape to CBs, and in fact are virtually indistinguishable under the microscope.<a href="#cite_note-MateraFrey-37">[37]</a> Unlike CBs, gems do not contain <a href="/wiki/SnRNP" title="SnRNP">small nuclear ribonucleoproteins</a> (snRNPs), but do contain a protein called <a href="/wiki/Survival_of_motor_neuron_protein" class="mw-redirect" title="Survival of motor neuron protein">survival of motor neuron</a> (SMN) whose function relates to snRNP biogenesis. Gems are believed to assist CBs in snRNP biogenesis,<a href="#cite_note-Matera-38">[38]</a> though it has also been suggested from microscopy evidence that CBs and gems are different manifestations of the same structure.<a href="#cite_note-MateraFrey-37">[37]</a> Later ultrastructural studies have shown gems to be twins of Cajal bodies with the difference being in the coilin component; Cajal bodies are SMN positive and coilin positive, and gems are SMN positive and coilin negative.<a href="#cite_note-Navascues-39">[39]</a> <div class="mw-heading mw-heading3"><h3 id="Other_nuclear_bodies">Other nuclear bodies</h3></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/Nuclear_bodies" title="Nuclear bodies">Nuclear bodies</a></div> <table class="wikitable" style="float:right; font-size:100%; margin-left:15px;"> <caption>Subnuclear structure sizes </caption> <tbody><tr bgcolor="#efefef"> <th style="width: 120px" abbr="name">Structure name </th> <th style="width: 130px" abbr="diameter">Structure diameter </th> <th scope="col"><abbr title="Reference">Ref.</abbr> </th></tr> <tr> <td>Cajal bodies</td> <td>0.2–2.0 μm</td> <td><a href="#cite_note-Cioce-35">[35]</a> </td></tr> <tr> <td>Clastosomes </td> <td>0.2–0.5 μm </td> <td><a href="#cite_note-Lafarga-2002-40">[40]</a> </td></tr> <tr> <td>PIKA</td> <td>5 μm</td> <td><a href="#cite_note-Pollard-36">[36]</a> </td></tr> <tr> <td>PML bodies</td> <td>0.2–1.0 μm</td> <td><a href="#cite_note-Dundr-41">[41]</a> </td></tr> <tr> <td>Paraspeckles</td> <td>0.5–1.0 μm</td> <td><a href="#cite_note-42">[42]</a> </td></tr> <tr> <td>Speckles</td> <td>20–25 nm</td> <td><a href="#cite_note-Pollard-36">[36]</a> </td></tr></tbody></table> Beyond the nuclear bodies first described by <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a> above (e.g., nucleolus, nuclear speckles, Cajal bodies) the nucleus contains a number of other nuclear bodies. These include polymorphic interphase karyosomal association (PIKA), promyelocytic leukaemia (PML) bodies, and <a href="/wiki/Paraspeckle" title="Paraspeckle">paraspeckles</a>. Although little is known about a number of these domains, they are significant in that they show that the nucleoplasm is not a uniform mixture, but rather contains organized functional subdomains.<a href="#cite_note-Dundr-41">[41]</a> Other subnuclear structures appear as part of abnormal disease processes. For example, the presence of small intranuclear rods has been reported in some cases of <a href="/wiki/Nemaline_myopathy" title="Nemaline myopathy">nemaline myopathy</a>. This condition typically results from mutations in <a href="/wiki/Actin" title="Actin">actin</a>, and the rods themselves consist of mutant actin as well as other cytoskeletal proteins.<a href="#cite_note-Goebel-43">[43]</a> <div class="mw-heading mw-heading4"><h4 id="PIKA_and_PTF_domains">PIKA and PTF domains</h4></div> PIKA domains, or polymorphic interphase karyosomal associations, were first described in microscopy studies in 1991. Their function remains unclear, though they were not thought to be associated with active DNA replication, transcription, or RNA processing.<a href="#cite_note-Saunders-44">[44]</a> They have been found to often associate with discrete domains defined by dense localization of the transcription factor PTF, which promotes transcription of <a href="/wiki/Small_nuclear_RNA" title="Small nuclear RNA">small nuclear RNA</a> (snRNA).<a href="#cite_note-Pombo-45">[45]</a> <div class="mw-heading mw-heading4"><h4 id="PML-nuclear_bodies">PML-nuclear bodies</h4></div> <a href="/wiki/Promyelocytic_leukemia_protein" title="Promyelocytic leukemia protein">Promyelocytic leukemia protein</a> (PML-nuclear bodies) are spherical bodies found scattered throughout the nucleoplasm, measuring around 0.1–1.0 μm. They are known by a number of other names, including nuclear domain 10 (ND10), Kremer bodies, and PML oncogenic domains.<a href="#cite_note-Zimber-46">[46]</a> PML-nuclear bodies are named after one of their major components, the promyelocytic leukemia protein (PML). They are often seen in the nucleus in association with Cajal bodies and cleavage bodies.<a href="#cite_note-Dundr-41">[41]</a> Pml-/- mice, which are unable to create PML-nuclear bodies, develop normally without obvious ill effects, showing that PML-nuclear bodies are not required for most essential biological processes.<a href="#cite_note-Lallemand2010-47">[47]</a> <div class="mw-heading mw-heading4"><h4 id="Paraspeckles">Paraspeckles</h4></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/Paraspeckle" title="Paraspeckle">Paraspeckle</a></div> Discovered by Fox et al. in 2002, paraspeckles are irregularly shaped compartments in the interchromatin space of the nucleus.<a href="#cite_note-Fox_2010-48">[48]</a> First documented in HeLa cells, where there are generally 10–30 per nucleus,<a href="#cite_note-para2-49">[49]</a> paraspeckles are now known to also exist in all human primary cells, transformed cell lines, and tissue sections.<a href="#cite_note-para3-50">[50]</a> Their name is derived from their distribution in the nucleus; the "para" is short for parallel and the "speckles" refers to the splicing speckles to which they are always in close proximity.<a href="#cite_note-para2-49">[49]</a> Paraspeckles sequester nuclear proteins and RNA and thus appear to function as a molecular sponge<a href="#cite_note-Nakagawa_2018-51">[51]</a> that is involved in the regulation of gene expression.<a href="#cite_note-Pisani_2019-52">[52]</a> Furthermore, paraspeckles are dynamic structures that are altered in response to changes in cellular metabolic activity. They are transcription dependent<a href="#cite_note-Fox_2010-48">[48]</a> and in the absence of RNA Pol II transcription, the paraspeckle disappears and all of its associated protein components (PSP1, p54nrb, PSP2, CFI(m)68, and PSF) form a crescent shaped perinucleolar cap in the nucleolus. This phenomenon is demonstrated during the cell cycle. In the <a href="/wiki/Cell_cycle" title="Cell cycle">cell cycle</a>, paraspeckles are present during <a href="/wiki/Interphase" title="Interphase">interphase</a> and during all of <a href="/wiki/Mitosis" title="Mitosis">mitosis</a> except for <a href="/wiki/Telophase" title="Telophase">telophase</a>. During telophase, when the two daughter nuclei are formed, there is no <a href="/wiki/RNA" title="RNA">RNA</a> Pol II <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcription</a> so the protein components instead form a perinucleolar cap.<a href="#cite_note-para3-50">[50]</a> <div class="mw-heading mw-heading4"><h4 id="Perichromatin_fibrils"> Perichromatin fibrils</h4></div> Perichromatin fibrils are visible only under electron microscope. They are located next to the transcriptionally active chromatin and are hypothesized to be the sites of active <a href="/wiki/Precursor_mRNA" class="mw-redirect" title="Precursor mRNA">pre-mRNA</a> processing.<a href="#cite_note-Matera2007_NatureMolCellBio-33">[33]</a> <div class="mw-heading mw-heading4"><h4 id="Clastosomes">Clastosomes</h4></div> Clastosomes are small nuclear bodies (0.2–0.5 μm) described as having a thick ring-shape due to the peripheral capsule around these bodies.<a href="#cite_note-Lafarga-2002-40">[40]</a> This name is derived from the Greek klastos, broken and soma, body.<a href="#cite_note-Lafarga-2002-40">[40]</a> Clastosomes are not typically present in normal cells, making them hard to detect. They form under high <a href="/wiki/Proteolysis" title="Proteolysis">proteolytic</a> conditions within the nucleus and degrade once there is a decrease in activity or if cells are treated with <a href="/wiki/Proteasome_inhibitor" title="Proteasome inhibitor">proteasome inhibitors</a>.<a href="#cite_note-Lafarga-2002-40">[40]</a><a href="#cite_note-53">[53]</a> The scarcity of clastosomes in cells indicates that they are not required for <a href="/wiki/Proteasome" title="Proteasome">proteasome</a> function.<a href="#cite_note-Carmo-Fonseca-2010-54">[54]</a> <a href="/wiki/Osmotic_shock" title="Osmotic shock">Osmotic stress</a> has also been shown to cause the formation of clastosomes.<a href="#cite_note-55">[55]</a> These nuclear bodies contain catalytic and regulatory subunits of the proteasome and its substrates, indicating that clastosomes are sites for degrading proteins.<a href="#cite_note-Carmo-Fonseca-2010-54">[54]</a> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2></div> The nucleus provides a site for <a href="/wiki/Transcription_(biology)" title="Transcription (biology)">genetic transcription</a> that is segregated from the location of <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a> in the cytoplasm, allowing levels of <a href="/wiki/Gene_regulation" class="mw-redirect" title="Gene regulation">gene regulation</a> that are not available to <a href="/wiki/Prokaryote" title="Prokaryote">prokaryotes</a>. The main function of the cell nucleus is to control <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> and mediate the <a href="/wiki/DNA_replication" title="DNA replication">replication of DNA</a> during the cell cycle.<a href="#cite_note-Lodish-1">[1]</a>: 171  <div class="mw-heading mw-heading3"><h3 id="Cell_compartmentalization">Cell compartmentalization</h3></div> The <a href="/wiki/Nuclear_envelope" title="Nuclear envelope">nuclear envelope</a> allows control of the nuclear contents, and separates them from the rest of the cytoplasm where necessary. This is important for controlling processes on either side of the nuclear membrane: In most cases where a cytoplasmic process needs to be restricted, a key participant is removed to the nucleus, where it interacts with transcription factors to downregulate the production of certain enzymes in the pathway. This regulatory mechanism occurs in the case of <a href="/wiki/Glycolysis" title="Glycolysis">glycolysis</a>, a cellular pathway for breaking down <a href="/wiki/Glucose" title="Glucose">glucose</a> to produce energy. <a href="/wiki/Hexokinase" title="Hexokinase">Hexokinase</a> is an enzyme responsible for the first step of glycolysis, forming <a href="/wiki/Glucose-6-phosphate" class="mw-redirect" title="Glucose-6-phosphate">glucose-6-phosphate</a> from glucose. At high concentrations of <a href="/wiki/Fructose-6-phosphate" class="mw-redirect" title="Fructose-6-phosphate">fructose-6-phosphate</a>, a molecule made later from glucose-6-phosphate, a regulator protein removes hexokinase to the nucleus,<a href="#cite_note-Lehninger-56">[56]</a> where it forms a transcriptional repressor complex with nuclear proteins to reduce the expression of genes involved in glycolysis.<a href="#cite_note-Moreno-57">[57]</a> In order to control which genes are being transcribed, the cell separates some transcription factor proteins responsible for regulating gene expression from physical access to the DNA until they are activated by other signaling pathways. This prevents even low levels of inappropriate gene expression. For example, in the case of <a href="/wiki/NF-%CE%BAB" title="NF-κB">NF-κB</a>-controlled genes, which are involved in most <a href="/wiki/Inflammation" title="Inflammation">inflammatory</a> responses, transcription is induced in response to a <a href="/wiki/Cell_signaling" title="Cell signaling">signal pathway</a> such as that initiated by the signaling molecule <a href="/wiki/TNF-%CE%B1" class="mw-redirect" title="TNF-α">TNF-α</a>, binds to a cell membrane receptor, resulting in the recruitment of signalling proteins, and eventually activating the transcription factor NF-κB. A <a href="/wiki/Nuclear_localisation_signal" class="mw-redirect" title="Nuclear localisation signal">nuclear localisation signal</a> on the NF-κB protein allows it to be transported through the nuclear pore and into the nucleus, where it stimulates the transcription of the target genes.<a href="#cite_note-MBoC-15">[15]</a> The compartmentalization allows the cell to prevent translation of unspliced mRNA.<a href="#cite_note-Gorlich-58">[58]</a> Eukaryotic mRNA contains introns that must be removed before being translated to produce functional proteins. The splicing is done inside the nucleus before the mRNA can be accessed by ribosomes for translation. Without the nucleus, ribosomes would translate newly transcribed (unprocessed) mRNA, resulting in malformed and nonfunctional proteins.<a href="#cite_note-Lodish-1">[1]</a>: 108–15  <div class="mw-heading mw-heading3"><h3 id="Replication">Replication</h3></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/Eukaryotic_DNA_replication" title="Eukaryotic DNA replication">Eukaryotic DNA replication</a></div> The main function of the cell nucleus is to control gene expression and mediate the replication of DNA during the cell cycle.<a href="#cite_note-Lodish-1">[1]</a>: 171  It has been found that replication happens in a localised way in the cell nucleus. In the S phase of interphase of the cell cycle; replication takes place. Contrary to the traditional view of moving replication forks along stagnant DNA, a concept of replication factories emerged, which means replication forks are concentrated towards some immobilised 'factory' regions through which the template DNA strands pass like conveyor belts.<a href="#cite_note-Hozák_1994-59">[59]</a> <div class="mw-heading mw-heading3"><h3 id="Gene_expression">Gene expression</h3></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/Gene_expression" title="Gene expression">Gene expression</a></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Transcription_factories" class="mw-redirect" title="Transcription factories">Transcription factories</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Basic_diagram_of_a_transcription_factory_during_transcription.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Basic_diagram_of_a_transcription_factory_during_transcription.png/290px-Basic_diagram_of_a_transcription_factory_during_transcription.png" decoding="async" width="290" height="189" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/3/35/Basic_diagram_of_a_transcription_factory_during_transcription.png 1.5x" data-file-width="422" data-file-height="275" /></a><figcaption>A generic <a href="/wiki/Transcription_factories" class="mw-redirect" title="Transcription factories">transcription factory</a> during transcription, highlighting the possibility of transcribing more than one gene at a time. The diagram includes 8 RNA polymerases however the number can vary depending on cell type. The image also includes transcription factors and a porous, protein core.</figcaption></figure> Gene expression first involves transcription, in which DNA is used as a template to produce RNA. In the case of genes encoding proteins, that RNA produced from this process is messenger RNA (mRNA), which then needs to be translated by ribosomes to form a protein. As ribosomes are located outside the nucleus, mRNA produced needs to be exported.<a href="#cite_note-60">[60]</a> Since the nucleus is the site of transcription, it also contains a variety of proteins that either directly mediate transcription or are involved in regulating the process. These proteins include <a href="/wiki/Helicase" title="Helicase">helicases</a>, which unwind the double-stranded DNA molecule to facilitate access to it, <a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerases</a>, which bind to the DNA promoter to synthesize the growing RNA molecule, <a href="/wiki/Topoisomerase" title="Topoisomerase">topoisomerases</a>, which change the amount of <a href="/wiki/Supercoil" class="mw-redirect" title="Supercoil">supercoiling</a> in DNA, helping it wind and unwind, as well as a large variety of transcription factors that regulate expression.<a href="#cite_note-61">[61]</a> <div class="mw-heading mw-heading3"><h3 id="Processing_of_pre-mRNA">Processing of pre-mRNA</h3></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/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcriptional modification</a></div> Newly synthesized mRNA molecules are known as <a href="/wiki/Primary_transcript" title="Primary transcript">primary transcripts</a> or pre-mRNA. They must undergo <a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">post-transcriptional modification</a> in the nucleus before being exported to the cytoplasm; mRNA that appears in the cytoplasm without these modifications is degraded rather than used for protein translation. The three main modifications are <a href="/wiki/5%27_cap" class="mw-redirect" title="5' cap">5' capping</a>, 3' <a href="/wiki/Polyadenylation" title="Polyadenylation">polyadenylation</a>, and <a href="/wiki/RNA_splicing" title="RNA splicing">RNA splicing</a>. While in the nucleus, pre-mRNA is associated with a variety of proteins in complexes known as <a href="/wiki/Heterogeneous_ribonucleoprotein_particle" title="Heterogeneous ribonucleoprotein particle">heterogeneous ribonucleoprotein particles</a> (hnRNPs). Addition of the 5' cap occurs co-transcriptionally and is the first step in post-transcriptional modification. The 3' poly-<a href="/wiki/Adenine" title="Adenine">adenine</a> tail is only added after transcription is complete.<a href="#cite_note-Lodish-1">[1]</a>: 509–18  RNA splicing, carried out by a complex called the <a href="/wiki/Spliceosome" title="Spliceosome">spliceosome</a>, is the process by which introns, or regions of DNA that do not code for protein, are removed from the pre-mRNA and the remaining <a href="/wiki/Exon" title="Exon">exons</a> connected to re-form a single continuous molecule. This process normally occurs after 5' capping and 3' polyadenylation but can begin before synthesis is complete in transcripts with many exons.<a href="#cite_note-Lodish-1">[1]</a>: 494  Many pre-mRNAs can be spliced in multiple ways to produce different mature mRNAs that encode different <a href="/wiki/Primary_structure" class="mw-redirect" title="Primary structure">protein sequences</a>. This process is known as <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a>, and allows production of a large variety of proteins from a limited amount of DNA.<a href="#cite_note-Black-62">[62]</a> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading2"><h2 id="Dynamics_and_regulation">Dynamics and regulation</h2></div> <div class="mw-heading mw-heading3"><h3 id="Nuclear_transport">Nuclear transport</h3></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/Nuclear_transport" title="Nuclear transport">Nuclear transport</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:RanGTPcycle.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/RanGTPcycle.png/360px-RanGTPcycle.png" decoding="async" width="360" height="243" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/RanGTPcycle.png/540px-RanGTPcycle.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1e/RanGTPcycle.png/720px-RanGTPcycle.png 2x" data-file-width="2000" data-file-height="1350" /></a><figcaption><a href="/wiki/Macromolecule" title="Macromolecule">Macromolecules</a>, such as <a href="/wiki/RNA" title="RNA">RNA</a> and <a href="/wiki/Protein" title="Protein">proteins</a>, are <a href="/wiki/Active_transport" title="Active transport">actively transported</a> across the nuclear membrane in a process called the <a href="/wiki/Ran_(biology)" class="mw-redirect" title="Ran (biology)">Ran</a>-<a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a> nuclear transport cycle.</figcaption></figure> The entry and exit of large molecules from the nucleus is tightly controlled by the nuclear pore complexes. Although small molecules can enter the nucleus without regulation,<a href="#cite_note-Watson-63">[63]</a> macromolecules such as RNA and proteins require association karyopherins called <a href="/wiki/Importin" title="Importin">importins</a> to enter the nucleus and <a href="/wiki/Exportin" class="mw-redirect" title="Exportin">exportins</a> to exit. "Cargo" proteins that must be translocated from the cytoplasm to the nucleus contain short amino acid sequences known as <a href="/wiki/Nuclear_localization_signal" class="mw-redirect" title="Nuclear localization signal">nuclear localization signals</a>, which are bound by importins, while those transported from the nucleus to the cytoplasm carry <a href="/wiki/Nuclear_export_signal" title="Nuclear export signal">nuclear export signals</a> bound by exportins. The ability of importins and exportins to transport their cargo is regulated by <a href="/wiki/GTPase" title="GTPase">GTPases</a>, enzymes that <a href="/wiki/Hydrolysis" title="Hydrolysis">hydrolyze</a> the molecule guanosine triphosphate (GTP) to release energy. The key GTPase in nuclear transport is <a href="/wiki/Ran_(protein)" title="Ran (protein)">Ran</a>, which is bound to either GTP or GDP (guanosine diphosphate), depending on whether it is located in the nucleus or the cytoplasm. Whereas importins depend on RanGTP to dissociate from their cargo, exportins require RanGTP in order to bind to their cargo.<a href="#cite_note-Pemberton-14">[14]</a> Nuclear import depends on the importin binding its cargo in the cytoplasm and carrying it through the nuclear pore into the nucleus. Inside the nucleus, RanGTP acts to separate the cargo from the importin, allowing the importin to exit the nucleus and be reused. Nuclear export is similar, as the exportin binds the cargo inside the nucleus in a process facilitated by RanGTP, exits through the nuclear pore, and separates from its cargo in the cytoplasm.<a href="#cite_note-Cavazza_2015-64">[64]</a> Specialized export proteins exist for translocation of mature mRNA and tRNA to the cytoplasm after post-transcriptional modification is complete. This quality-control mechanism is important due to these molecules' central role in protein translation. Mis-expression of a protein due to incomplete excision of exons or mis-incorporation of amino acids could have negative consequences for the cell; thus, incompletely modified RNA that reaches the cytoplasm is degraded rather than used in translation.<a href="#cite_note-Lodish-1">[1]</a> <div class="mw-heading mw-heading3"><h3 id="Assembly_and_disassembly">Assembly and disassembly</h3></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Mitosis-fluorescent.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5b/Mitosis-fluorescent.jpg/230px-Mitosis-fluorescent.jpg" decoding="async" width="230" height="202" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/5/5b/Mitosis-fluorescent.jpg 1.5x" data-file-width="300" data-file-height="264" /></a><figcaption>An image of a <a href="/wiki/Newt" title="Newt">newt</a> <a href="/wiki/Lung" title="Lung">lung</a> cell <a href="/wiki/Staining_(biology)" class="mw-redirect" title="Staining (biology)">stained</a> with <a href="/wiki/Fluorescent" class="mw-redirect" title="Fluorescent">fluorescent</a> <a href="/wiki/Dye" title="Dye">dyes</a> during <a href="/wiki/Metaphase" title="Metaphase">metaphase</a>. The <a href="/wiki/Mitotic_spindle" class="mw-redirect" title="Mitotic spindle">mitotic spindle</a> can be seen, stained green, attached to the two sets of <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a>, stained light blue. All chromosomes but one are already at the metaphase plate. </figcaption></figure> During its lifetime, a nucleus may be broken down or destroyed, either in the process of <a href="/wiki/Cell_division" title="Cell division">cell division</a> or as a consequence of <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> (the process of <a href="/wiki/Programmed_cell_death" title="Programmed cell death">programmed cell death</a>). During these events, the structural components of the nucleus — the envelope and lamina — can be systematically degraded. In most cells, the disassembly of the nuclear envelope marks the end of the <a href="/wiki/Prophase" title="Prophase">prophase</a> of mitosis. However, this disassembly of the nucleus is not a universal feature of mitosis and does not occur in all cells. Some unicellular eukaryotes (e.g., yeasts) undergo so-called <a href="/wiki/Closed_mitosis" class="mw-redirect" title="Closed mitosis">closed mitosis</a>, in which the nuclear envelope remains intact. In closed mitosis, the daughter chromosomes migrate to opposite poles of the nucleus, which then divides in two. The cells of higher eukaryotes, however, usually undergo <a href="/wiki/Open_mitosis" class="mw-redirect" title="Open mitosis">open mitosis</a>, which is characterized by breakdown of the nuclear envelope. The daughter chromosomes then migrate to opposite poles of the mitotic spindle, and new nuclei reassemble around them.<a href="#cite_note-Lodish-1">[1]</a>: 854  At a certain point during the cell cycle in open mitosis, the cell divides to form two cells. In order for this process to be possible, each of the new daughter cells must have a full set of genes, a process requiring replication of the chromosomes as well as segregation of the separate sets. This occurs by the replicated chromosomes, the <a href="/wiki/Sister_chromatid" class="mw-redirect" title="Sister chromatid">sister chromatids</a>, attaching to <a href="/wiki/Microtubule" title="Microtubule">microtubules</a>, which in turn are attached to different <a href="/wiki/Centrosome" title="Centrosome">centrosomes</a>. The sister chromatids can then be pulled to separate locations in the cell. In many cells, the centrosome is located in the cytoplasm, outside the nucleus; the microtubules would be unable to attach to the chromatids in the presence of the nuclear envelope.<a href="#cite_note-Lippincott-Schwartz-65">[65]</a> Therefore, the early stages in the cell cycle, beginning in prophase and until around <a href="/wiki/Prometaphase" title="Prometaphase">prometaphase</a>, the nuclear membrane is dismantled.<a href="#cite_note-RGoldman-18">[18]</a> Likewise, during the same period, the nuclear lamina is also disassembled, a process regulated by phosphorylation of the lamins by protein kinases such as the <a href="/wiki/CDC2" class="mw-redirect" title="CDC2">CDC2 protein kinase</a>.<a href="#cite_note-Boulikas-66">[66]</a> Towards the end of the cell cycle, the nuclear membrane is reformed, and around the same time, the nuclear lamina are reassembled by dephosphorylating the lamins.<a href="#cite_note-Boulikas-66">[66]</a> However, in <a href="/wiki/Dinoflagellates" class="mw-redirect" title="Dinoflagellates">dinoflagellates</a>, the nuclear envelope remains intact, the centrosomes are located in the cytoplasm, and the microtubules come in contact with chromosomes, whose centromeric regions are incorporated into the nuclear envelope (the so-called closed mitosis with extranuclear spindle). In many other protists (e.g., <a href="/wiki/Ciliate" title="Ciliate">ciliates</a>, <a href="/wiki/Apicomplexa" title="Apicomplexa">sporozoans</a>) and fungi, the centrosomes are intranuclear, and their nuclear envelope also does not disassemble during cell division.<a href="#cite_note-Boettcher_2013-67">[67]</a> Apoptosis is a controlled process in which the cell's structural components are destroyed, resulting in death of the cell. Changes associated with apoptosis directly affect the nucleus and its contents, for example, in the condensation of chromatin and the disintegration of the nuclear envelope and lamina. The destruction of the lamin networks is controlled by specialized apoptotic <a href="/wiki/Protease" title="Protease">proteases</a> called <a href="/wiki/Caspase" title="Caspase">caspases</a>, which cleave the lamin proteins and, thus, degrade the nucleus' structural integrity. Lamin cleavage is sometimes used as a laboratory indicator of caspase activity in <a href="/wiki/Assay" title="Assay">assays</a> for early apoptotic activity.<a href="#cite_note-RGoldman-18">[18]</a> Cells that express mutant caspase-resistant lamins are deficient in nuclear changes related to apoptosis, suggesting that lamins play a role in initiating the events that lead to apoptotic degradation of the nucleus.<a href="#cite_note-RGoldman-18">[18]</a> Inhibition of lamin assembly itself is an inducer of apoptosis.<a href="#cite_note-Steen-68">[68]</a> The nuclear envelope acts as a barrier that prevents both DNA and RNA viruses from entering the nucleus. Some viruses require access to proteins inside the nucleus in order to replicate and/or assemble. DNA viruses, such as <a href="/wiki/Herpesvirus" class="mw-redirect" title="Herpesvirus">herpesvirus</a> replicate and assemble in the cell nucleus, and exit by budding through the inner nuclear membrane. This process is accompanied by disassembly of the lamina on the nuclear face of the inner membrane.<a href="#cite_note-RGoldman-18">[18]</a> <div class="mw-heading mw-heading3"><h3 id="Disease-related_dynamics">Disease-related dynamics</h3></div> Initially, it has been suspected that <a href="/wiki/Immunoglobulins" class="mw-redirect" title="Immunoglobulins">immunoglobulins</a> in general and <a href="/wiki/Autoantibodies" class="mw-redirect" title="Autoantibodies">autoantibodies</a> in particular do not enter the nucleus. Now there is a body of evidence that under pathological conditions (e.g. <a href="/wiki/Lupus_erythematosus" title="Lupus erythematosus">lupus erythematosus</a>) IgG can enter the nucleus.<a href="#cite_note-pmid17364135-69">[69]</a> <div class="mw-heading mw-heading2"><h2 id="Nuclei_per_cell">Nuclei per cell</h2></div> Most <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> cell types usually have a single nucleus, but some have no nuclei, while others have several. This can result from normal development, as in the maturation of mammalian <a href="/wiki/Red_blood_cell" title="Red blood cell">red blood cells</a>, or from faulty cell division.<a href="#cite_note-70">[70]</a> <div class="mw-heading mw-heading3"><h3 id="Anucleated_cells">Anucleated cells</h3></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Redbloodcells.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/1/13/Redbloodcells.jpg" decoding="async" width="144" height="166" class="mw-file-element" data-file-width="144" data-file-height="166" /></a><figcaption>Human red blood cells, like those of other mammals, lack nuclei. This occurs as a normal part of the cells' development.</figcaption></figure> An anucleated cell contains no nucleus and is, therefore, incapable of dividing to produce daughter cells. The best-known anucleated cell is the mammalian red blood cell, or <a href="/wiki/Erythrocyte" class="mw-redirect" title="Erythrocyte">erythrocyte</a>, which also lacks other organelles such as mitochondria, and serves primarily as a transport vessel to ferry <a href="/wiki/Oxygen" title="Oxygen">oxygen</a> from the <a href="/wiki/Lungs" class="mw-redirect" title="Lungs">lungs</a> to the body's tissues. Erythrocytes mature through <a href="/wiki/Erythropoiesis" title="Erythropoiesis">erythropoiesis</a> in the <a href="/wiki/Bone_marrow" title="Bone marrow">bone marrow</a>, where they lose their nuclei, organelles, and ribosomes. The nucleus is expelled during the process of differentiation from an <a href="/wiki/Erythroblast" class="mw-redirect" title="Erythroblast">erythroblast</a> to a <a href="/wiki/Reticulocyte" title="Reticulocyte">reticulocyte</a>, which is the immediate precursor of the mature erythrocyte.<a href="#cite_note-Skutelsky-71">[71]</a> The presence of <a href="/wiki/Mutagen" title="Mutagen">mutagens</a> may induce the release of some immature "micronucleated" erythrocytes into the bloodstream.<a href="#cite_note-Torous-72">[72]</a><a href="#cite_note-Hutter-73">[73]</a> Anucleated cells can also arise from flawed cell division in which one daughter lacks a nucleus and the other has two nuclei. In <a href="/wiki/Flowering_plant" title="Flowering plant">flowering plants</a>, this condition occurs in <a href="/wiki/Sieve_tube_element" title="Sieve tube element">sieve tube elements</a>.<a href="#cite_note-74">[74]</a> <div class="mw-heading mw-heading3"><h3 id="Multinucleated_cells">Multinucleated cells</h3></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/Multinucleate" title="Multinucleate">Multinucleate</a></div> <a href="/wiki/Multinucleated" class="mw-redirect" title="Multinucleated">Multinucleated</a> cells contain multiple nuclei. Most <a href="/wiki/Acantharea" class="mw-redirect" title="Acantharea">acantharean</a> species of <a href="/wiki/Protozoa" title="Protozoa">protozoa</a><a href="#cite_note-Zettler-75">[75]</a> and some <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> in <a href="/wiki/Mycorrhizae" class="mw-redirect" title="Mycorrhizae">mycorrhizae</a><a href="#cite_note-Horton-76">[76]</a> have naturally multinucleated cells. Other examples include the <a href="/wiki/Intestinal_parasite" class="mw-redirect" title="Intestinal parasite">intestinal parasites</a> in the genus <a href="/wiki/Giardia" title="Giardia">Giardia</a>, which have two nuclei per cell.<a href="#cite_note-77">[77]</a> <a href="/wiki/Ciliate" title="Ciliate">Ciliates</a> have two kinds of nuclei in a single cell, a somatic <a href="/wiki/Macronucleus" title="Macronucleus">macronucleus</a> and a germline <a href="/wiki/Micronucleus" title="Micronucleus">micronucleus</a>.<a href="#cite_note-78">[78]</a> In humans, <a href="/wiki/Skeletal_muscle#Skeletal_muscle_cells" title="Skeletal muscle">skeletal muscle cells</a>, also called <a href="/wiki/Myocyte" class="mw-redirect" title="Myocyte">myocytes</a> and <a href="/wiki/Syncytium" title="Syncytium">syncytium</a>, become multinucleated during development; the resulting arrangement of nuclei near the periphery of the cells allows maximal intracellular space for <a href="/wiki/Myofibrils" class="mw-redirect" title="Myofibrils">myofibrils</a>.<a href="#cite_note-Lodish-1">[1]</a> Other multinucleate cells in the human are <a href="/wiki/Osteoclast" title="Osteoclast">osteoclasts</a> a type of <a href="/wiki/Bone_cell" class="mw-redirect" title="Bone cell">bone cell</a>. Multinucleated and <a href="/wiki/Binucleated_cells" title="Binucleated cells">binucleated cells</a> can also be abnormal in humans; for example, cells arising from the fusion of <a href="/wiki/Monocyte" title="Monocyte">monocytes</a> and <a href="/wiki/Macrophage" title="Macrophage">macrophages</a>, known as <a href="/wiki/Giant_multinucleated_cell" class="mw-redirect" title="Giant multinucleated cell">giant multinucleated cells</a>, sometimes accompany inflammation<a href="#cite_note-McInnes-79">[79]</a> and are also implicated in tumor formation.<a href="#cite_note-Goldring-80">[80]</a> A number of <a href="/wiki/Dinoflagellate" title="Dinoflagellate">dinoflagellates</a> are known to have two nuclei. Unlike other multinucleated cells these nuclei contain two distinct lineages of DNA: one from the dinoflagellate and the other from a symbiotic <a href="/wiki/Diatom" title="Diatom">diatom</a>.<a href="#cite_note-Imanian2012-81">[81]</a> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2></div> As the major defining characteristic of the eukaryotic cell, the nucleus's <a href="/wiki/Evolution" title="Evolution">evolutionary</a> origin has been the subject of much speculation. Four major hypotheses have been proposed to explain the existence of the nucleus, although none have yet earned widespread support.<a href="#cite_note-Pennisi-82">[82]</a><a href="#cite_note-Devos_2014-83">[83]</a><a href="#cite_note-López-García_2015-84">[84]</a> The first model known as the "syntrophic model" proposes that a <a href="/wiki/Symbiosis" title="Symbiosis">symbiotic</a> relationship between the <a href="/wiki/Archaea" title="Archaea">archaea</a> and <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> created the nucleus-containing eukaryotic cell. (Organisms of the Archaeal and Bacterial domains have no cell nucleus.<a href="#cite_note-85">[85]</a>) It is hypothesized that the symbiosis originated when ancient archaea similar to modern <a href="/wiki/Methanogenesis" title="Methanogenesis">methanogenic</a> archaea, invaded and lived within bacteria similar to modern <a href="/wiki/Myxobacteria" title="Myxobacteria">myxobacteria</a>, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a>, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria.<a href="#cite_note-Margulis-86">[86]</a> One possibility is that the nuclear membrane arose as a new membrane system following the origin of <a href="/wiki/Mitochondrion" title="Mitochondrion">mitochondria</a> in an <a href="/wiki/Archaea" title="Archaea">archaebacterial</a> host.<a href="#cite_note-Martin-87">[87]</a> The nuclear membrane may have served to protect the genome from damaging <a href="/wiki/Reactive_oxygen_species" title="Reactive oxygen species">reactive oxygen species</a> produced by the protomitochondria.<a href="#cite_note-88">[88]</a> The archaeal origin of the nucleus is supported by observations that archaea and eukarya have similar genes for certain proteins, including <a href="/wiki/Histone" title="Histone">histones</a>. Observations that myxobacteria are motile, can form multicellular complexes, and possess <a href="/wiki/Kinase" title="Kinase">kinases</a> and <a href="/wiki/G_protein" title="G protein">G proteins</a> similar to eukarya, support a bacterial origin for the eukaryotic cell.<a href="#cite_note-Lopez-Garcia-89">[89]</a> A second model proposes that proto-eukaryotic cells evolved from bacteria without an endosymbiotic stage. This model is based on the existence of modern <a href="/wiki/Planctomycetota" title="Planctomycetota">Planctomycetota</a> bacteria that possess a nuclear structure with primitive pores and other compartmentalized membrane structures.<a href="#cite_note-Fuerst-90">[90]</a> A similar proposal states that a eukaryote-like cell, the <a href="/wiki/Eukaryote#External_phylogeny:_relationship_to_Archaea_and_Bacteria" title="Eukaryote">chronocyte</a>, evolved first and <a href="/wiki/Phagocytosed" class="mw-redirect" title="Phagocytosed">phagocytosed</a> archaea and bacteria to generate the nucleus and the eukaryotic cell.<a href="#cite_note-Hartman-91">[91]</a> The most controversial model, known as <a href="/wiki/Viral_eukaryogenesis" title="Viral eukaryogenesis">viral eukaryogenesis</a>, posits that the membrane-bound nucleus, along with other eukaryotic features, originated from the infection of a prokaryote by a virus. The suggestion is based on similarities between eukaryotes and viruses such as linear DNA strands, mRNA capping, and tight binding to proteins (analogizing histones to <a href="/wiki/Viral_envelope" title="Viral envelope">viral envelopes</a>). One version of the proposal suggests that the nucleus evolved in concert with <a href="/wiki/Phagocytosis" title="Phagocytosis">phagocytosis</a> to form an early cellular "<a href="/wiki/Predator" class="mw-redirect" title="Predator">predator</a>".<a href="#cite_note-Bell-92">[92]</a> Another variant proposes that eukaryotes originated from early archaea infected by <a href="/wiki/Poxvirus" class="mw-redirect" title="Poxvirus">poxviruses</a>, on the basis of observed similarity between the <a href="/wiki/DNA_polymerase" title="DNA polymerase">DNA polymerases</a> in modern poxviruses and eukaryotes.<a href="#cite_note-Takemura-93">[93]</a><a href="#cite_note-Villareal-94">[94]</a> It has been suggested that the unresolved question of the <a href="/wiki/Evolution_of_sex" class="mw-redirect" title="Evolution of sex">evolution of sex</a> could be related to the viral eukaryogenesis hypothesis.<a href="#cite_note-Bell2-95">[95]</a> A more recent proposal, the exomembrane hypothesis, suggests that the nucleus instead originated from a single ancestral cell that evolved a second exterior cell membrane; the interior membrane enclosing the original cell then became the nuclear membrane and evolved increasingly elaborate pore structures for passage of internally synthesized cellular components such as ribosomal subunits.<a href="#cite_note-deRoos-96">[96]</a> <div class="mw-heading mw-heading2"><h2 id="History">History</h2></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Leeuwenhoek1719RedBloodCells.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Leeuwenhoek1719RedBloodCells.jpg/330px-Leeuwenhoek1719RedBloodCells.jpg" decoding="async" width="330" height="112" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Leeuwenhoek1719RedBloodCells.jpg/495px-Leeuwenhoek1719RedBloodCells.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/39/Leeuwenhoek1719RedBloodCells.jpg/660px-Leeuwenhoek1719RedBloodCells.jpg 2x" data-file-width="1440" data-file-height="489" /></a><figcaption>Oldest known depiction of cells and their nuclei by <a href="/wiki/Antonie_van_Leeuwenhoek" title="Antonie van Leeuwenhoek">Antonie van Leeuwenhoek</a>, 1719</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Flemming1882Tafel1Fig14.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Flemming1882Tafel1Fig14.jpg/220px-Flemming1882Tafel1Fig14.jpg" decoding="async" width="220" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Flemming1882Tafel1Fig14.jpg/330px-Flemming1882Tafel1Fig14.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/69/Flemming1882Tafel1Fig14.jpg/440px-Flemming1882Tafel1Fig14.jpg 2x" data-file-width="458" data-file-height="469" /></a><figcaption>Drawing of a <a href="/wiki/Chironomus" title="Chironomus">Chironomus</a> <a href="/wiki/Salivary_gland" title="Salivary gland">salivary gland</a> cell published by <a href="/wiki/Walther_Flemming" title="Walther Flemming">Walther Flemming</a> in 1882. The nucleus contains <a href="/wiki/Polytene_chromosome" title="Polytene chromosome">polytene chromosomes</a>. </figcaption></figure> The nucleus was the first organelle to be discovered. What is most likely the oldest preserved drawing dates back to the early microscopist Antonie van Leeuwenhoek (1632–1723). He observed a "lumen", the nucleus, in the red blood cells of <a href="/wiki/Salmon" title="Salmon">salmon</a>.<a href="#cite_note-97">[97]</a> Unlike mammalian red blood cells, those of other vertebrates still contain nuclei.<a href="#cite_note-98">[98]</a> The nucleus was also described by <a href="/wiki/Franz_Bauer" title="Franz Bauer">Franz Bauer</a> in 1804<a href="#cite_note-Harris-99">[99]</a> and in more detail in 1831 by Scottish <a href="/wiki/Botanist" class="mw-redirect" title="Botanist">botanist</a> <a href="/wiki/Robert_Brown_(Scottish_botanist_from_Montrose)" class="mw-redirect" title="Robert Brown (Scottish botanist from Montrose)">Robert Brown</a> in a talk at the <a href="/wiki/Linnean_Society_of_London" title="Linnean Society of London">Linnean Society of London</a>. Brown was studying <a href="/wiki/Orchid" title="Orchid">orchids</a> under the microscope when he observed an opaque area, which he called the "areola" or "nucleus", in the cells of the flower's outer layer.<a href="#cite_note-Robert_Brown-100">[100]</a> He did not suggest a potential function. In 1838, <a href="/wiki/Matthias_Schleiden" class="mw-redirect" title="Matthias Schleiden">Matthias Schleiden</a> proposed that the nucleus plays a role in generating cells, thus he introduced the name "cytoblast" ("cell builder"). He believed that he had observed new cells assembling around "cytoblasts". <a href="/wiki/Franz_Meyen" title="Franz Meyen">Franz Meyen</a> was a strong opponent of this view, having already described cells multiplying by division and believing that many cells would have no nuclei. The idea that cells can be generated de novo, by the "cytoblast" or otherwise, contradicted work by <a href="/wiki/Robert_Remak" title="Robert Remak">Robert Remak</a> (1852) and <a href="/wiki/Rudolf_Virchow" title="Rudolf Virchow">Rudolf Virchow</a> (1855) who decisively propagated the new paradigm that cells are generated solely by cells ("Omnis cellula e cellula"). The function of the nucleus remained unclear.<a href="#cite_note-Cremer-101">[101]</a> Between 1877 and 1878, <a href="/wiki/Oscar_Hertwig" title="Oscar Hertwig">Oscar Hertwig</a> published several studies on the <a href="/wiki/Fertilization" class="mw-redirect" title="Fertilization">fertilization</a> of <a href="/wiki/Sea_urchin" title="Sea urchin">sea urchin</a> eggs, showing that the nucleus of the <a href="/wiki/Sperm" title="Sperm">sperm</a> enters the <a href="/wiki/Oocyte" title="Oocyte">oocyte</a> and fuses with its nucleus. This was the first time it was suggested that an individual develops from a (single) nucleated cell. This was in contradiction to <a href="/wiki/Ernst_Haeckel" title="Ernst Haeckel">Ernst Haeckel</a>'s theory that the complete <a href="/wiki/Phylogeny" class="mw-redirect" title="Phylogeny">phylogeny</a> of a species would be repeated during embryonic development, including generation of the first nucleated cell from a "monerula", a structureless mass of primordial protoplasm ("<a href="/wiki/Urschleim" class="mw-redirect" title="Urschleim">Urschleim</a>"). Therefore, the necessity of the sperm nucleus for fertilization was discussed for quite some time. However, Hertwig confirmed his observation in other animal groups, including <a href="/wiki/Amphibians" class="mw-redirect" title="Amphibians">amphibians</a> and <a href="/wiki/Molluscs" class="mw-redirect" title="Molluscs">molluscs</a>. <a href="/wiki/Eduard_Strasburger" title="Eduard Strasburger">Eduard Strasburger</a> produced the same results for plants in 1884. This paved the way to assign the nucleus an important role in heredity. In 1873, <a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a> postulated the equivalence of the maternal and paternal germ cells for heredity. The function of the nucleus as carrier of genetic information became clear only later, after mitosis was discovered and the <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian rules</a> were rediscovered at the beginning of the 20th century; the <a href="/wiki/Chromosome_theory_of_heredity" class="mw-redirect" title="Chromosome theory of heredity">chromosome theory of heredity</a> was therefore developed.<a href="#cite_note-Cremer-101">[101]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div> <ul><li><a href="/wiki/Computed_Corpuscle_Sectioning" title="Computed Corpuscle Sectioning">Computed Corpuscle Sectioning</a></li> <li><a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">Nucleus (neuroanatomy)</a></li> <li><a href="/wiki/Nucleoid" title="Nucleoid">Nucleoid</a></li> <li><a href="/wiki/Nucleomorph" title="Nucleomorph">Nucleomorph</a></li></ul> <div class="mw-heading mw-heading2"><h2 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Journal of Molecular Evolution. 53 (3): 251–6. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2001JMolE..53..251L">2001JMolE..53..251L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs002390010215">10.1007/s002390010215</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11523012">11523012</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:20542871">20542871</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Molecular+Evolution&rft.atitle=Viral+eukaryogenesis%3A+was+the+ancestor+of+the+nucleus+a+complex+DNA+virus%3F&rft.volume=53&rft.issue=3&rft.pages=251-6&rft.date=2001-09&rft_id=info%3Adoi%2F10.1007%2Fs002390010215&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A20542871%23id-name%3DS2CID&rft_id=info%3Apmid%2F11523012&rft_id=info%3Abibcode%2F2001JMolE..53..251L&rft.aulast=Bell&rft.aufirst=PJ&rft_id=https%3A%2F%2Fdoi.org%2F10.1007%252Fs002390010215&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-Takemura-93"><a href="#cite_ref-Takemura_93-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTakemura2001" class="citation journal cs1">Takemura M (May 2001). 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Journal of Virology. 74 (15): 7079–84. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1128%2FJVI.74.15.7079-7084.2000">10.1128/JVI.74.15.7079-7084.2000</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC112226">112226</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/10888648">10888648</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Virology&rft.atitle=A+hypothesis+for+DNA+viruses+as+the+origin+of+eukaryotic+replication+proteins&rft.volume=74&rft.issue=15&rft.pages=7079-84&rft.date=2000-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC112226%23id-name%3DPMC&rft_id=info%3Apmid%2F10888648&rft_id=info%3Adoi%2F10.1128%2FJVI.74.15.7079-7084.2000&rft.aulast=Villarreal&rft.aufirst=LP&rft.au=DeFilippis%2C+VR&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC112226&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-Bell2-95"><a href="#cite_ref-Bell2_95-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBell2006" class="citation journal cs1">Bell PJ (November 2006). "Sex and the eukaryotic cell cycle is consistent with a viral ancestry for the eukaryotic nucleus". Primary. Journal of Theoretical Biology. 243 (1): 54–63. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2006JThBi.243...54B">2006JThBi.243...54B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jtbi.2006.05.015">10.1016/j.jtbi.2006.05.015</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16846615">16846615</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Theoretical+Biology&rft.atitle=Sex+and+the+eukaryotic+cell+cycle+is+consistent+with+a+viral+ancestry+for+the+eukaryotic+nucleus&rft.volume=243&rft.issue=1&rft.pages=54-63&rft.date=2006-11&rft_id=info%3Apmid%2F16846615&rft_id=info%3Adoi%2F10.1016%2Fj.jtbi.2006.05.015&rft_id=info%3Abibcode%2F2006JThBi.243...54B&rft.aulast=Bell&rft.aufirst=PJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-deRoos-96"><a href="#cite_ref-deRoos_96-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFde_Roos2006" class="citation journal cs1">de Roos AD (2006). "The origin of the eukaryotic cell based on conservation of existing interfaces". Primary. Artificial Life. 12 (4): 513–23. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1162%2Fartl.2006.12.4.513">10.1162/artl.2006.12.4.513</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16953783">16953783</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:5963228">5963228</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Artificial+Life&rft.atitle=The+origin+of+the+eukaryotic+cell+based+on+conservation+of+existing+interfaces&rft.volume=12&rft.issue=4&rft.pages=513-23&rft.date=2006&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A5963228%23id-name%3DS2CID&rft_id=info%3Apmid%2F16953783&rft_id=info%3Adoi%2F10.1162%2Fartl.2006.12.4.513&rft.aulast=de+Roos&rft.aufirst=AD&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-97"><a href="#cite_ref-97">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVan_Leeuwenhoek" class="citation book cs1 cs1-prop-foreign-lang-source">Van Leeuwenhoek A. Opera Omnia, seu Arcana Naturae ope exactissimorum Microscopiorum detecta, experimentis variis comprobata, Epistolis ad varios illustres viros J. Arnold et Delphis, A. Beman, Lugdinum Batavorum [The Works of, or arcana of nature by means of exactissimorum microscopes had been detected and confirmed by a variety of experiments, the Epistles to the various illustrious men of valor J. Arnold and Delphi, A. Beman, Lugdina York 1719-1730] (in Latin).</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Opera+Omnia%2C+seu+Arcana+Naturae+ope+exactissimorum+Microscopiorum+detecta%2C+experimentis+variis+comprobata%2C+Epistolis+ad+varios+illustres+viros+J.+Arnold+et+Delphis%2C+A.+Beman%2C+Lugdinum+Batavorum&rft.aulast=Van+Leeuwenhoek&rft.aufirst=A&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> Cited in <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGerlach2009" class="citation book cs1">Gerlach D (2009). Geschichte der Mikroskopie. Frankfurt am Main, Germany: <a href="/wiki/Verlag_Harri_Deutsch" title="Verlag Harri Deutsch">Verlag Harri Deutsch</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-8171-1781-9" title="Special:BookSources/978-3-8171-1781-9"><bdi>978-3-8171-1781-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Geschichte+der+Mikroskopie&rft.place=Frankfurt+am+Main%2C+Germany&rft.pub=Verlag+Harri+Deutsch&rft.date=2009&rft.isbn=978-3-8171-1781-9&rft.aulast=Gerlach&rft.aufirst=D&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-98"><a href="#cite_ref-98">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCohen1982" class="citation journal cs1">Cohen WD (1982). "The cytomorphic system of anucleate non-mammalian erythrocytes". Protoplasma. 113: 23–32. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF01283036">10.1007/BF01283036</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:41287948">41287948</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Protoplasma&rft.atitle=The+cytomorphic+system+of+anucleate+non-mammalian+erythrocytes&rft.volume=113&rft.pages=23-32&rft.date=1982&rft_id=info%3Adoi%2F10.1007%2FBF01283036&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A41287948%23id-name%3DS2CID&rft.aulast=Cohen&rft.aufirst=WD&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-Harris-99"><a href="#cite_ref-Harris_99-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarris1999" class="citation book cs1">Harris H (1999). <a rel="nofollow" class="external text" href="https://archive.org/details/birthofcell0000harr">The Birth of the Cell</a>. New Haven: Yale University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-300-07384-3" title="Special:BookSources/978-0-300-07384-3"><bdi>978-0-300-07384-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Birth+of+the+Cell&rft.place=New+Haven&rft.pub=Yale+University+Press&rft.date=1999&rft.isbn=978-0-300-07384-3&rft.aulast=Harris&rft.aufirst=H&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fbirthofcell0000harr&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-Robert_Brown-100"><a href="#cite_ref-Robert_Brown_100-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrown1866" class="citation journal cs1">Brown R (1866). "On the Organs and Mode of Fecundation of Orchidex and Asclepiadea". Miscellaneous Botanical Works I: 511–514.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Miscellaneous+Botanical+Works+I&rft.atitle=On+the+Organs+and+Mode+of+Fecundation+of+Orchidex+and+Asclepiadea&rft.pages=511-514&rft.date=1866&rft.aulast=Brown&rft.aufirst=Robert&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> </li> <li id="cite_note-Cremer-101">^ <a href="#cite_ref-Cremer_101-0">a</a> <a href="#cite_ref-Cremer_101-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCremer1985" class="citation book cs1">Cremer T (1985). Von der Zellenlehre zur Chromosomentheorie. Berlin, Heidelberg, New York, Tokyo: Springer Verlag. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-540-13987-4" title="Special:BookSources/978-3-540-13987-4"><bdi>978-3-540-13987-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Von+der+Zellenlehre+zur+Chromosomentheorie&rft.place=Berlin%2C+Heidelberg%2C+New+York%2C+Tokyo&rft.pub=Springer+Verlag&rft.date=1985&rft.isbn=978-3-540-13987-4&rft.aulast=Cremer&rft.aufirst=Thomas&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> Online Version <a rel="nofollow" class="external text" href="http://www.t-cremer.de/main_de/cremer/personen/info_T_Cremer.htm#book">here</a> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></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 refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGoldmanGruenbaumMoirShumaker2002" class="citation journal cs1">Goldman RD, Gruenbaum Y, Moir RD, Shumaker DK, Spann TP (March 2002). <a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgad.960502">"Nuclear lamins: building blocks of nuclear architecture"</a>. Genes & Development. 16 (5): 533–47. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgad.960502">10.1101/gad.960502</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11877373">11877373</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Genes+%26+Development&rft.atitle=Nuclear+lamins%3A+building+blocks+of+nuclear+architecture&rft.volume=16&rft.issue=5&rft.pages=533-47&rft.date=2002-03&rft_id=info%3Adoi%2F10.1101%2Fgad.960502&rft_id=info%3Apmid%2F11877373&rft.aulast=Goldman&rft.aufirst=RD&rft.au=Gruenbaum%2C+Y&rft.au=Moir%2C+RD&rft.au=Shumaker%2C+DK&rft.au=Spann%2C+TP&rft_id=https%3A%2F%2Fdoi.org%2F10.1101%252Fgad.960502&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> <dl><dd>A review article about nuclear lamins, explaining their structure and various roles</dd></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGörlichKutay1999" class="citation journal cs1">Görlich D, Kutay U (1999). "Transport between the cell nucleus and the cytoplasm". Annual Review of Cell and Developmental Biology. 15: 607–60. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.cellbio.15.1.607">10.1146/annurev.cellbio.15.1.607</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/10611974">10611974</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Cell+and+Developmental+Biology&rft.atitle=Transport+between+the+cell+nucleus+and+the+cytoplasm&rft.volume=15&rft.pages=607-60&rft.date=1999&rft_id=info%3Adoi%2F10.1146%2Fannurev.cellbio.15.1.607&rft_id=info%3Apmid%2F10611974&rft.aulast=G%C3%B6rlich&rft.aufirst=D&rft.au=Kutay%2C+U&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> <dl><dd>A review article about nuclear transport, explains the principles of the mechanism, and the various transport pathways</dd></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLamondEarnshaw1998" class="citation journal cs1">Lamond AI, Earnshaw WC (April 1998). <a rel="nofollow" class="external text" href="http://azolla.fc.ul.pt/aulas/BiologiaCelular/docs/nucleo.pdf">"Structure and function in the nucleus"</a> (PDF). Science. 280 (5363): 547–53. <a href="/wiki/CiteSeerX_(identifier)" class="mw-redirect" title="CiteSeerX (identifier)">CiteSeerX</a> <a rel="nofollow" class="external text" href="https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.323.5543">10.1.1.323.5543</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.280.5363.547">10.1126/science.280.5363.547</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9554838">9554838</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Structure+and+function+in+the+nucleus&rft.volume=280&rft.issue=5363&rft.pages=547-53&rft.date=1998-04&rft_id=https%3A%2F%2Fciteseerx.ist.psu.edu%2Fviewdoc%2Fsummary%3Fdoi%3D10.1.1.323.5543%23id-name%3DCiteSeerX&rft_id=info%3Apmid%2F9554838&rft_id=info%3Adoi%2F10.1126%2Fscience.280.5363.547&rft.aulast=Lamond&rft.aufirst=AI&rft.au=Earnshaw%2C+WC&rft_id=http%3A%2F%2Fazolla.fc.ul.pt%2Faulas%2FBiologiaCelular%2Fdocs%2Fnucleo.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> <dl><dd>A review article about the nucleus, explaining the structure of chromosomes within the organelle, and describing the nucleolus and other subnuclear bodies</dd></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPennisi2004" class="citation journal cs1">Pennisi E (August 2004). "Evolutionary biology. The birth of the nucleus". Science. 305 (5685): 766–8. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.305.5685.766">10.1126/science.305.5685.766</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15297641">15297641</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:83769250">83769250</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Evolutionary+biology.+The+birth+of+the+nucleus&rft.volume=305&rft.issue=5685&rft.pages=766-8&rft.date=2004-08&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83769250%23id-name%3DS2CID&rft_id=info%3Apmid%2F15297641&rft_id=info%3Adoi%2F10.1126%2Fscience.305.5685.766&rft.aulast=Pennisi&rft.aufirst=E&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> <dl><dd>A review article about the evolution of the nucleus, explaining a number of different theories</dd></dl> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPollardEarnshaw2004" class="citation book cs1">Pollard TD, Earnshaw WC (2004). Cell Biology. Philadelphia: Saunders. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7216-3360-2" title="Special:BookSources/978-0-7216-3360-2"><bdi>978-0-7216-3360-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Cell+Biology&rft.place=Philadelphia&rft.pub=Saunders&rft.date=2004&rft.isbn=978-0-7216-3360-2&rft.aulast=Pollard&rft.aufirst=TD&rft.au=Earnshaw%2C+WC&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> <dl><dd>A university level textbook focusing on cell biology. Contains information on nucleus structure and function, including nuclear transport, and subnuclear domains</dd></dl> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <a href="https://commons.wikimedia.org/wiki/Category:Cell_nucleus" class="extiw" title="commons:Category:Cell nucleus">Cell nucleus</a>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239549316"><div class="refbegin" style=""> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><div class="side-box metadata side-box-right"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-abovebelow"> <a href="/wiki/Wikipedia:The_Wikipedia_Library" title="Wikipedia:The Wikipedia Library">Library resources</a> about Cell nucleus <hr /></div> <div class="side-box-flex"> <div class="side-box-text plainlist"><ul><li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Cell+nuclei&library=OLBP">Online books</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Cell+nuclei">Resources in your library</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Cell+nuclei&library=0CHOOSE0">Resources in other libraries</a></li> </ul></div></div> </div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.mechanobio.info/topics/cellular-organization/go-0005634">"The Nucleus"</a>. MBInfo.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=MBInfo&rft.atitle=The+Nucleus&rft_id=http%3A%2F%2Fwww.mechanobio.info%2Ftopics%2Fcellular-organization%2Fgo-0005634&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.cellnucleus.com/education_main.htm">"Learn about the Cell Nucleus"</a>. cellnucleus.com.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=cellnucleus.com&rft.atitle=Learn+about+the+Cell+Nucleus&rft_id=http%3A%2F%2Fwww.cellnucleus.com%2Feducation_main.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> Website covering structure and function of the nucleus from the Department of Oncology at the University of Alberta.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBickmore" class="citation web cs1">Bickmore W. <a rel="nofollow" class="external text" href="http://npd.hgu.mrc.ac.uk/user/?page=compartment">"The Nuclear Protein Database"</a>. Medical Research Council Human Genetics Unit.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=The+Nuclear+Protein+Database&rft.pub=Medical+Research+Council+Human+Genetics+Unit&rft.aulast=Bickmore&rft.aufirst=W&rft_id=http%3A%2F%2Fnpd.hgu.mrc.ac.uk%2Fuser%2F%3Fpage%3Dcompartment&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> Information on nuclear components.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20061112023405/http://cellimages.ascb.org/cdm4/browse.php?CISOROOT=%2Fp4041coll6">"The Nucleus Collection"</a>. Image & Video Library. The American Society for Cell Biology. Archived from <a rel="nofollow" class="external text" href="http://cellimages.ascb.org/cdm4/browse.php?CISOROOT=%2Fp4041coll6">the original</a> on 12 November 2006.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Image+%26+Video+Library&rft.atitle=The+Nucleus+Collection&rft_id=http%3A%2F%2Fcellimages.ascb.org%2Fcdm4%2Fbrowse.php%3FCISOROOT%3D%252Fp4041coll6&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> contains peer-reviewed still images and video clips that illustrate the nucleus.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGallMcIntosh" class="citation web cs1">Gall JG, McIntosh JR (eds.). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20061117175431/http://cellimages.ascb.org/u/?%2Fp4041coll11%2C62">"Nuclear Envelope and Nuclear Import Section"</a>. Landmark Papers in Cell Biology. Archived from <a rel="nofollow" class="external text" href="http://cellimages.ascb.org/u/?%2Fp4041coll11%2C62">the original</a> on 17 November 2006.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Landmark+Papers+in+Cell+Biology&rft.atitle=Nuclear+Envelope+and+Nuclear+Import+Section&rft_id=http%3A%2F%2Fcellimages.ascb.org%2Fu%2F%3F%252Fp4041coll11%252C62&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"> contains digitized commentaries and links to seminal research papers on the nucleus. Published online in the <a rel="nofollow" class="external text" href="http://cellimages.ascb.org/">Image & Video Library</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110610012208/http://cellimages.ascb.org/">Archived</a> 10 June 2011 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> of <a rel="nofollow" class="external text" href="http://www.ascb.org/">The American Society for Cell Biology</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070102032704/http://www.antibodypatterns.com/cytoplasmic.php">"Cytoplasmic patterns generated by human antibodies"</a>. AntibodyPatterns.com. Archived from <a rel="nofollow" class="external text" href="http://www.antibodypatterns.com/cytoplasmic.php">the original</a> on 2 January 2007.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=AntibodyPatterns.com&rft.atitle=Cytoplasmic+patterns+generated+by+human+antibodies&rft_id=http%3A%2F%2Fwww.antibodypatterns.com%2Fcytoplasmic.php&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACell+nucleus" class="Z3988"></li></ul> </div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl 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href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">Endoplasmic reticulum</a></li> <li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a></li> <li><a href="/wiki/Parenthesome" title="Parenthesome">Parenthesome</a></li> <li><a href="/wiki/Autophagosome" title="Autophagosome">Autophagosome</a></li> <li><a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">Vesicle</a> <ul><li><a href="/wiki/Exosome_(vesicle)" title="Exosome (vesicle)">Exosome</a></li> <li><a href="/wiki/Lysosome" title="Lysosome">Lysosome</a></li> <li><a href="/wiki/Endosome" title="Endosome">Endosome</a></li> <li><a href="/wiki/Phagosome" title="Phagosome">Phagosome</a></li> <li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li> <li><a href="/wiki/Acrosome" title="Acrosome">Acrosome</a></li></ul></li> <li><a href="/wiki/Granule_(cell_biology)" title="Granule (cell biology)">Cytoplasmic granule</a> <ul><li><a href="/wiki/Melanosome" title="Melanosome">Melanosome</a></li> <li><a href="/wiki/Microbody" title="Microbody">Microbody</a></li> <li><a href="/wiki/Glyoxysome" title="Glyoxysome">Glyoxysome</a></li> <li><a href="/wiki/Peroxisome" title="Peroxisome">Peroxisome</a></li> <li><a href="/wiki/Weibel%E2%80%93Palade_body" title="Weibel–Palade body">Weibel–Palade body</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cytoskeleton" title="Cytoskeleton">Cytoskeleton</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/Microfilament" title="Microfilament">Microfilament</a></li> <li><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate filament</a></li> <li><a href="/wiki/Microtubule" title="Microtubule">Microtubule</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a></li> <li><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organizing center</a> <ul><li><a href="/wiki/Centrosome" title="Centrosome">Centrosome</a></li> <li><a href="/wiki/Centriole" title="Centriole">Centriole</a></li> <li><a href="/wiki/Basal_body" title="Basal body">Basal body</a></li> <li><a href="/wiki/Spindle_pole_body" title="Spindle pole body">Spindle pole body</a></li></ul></li> <li><a href="/wiki/Myofibril" title="Myofibril">Myofibril</a></li> <li><a href="/wiki/Undulipodium" title="Undulipodium">Undulipodium</a> <ul><li><a href="/wiki/Cilium" title="Cilium">Cilium</a></li> <li><a href="/wiki/Flagellum#Eukaryotic" title="Flagellum">Flagellum</a></li> <li><a href="/wiki/Axoneme" title="Axoneme">Axoneme</a></li> <li><a href="/wiki/Radial_spoke" title="Radial spoke">Radial spoke</a></li></ul></li> <li><a href="/wiki/Pseudopodium" class="mw-redirect" title="Pseudopodium">Pseudopodium</a> <ul><li><a href="/wiki/Lamellipodium" title="Lamellipodium">Lamellipodium</a></li> <li><a href="/wiki/Filopodium" class="mw-redirect" title="Filopodium">Filopodium</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endosymbiont" title="Endosymbiont">Endosymbionts</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/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li> <li><a href="/wiki/Plastid" title="Plastid">Plastid</a> <ul><li><a href="/wiki/Chloroplast" title="Chloroplast">Chloroplast</a></li> <li><a href="/wiki/Chromoplast" title="Chromoplast">Chromoplast</a></li> <li><a href="/wiki/Gerontoplast" title="Gerontoplast">Gerontoplast</a></li> <li><a href="/wiki/Leucoplast" title="Leucoplast">Leucoplast</a></li> <li><a href="/wiki/Amyloplast" title="Amyloplast">Amyloplast</a></li> <li><a href="/wiki/Elaioplast" title="Elaioplast">Elaioplast</a></li> <li><a href="/wiki/Proteinoplast" title="Proteinoplast">Proteinoplast</a></li> <li><a href="/wiki/Tannosome" title="Tannosome">Tannosome</a></li> <li><a href="/wiki/Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li> <li><a href="/wiki/Nitroplast" title="Nitroplast">Nitroplast</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other internal</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/Nucleolus" title="Nucleolus">Nucleolus</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Spliceosome" title="Spliceosome">Spliceosome</a></li> <li><a href="/wiki/Vault_(organelle)" title="Vault (organelle)">Vault</a></li></ul></li> <li><a href="/wiki/Cytoplasm" title="Cytoplasm">Cytoplasm</a> <ul><li><a href="/wiki/Cytosol" title="Cytosol">Cytosol</a></li> <li><a href="/wiki/Cytoplasmic_inclusion" class="mw-redirect" title="Cytoplasmic inclusion">Inclusions</a></li></ul></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</a></li> <li><a href="/wiki/Magnetosome" title="Magnetosome">Magnetosome</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">External</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_wall" title="Cell wall">Cell wall</a></li> <li><a href="/wiki/Extracellular_matrix" title="Extracellular matrix">Extracellular matrix</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="Structures_of_the_cell_nucleus_/_nuclear_protein" 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:Nucleus" title="Template:Nucleus"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nucleus" title="Template talk:Nucleus"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nucleus" title="Special:EditPage/Template:Nucleus"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Structures_of_the_cell_nucleus_/_nuclear_protein" style="font-size:114%;margin:0 4em">Structures of the <a class="mw-selflink selflink">cell nucleus</a> / <a href="/wiki/Nuclear_protein" title="Nuclear protein">nuclear protein</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nuclear_envelope" title="Nuclear envelope">Envelope (membrane)</a>/ <a href="/wiki/Nuclear_lamina" title="Nuclear lamina">nuclear lamina</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/Nuclear_pore" title="Nuclear pore">Pore complex</a>:</li> <li><a href="/wiki/Nucleoporin" title="Nucleoporin">Nucleoporin</a> <ul><li><a href="/wiki/NUP35" class="mw-redirect" title="NUP35">NUP35</a></li> <li><a href="/wiki/NUP37" class="mw-redirect" title="NUP37">NUP37</a></li> <li><a href="/wiki/NUP43" class="mw-redirect" title="NUP43">NUP43</a></li> <li><a href="/wiki/NUP50" class="mw-redirect" title="NUP50">NUP50</a></li> <li><a href="/wiki/NUP54" class="mw-redirect" title="NUP54">NUP54</a></li> <li><a href="/wiki/Nucleoporin_62" class="mw-redirect" title="Nucleoporin 62">NUP62</a></li> <li><a href="/wiki/NUP85" class="mw-redirect" title="NUP85">NUP85</a></li> <li><a href="/wiki/NUP88" class="mw-redirect" title="NUP88">NUP88</a></li> <li><a href="/wiki/NUP93" class="mw-redirect" title="NUP93">NUP93</a></li> <li><a href="/wiki/NUP98" title="NUP98">NUP98</a></li> <li><a href="/wiki/NUP107" class="mw-redirect" title="NUP107">NUP107</a></li> <li><a href="/wiki/NUP133" class="mw-redirect" title="NUP133">NUP133</a></li> <li><a href="/wiki/NUP153" class="mw-redirect" title="NUP153">NUP153</a></li> <li><a href="/wiki/NUP155" class="mw-redirect" title="NUP155">NUP155</a></li> <li><a href="/wiki/NUP160" class="mw-redirect" title="NUP160">NUP160</a></li> <li><a href="/wiki/NUP188" class="mw-redirect" title="NUP188">NUP188</a></li> <li><a href="/wiki/NUP205" class="mw-redirect" title="NUP205">NUP205</a></li> <li><a href="/wiki/Nucleoporin_210kDa" title="Nucleoporin 210kDa">NUP210</a></li> <li><a href="/wiki/NUP214" class="mw-redirect" title="NUP214">NUP214</a></li></ul></li> <li><a href="/wiki/AAAS_(gene)" class="mw-redirect" title="AAAS (gene)">AAAS</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nucleolus" title="Nucleolus">Nucleolus</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/Cajal_body" title="Cajal body">Cajal (coiled) body</a> <ul><li><a href="/wiki/Survival_of_motor_neuron" title="Survival of motor neuron">SMN</a></li> <li><a href="/wiki/Gem-associated_protein_2" title="Gem-associated protein 2">GEMIN2</a></li> <li><a href="/wiki/Gem-associated_protein_4" title="Gem-associated protein 4">GEMIN4</a></li> <li><a href="/wiki/Gem-associated_protein_5" title="Gem-associated protein 5">GEMIN5</a></li> <li><a href="/wiki/Gem-associated_protein_6" title="Gem-associated protein 6">GEMIN6</a></li> <li><a href="/wiki/Gem-associated_protein_7" title="Gem-associated protein 7">GEMIN7</a></li> <li><a href="/wiki/DDX20" title="DDX20">DDX20</a></li> <li><a href="/wiki/Coilin" title="Coilin">COIL</a></li></ul></li> <li><a href="/wiki/Perinucleolar_compartment" title="Perinucleolar compartment">Perinucleolar compartment</a> <ul><li><a href="/wiki/PTBP1" title="PTBP1">PTBP1</a></li> <li><a href="/wiki/CUGBP1" title="CUGBP1">CUGBP1</a></li></ul></li> <li><a href="/wiki/Treacle_protein" title="Treacle protein">TCOF</a></li> <li><a href="/wiki/Ataxin_7" title="Ataxin 7">ATXN7</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd 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-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chromatin" title="Chromatin">Chromatin</a></li> <li><a href="/wiki/Nuclear_dots" class="mw-redirect" title="Nuclear dots">Dot (PML body)</a></li> <li><a href="/wiki/Paraspeckle" title="Paraspeckle">Paraspeckle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/SMC_protein" title="SMC protein">SMC protein</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/Cohesin" title="Cohesin">Cohesin</a> <ul><li><a href="/wiki/SMC1A" title="SMC1A">SMC1A</a></li> <li><a href="/wiki/SMC1B" title="SMC1B">SMC1B</a></li> <li><a href="/wiki/SMC3" title="SMC3">SMC3</a></li></ul></li> <li><a href="/wiki/Condensin" title="Condensin">Condensin</a> <ul><li><a href="/wiki/NCAPD2" title="NCAPD2">NCAPD2</a></li> <li><a href="/wiki/NCAPD3" title="NCAPD3">NCAPD3</a></li> <li><a href="/wiki/NCAPG" title="NCAPG">NCAPG</a></li> <li><a href="/wiki/NCAPG2" title="NCAPG2">NCAPG2</a></li> <li><a href="/wiki/NCAPH" title="NCAPH">NCAPH</a></li> <li><a href="/wiki/NCAPH2" title="NCAPH2">NCAPH2</a></li> <li><a href="/wiki/SMC2" title="SMC2">SMC2</a></li> <li><a href="/wiki/SMC4" title="SMC4">SMC4</a></li></ul></li> <li><a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> <ul><li><a href="/wiki/SMC5" title="SMC5">SMC5</a></li> <li><a href="/wiki/SMC6" title="SMC6">SMC6</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transition_nuclear_protein" title="Transition nuclear protein">Transition nuclear protein</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><ul><li><a href="/wiki/TNP1" title="TNP1">TNP1</a></li> <li><a href="/wiki/TNP2" title="TNP2">TNP2</a></li></ul></li></ul> </div></td></tr><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/Nuclear_matrix" title="Nuclear matrix">Nuclear matrix</a> (Nucleoskeleton)</li> <li><a href="/wiki/Nucleoplasm" title="Nucleoplasm">Nucleoplasm</a> (Nucleosol)</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/LITAF" title="LITAF">LITAF</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>see also <a href="/wiki/Template:Transcription_factors_and_intracellular_receptors" title="Template:Transcription factors and intracellular receptors">transcription factors and intracellular receptors</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div>see also <a href="/wiki/Template:Nucleus_diseases" title="Template:Nucleus diseases">nucleus diseases</a></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" class="navbox authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a>: National <a href="https://www.wikidata.org/wiki/Q40260#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" 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