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data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-Tropomyosin_and_the_actin_skeleton" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Tropomyosin_and_the_actin_skeleton"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>Tropomyosin and the actin skeleton</span> </div> </a> <ul id="toc-Tropomyosin_and_the_actin_skeleton-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Isoforms_and_evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Isoforms_and_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Isoforms and evolution</span> </div> </a> <ul id="toc-Isoforms_and_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genes_and_isoforms_(isoform_complexity)" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Genes_and_isoforms_(isoform_complexity)"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Genes and isoforms (isoform complexity)</span> </div> </a> <ul id="toc-Genes_and_isoforms_(isoform_complexity)-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Splicing" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Splicing"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Splicing</span> </div> </a> <ul id="toc-Splicing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution_of_isoform_generation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution_of_isoform_generation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Evolution of isoform generation</span> </div> </a> <ul id="toc-Evolution_of_isoform_generation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Spatial_sorting_of_tropomyosin_isoforms" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Spatial_sorting_of_tropomyosin_isoforms"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Spatial sorting of tropomyosin isoforms</span> </div> </a> <button aria-controls="toc-Spatial_sorting_of_tropomyosin_isoforms-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Spatial sorting of tropomyosin isoforms subsection</span> </button> <ul id="toc-Spatial_sorting_of_tropomyosin_isoforms-sublist" class="vector-toc-list"> <li id="toc-Regulation_of_sorting" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Regulation_of_sorting"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Regulation of sorting</span> </div> </a> <ul id="toc-Regulation_of_sorting-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanism_of_sorting" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mechanism_of_sorting"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Mechanism of sorting</span> </div> </a> <ul id="toc-Mechanism_of_sorting-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Isoforms_are_not_functionally_redundant" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Isoforms_are_not_functionally_redundant"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Isoforms are not functionally redundant</span> </div> </a> <ul id="toc-Isoforms_are_not_functionally_redundant-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Specific_roles_and_functions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Specific_roles_and_functions"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Specific roles and functions</span> </div> </a> <button aria-controls="toc-Specific_roles_and_functions-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Specific roles and functions subsection</span> </button> <ul id="toc-Specific_roles_and_functions-sublist" class="vector-toc-list"> <li id="toc-Influencing_binding_of_actin_binding_proteins_to_actin_filaments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Influencing_binding_of_actin_binding_proteins_to_actin_filaments"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Influencing binding of actin binding proteins to actin filaments</span> </div> </a> <ul id="toc-Influencing_binding_of_actin_binding_proteins_to_actin_filaments-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Function_in_skeletal_muscle_contraction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Function_in_skeletal_muscle_contraction"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Function in skeletal muscle contraction</span> </div> </a> <ul id="toc-Function_in_skeletal_muscle_contraction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Regulation_of_contraction_in_smooth_muscle" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Regulation_of_contraction_in_smooth_muscle"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.3</span> <span>Regulation of contraction in smooth muscle</span> </div> </a> <ul id="toc-Regulation_of_contraction_in_smooth_muscle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Role_in_cytoskeleton_function" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Role_in_cytoskeleton_function"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.4</span> <span>Role in cytoskeleton function</span> </div> </a> <ul id="toc-Role_in_cytoskeleton_function-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Role_in_diseases" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Role_in_diseases"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Role in diseases</span> </div> </a> <button aria-controls="toc-Role_in_diseases-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Role in diseases subsection</span> </button> <ul id="toc-Role_in_diseases-sublist" class="vector-toc-list"> <li id="toc-Cancer" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cancer"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.1</span> <span>Cancer</span> </div> </a> <ul id="toc-Cancer-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Autoimmunity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Autoimmunity"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.2</span> <span>Autoimmunity</span> </div> </a> <ul id="toc-Autoimmunity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Muscle_diseases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Muscle_diseases"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.3</span> <span>Muscle diseases</span> </div> </a> <ul id="toc-Muscle_diseases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Allergies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Allergies"> <div class="vector-toc-text"> <span class="vector-toc-numb">9.4</span> <span>Allergies</span> </div> </a> <ul id="toc-Allergies-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Tools_and_technologies_to_study_tropomyosins" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Tools_and_technologies_to_study_tropomyosins"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Tools and technologies to study tropomyosins</span> </div> </a> <button aria-controls="toc-Tools_and_technologies_to_study_tropomyosins-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Tools and technologies to study tropomyosins subsection</span> </button> <ul id="toc-Tools_and_technologies_to_study_tropomyosins-sublist" class="vector-toc-list"> <li id="toc-Antibodies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Antibodies"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Antibodies</span> </div> </a> <ul id="toc-Antibodies-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" 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href="https://el.wikipedia.org/wiki/%CE%A4%CF%81%CE%BF%CF%80%CE%BF%CE%BC%CF%85%CE%BF%CF%83%CE%AF%CE%BD%CE%B7" title="Τροπομυοσίνη – Greek" lang="el" hreflang="el" data-title="Τροπομυοσίνη" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Tropomiosina" title="Tropomiosina – Spanish" lang="es" hreflang="es" data-title="Tropomiosina" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Tropomyosine" title="Tropomyosine – French" lang="fr" hreflang="fr" data-title="Tropomyosine" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Tropomiosina" title="Tropomiosina – Galician" lang="gl" hreflang="gl" data-title="Tropomiosina" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Tropomiosin" title="Tropomiosin – Indonesian" lang="id" hreflang="id" data-title="Tropomiosin" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Tropomiosina" title="Tropomiosina – Italian" lang="it" hreflang="it" data-title="Tropomiosina" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%83%88%E3%83%AD%E3%83%9D%E3%83%9F%E3%82%AA%E3%82%B7%E3%83%B3" title="トロポミオシン – Japanese" lang="ja" hreflang="ja" data-title="トロポミオシン" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Tropomiozyna" title="Tropomiozyna – Polish" lang="pl" hreflang="pl" data-title="Tropomiozyna" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Tropomiosina" title="Tropomiosina – Portuguese" lang="pt" hreflang="pt" data-title="Tropomiosina" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A2%D1%80%D0%BE%D0%BF%D0%BE%D0%BC%D0%B8%D0%BE%D0%B7%D0%B8%D0%BD" title="Тропомиозин – Russian" lang="ru" hreflang="ru" data-title="Тропомиозин" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Tropomyoz%C3%ADn" title="Tropomyozín – Slovak" lang="sk" hreflang="sk" data-title="Tropomyozín" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Tropomyosin" title="Tropomyosin – Swedish" lang="sv" hreflang="sv" data-title="Tropomyosin" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A2%D1%80%D0%BE%D0%BF%D0%BE%D0%BC%D1%96%D0%BE%D0%B7%D0%B8%D0%BD" title="Тропоміозин – Ukrainian" lang="uk" hreflang="uk" data-title="Тропоміозин" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Tropomyosin" title="Tropomyosin – Vietnamese" lang="vi" hreflang="vi" data-title="Tropomyosin" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q418454#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> 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<style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">Tropomyosin</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:1C1G_Tropomyosin_crystal.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/1C1G_Tropomyosin_crystal.png/220px-1C1G_Tropomyosin_crystal.png" decoding="async" width="220" height="73" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/1C1G_Tropomyosin_crystal.png/330px-1C1G_Tropomyosin_crystal.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8e/1C1G_Tropomyosin_crystal.png/440px-1C1G_Tropomyosin_crystal.png 2x" data-file-width="750" data-file-height="250" /></a></span><div class="infobox-caption"><span class="plainlinks"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a>: <a rel="nofollow" class="external text" href="https://www.rcsb.org/structure/1C1G">1C1G</a></span></div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3">Symbol</th><td class="infobox-data" style="background-color: #eee">Tropomyosin</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00261">PF00261</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR000533">IPR000533</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/PROSITE" title="PROSITE">PROSITE</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="https://prosite.expasy.org/PDOC00290">PDOC00290</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3"><a href="/wiki/Structural_Classification_of_Proteins" class="mw-redirect" title="Structural Classification of Proteins">SCOP2</a></th><td class="infobox-data" style="background-color: #eee"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=2tma">2tma</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=2tma">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=2tma">SUPFAM</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><table class="infobox mw-collapsible mw-collapsed" style="float:none; clear:none; margin:0; border-width:0; border-collapse:collapse; text-align:left; width:100%"><tbody><tr><th colspan="2" class="infobox-header" style="background-color: #ddd">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Pfam" title="Pfam">Pfam</a>  </th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF00261?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00261">ECOD</a>  </td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.rcsb.org/search?q=rcsb_polymer_entity_annotation.annotation_id:PF00261%20AND%20rcsb_polymer_entity_annotation.type:Pfam">RCSB PDB</a>; <a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/pdbe/entry/search/index?pfam_accession:PF00261">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF00261">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; border:#fafafa 2px solid; border-width:3px 2px 0 0;"><a href="/wiki/PDBsum" title="PDBsum">PDBsum</a></th><td class="infobox-data" style="background-color: #eee; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/thornton-srv/databases/cgi-bin/pdbsum/GetPfamStr.pl?pfam_id=PF00261">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p><b>Tropomyosin</b> is a two-stranded <a href="/wiki/Alpha-helical" class="mw-redirect" title="Alpha-helical">alpha-helical</a>, <a href="/wiki/Coiled_coil" title="Coiled coil">coiled coil</a> <a href="/wiki/Protein" title="Protein">protein</a> found in many <a href="/wiki/Animal" title="Animal">animal</a> and <a href="/wiki/Fungal" class="mw-redirect" title="Fungal">fungal</a> cells. In animals, it is an important component of the muscular system which works in conjunction with <a href="/wiki/Troponin" title="Troponin">troponin</a> to regulate muscle contraction. It is present in smooth and striated muscle tissues, which can be found in various organs and body systems, including the heart, blood vessels, respiratory system, and digestive system. In fungi, tropomyosin is found in cell walls and helps maintain the structural integrity of cells. </p><p>Tropomyosin is found in other <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> too, but not in plants. Overall, tropomyosin is an important protein that plays a vital role in the proper functioning of many different organisms. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Tropomyosin_and_the_actin_skeleton">Tropomyosin and the actin skeleton</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=1" title="Edit section: Tropomyosin and the actin skeleton"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Cardiac_sarcomere_structure.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Cardiac_sarcomere_structure.png/336px-Cardiac_sarcomere_structure.png" decoding="async" width="336" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Cardiac_sarcomere_structure.png/504px-Cardiac_sarcomere_structure.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7f/Cardiac_sarcomere_structure.png/672px-Cardiac_sarcomere_structure.png 2x" data-file-width="2903" data-file-height="1417" /></a><figcaption>Cardiac sarcomere structure featuring tropomyosin </figcaption></figure> <p>All organisms contain organelles that provide physical integrity to their cells. These type of organelles are collectively known as the cytoskeleton, and one of the most ancient systems is based on filamentous polymers of the protein <a href="/wiki/Actin" title="Actin">actin</a>. A polymer of a second protein, tropomyosin, is an integral part of most actin filaments in animals. </p><p>Tropomyosins are a large family of integral components of actin filaments that play a critical role in regulating the function of actin filaments in both muscle and nonmuscle cells. These proteins consist of rod-shaped coiled-coil hetero- or homo-<a href="/wiki/Protein_dimer" title="Protein dimer">dimers</a> that lie along the <a href="/wiki/Alpha_helix" title="Alpha helix">α-helical groove</a> of most actin filaments. Interaction occurs along the length of the actin filament, with dimers aligning in a head-to-tail fashion. </p><p>Tropomyosins are often categorised into two groups: muscle tropomyosin <a href="/wiki/Isoforms" class="mw-redirect" title="Isoforms">isoforms</a> and nonmuscle tropomyosin isoforms. Muscle tropomyosin isoforms are involved in regulating interactions between actin and <a href="/wiki/Myosin" title="Myosin">myosin</a> in the muscle <a href="/wiki/Sarcomere" title="Sarcomere">sarcomere</a> and play a pivotal role in regulated <a href="/wiki/Muscle_contraction" title="Muscle contraction">muscle contraction</a>. Nonmuscle tropomyosin isoforms function in both muscle and nonmuscle cells, and are involved in a range of cellular pathways that control and regulate the cell's <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a> and other key cellular functions. </p><p>The actin filament system that is involved in regulating these cellular pathways is more complex than the actin filament systems that regulates muscle contraction. The contractile system relies upon 4 actin filament isoforms and 5 tropomyosin isoforms,<sup id="cite_ref-Pittenger_1-0" class="reference"><a href="#cite_note-Pittenger-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> whereas the actin filament system of the cytoskeleton uses two actin filament isoforms and over 40 tropomyosin isoforms.<sup id="cite_ref-Pittenger_1-1" class="reference"><a href="#cite_note-Pittenger-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Gunning_2-0" class="reference"><a href="#cite_note-Gunning-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Tropomyosin_bound_to_actin.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Tropomyosin_bound_to_actin.png/120px-Tropomyosin_bound_to_actin.png" decoding="async" width="120" height="86" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/69/Tropomyosin_bound_to_actin.png/180px-Tropomyosin_bound_to_actin.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/69/Tropomyosin_bound_to_actin.png/240px-Tropomyosin_bound_to_actin.png 2x" data-file-width="933" data-file-height="666" /></a></span></div> <div class="gallerytext"></div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Tropomyosin_unbound_to_actin.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Tropomyosin_unbound_to_actin.png/120px-Tropomyosin_unbound_to_actin.png" decoding="async" width="120" height="73" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Tropomyosin_unbound_to_actin.png/180px-Tropomyosin_unbound_to_actin.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/Tropomyosin_unbound_to_actin.png/240px-Tropomyosin_unbound_to_actin.png 2x" data-file-width="933" data-file-height="564" /></a></span></div> <div class="gallerytext"></div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="Isoforms_and_evolution">Isoforms and evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=2" title="Edit section: Isoforms and evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In direct contrast with the 'one <a href="/wiki/Gene" title="Gene">gene</a>, one <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptide</a>' rule, it is now known from a combination of <a href="/wiki/DNA_sequencing" title="DNA sequencing">genomic sequencing</a>, such as the <a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a> and <a href="/wiki/Expressed_sequence_tag" title="Expressed sequence tag">EST</a> data of expressed proteins, that many <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a> produce a range of proteins from a single gene. This plays a crucial role in the functionality of higher eukaryotes, with humans expressing more than 5 times as many different proteins (isoforms) as genes through <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a>. From a mechanistic point of view, it is much easier for an organism to expand on a current gene/protein family (creating protein isoforms) than it is to create an entirely new gene. From an evolutionary point of view, tropomyosins in higher eukaryotes are notable in retaining all 4 of the potential genes produced by the dual genomic duplication event that took place in early eukaryotic evolution. <sup id="cite_ref-gunning_3-0" class="reference"><a href="#cite_note-gunning-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genes_and_isoforms_(isoform_complexity)"><span id="Genes_and_isoforms_.28isoform_complexity.29"></span>Genes and isoforms (isoform complexity)</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=3" title="Edit section: Genes and isoforms (isoform complexity)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1257001546"><table class="infobox" style="clear: right; float:right;margin:0 0 1.5em 1.5em"><tbody><tr><th colspan="2" class="infobox-above" style="font-size:115%">External image</th></tr><tr><td colspan="2" class="infobox-full-data" style="text-align: left"><span typeof="mw:File"><span><img alt="image icon" src="//upload.wikimedia.org/wikipedia/en/thumb/6/61/Searchtool.svg/16px-Searchtool.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/61/Searchtool.svg/24px-Searchtool.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/61/Searchtool.svg/32px-Searchtool.svg.png 2x" data-file-width="512" data-file-height="512" /></span></span> <a href="#CITEREFGunningSchevzovKeeHardeman2005">Gunning et al. 2005</a>, figure 1.<div class="paragraphbreak" style="margin-top:0.5em"></div> Alternative splicing generates multiple products through the use of alternative promoters, resulting in different amino termini, mutually exclusive internal splicing of 6a versus 6b and alternative carboxyl termini. Colour coding is used to indicate that the 1a exon, for example, from the α-Tropomyosin gene is more similar to the 1a exon from the β-Tropomyosin and α3-Tropomyosin genes than it is to the alternative N-terminal 1b exon from the α-Tropomyosin gene. Not all isoforms generated from these genes are shown, although the existence of those shown has been confirmed by northern blots. In most cases, the isoforms arising from alternative splicing do not contain an exon unique to just one isoform. Rather, the isoforms gain their individuality from a unique combination of exons.<sup id="cite_ref-Gunning_2-1" class="reference"><a href="#cite_note-Gunning-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></td></tr></tbody></table> <p>Within mammals, four genes are responsible for generating more than 40 different tropomyosin isoforms. In terms of structure, the genes are very similar, suggesting that they arose through <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a> of an ancestral gene. In humans, these genes are no longer linked and are widely dispersed. In humans, the α1-, β-, α3-, and α4-genes are formally known as <a href="/wiki/TPM1" title="TPM1">TPM1</a>, <a href="/wiki/TPM2" title="TPM2">TPM2</a>, <a href="/wiki/Tropomyosin_3" title="Tropomyosin 3">TPM3</a>, and <a href="/wiki/TPM4" title="TPM4">TPM4</a> and are located at 15q22,<sup id="cite_ref-Eyre_4-0" class="reference"><a href="#cite_note-Eyre-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> 9p13,<sup id="cite_ref-Hunt_5-0" class="reference"><a href="#cite_note-Hunt-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> 1q22<sup id="cite_ref-Wilton_6-0" class="reference"><a href="#cite_note-Wilton-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> and 19p13,<sup id="cite_ref-Wilton1_7-0" class="reference"><a href="#cite_note-Wilton1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> respectively. An alternative nomenclature names the four genes (α,β,γ,δ).<sup id="cite_ref-Gunning_2-2" class="reference"><a href="#cite_note-Gunning-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>Isoforms are defined as highly related gene products that perform, in essence, similar biological functions, with variations existing between the isoforms in terms of biological activity, regulatory properties, temporal and spatial expression, and/or the intercellular location. Isoforms are produced by two distinct mechanisms, gene duplication and alternative splicing. The former mechanism is a process by which multiple copies of a gene are generated through unequal crossing over, through tandem duplication, or by translocation. Alternative splicing is a mechanism wherein exons are either retained in the mRNA or targeted for removal in different combinations to create a diverse array of mRNAs from a single pre-mRNA. </p> <div class="mw-heading mw-heading2"><h2 id="Splicing">Splicing</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=4" title="Edit section: Splicing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A vast array of tropomyosin isoforms are generated by using a combination of different genes and alternative splicing.<sup id="cite_ref-Lees-Miller_8-0" class="reference"><a href="#cite_note-Lees-Miller-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In mammals, regardless of the gene, transcription is initiated at the start of either exon 1a or exon 1b. Depending on the <a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">promoter</a> and initial exon used, tropomyosin isoforms can be categorized as either high-<a href="/wiki/Molecular_weight" class="mw-redirect" title="Molecular weight">molecular-weight</a> (HMW, 284 amino acids) or low-molecular-weight (LMW, 248).<sup id="cite_ref-Pittenger_1-2" class="reference"><a href="#cite_note-Pittenger-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Martin_9-0" class="reference"><a href="#cite_note-Martin-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> HMW isoforms express exon 1a and either 2a or 2b, while LMW isoforms express exon 1b.<sup id="cite_ref-Martin_9-1" class="reference"><a href="#cite_note-Martin-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> To date, all known tropomyosins contain exons 3-9. Alternative splicing can occur at exon 6, with the mutually exclusive choice of exon 6a or 6b.<sup id="cite_ref-Gunning1_10-0" class="reference"><a href="#cite_note-Gunning1-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> At the c-terminus, the transcript is spliced again at exon 9, with the choice of exon 9a, 9b, 9c, or 9d.<sup id="cite_ref-Gunning1_10-1" class="reference"><a href="#cite_note-Gunning1-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_isoform_generation">Evolution of isoform generation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=5" title="Edit section: Evolution of isoform generation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In terms of structure, the genes are very similar, suggesting that they arose through gene duplication of an ancestral gene. The most highly related genes are the α- and γ-genes, utilizing two promoters and differing only with the presence of the unique 2a exon in the α-gene.<sup id="cite_ref-Ruiz-Opazo_11-0" class="reference"><a href="#cite_note-Ruiz-Opazo-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Ruiz-Opazo1_12-0" class="reference"><a href="#cite_note-Ruiz-Opazo1-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Although substantial differences between alternative exons from the same gene have been revealed by sequence comparison (1a and 1b, 6a and 6b, and the exon 9s), most exons are, however, highly conserved between the different genes.<sup id="cite_ref-Pittenger_1-3" class="reference"><a href="#cite_note-Pittenger-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lees-Miller_8-1" class="reference"><a href="#cite_note-Lees-Miller-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Beisel_13-0" class="reference"><a href="#cite_note-Beisel-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lees-Miller1_14-0" class="reference"><a href="#cite_note-Lees-Miller1-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> For example, exon 1a and 1b from the α-gene vary considerably in sequence; however, the sequence from exon 1a from the α-, β-, γ-, and δ-genes is highly conserved. </p><p>Due to the conservative nature of the genes, it is believed that the genes evolved from a common ancestral gene, giving rise to over 40 functionally distinct isoforms. The expression of these isoforms is highly regulated and variable throughout development. The diversity of tropomyosin expression, both in space and in time, provides the potential not only to regulate actin filament function but to create specialised actin filament populations.<sup id="cite_ref-gunning_3-1" class="reference"><a href="#cite_note-gunning-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Spatial_sorting_of_tropomyosin_isoforms">Spatial sorting of tropomyosin isoforms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=6" title="Edit section: Spatial sorting of tropomyosin isoforms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Numerous reports detail that tropomyosin isoforms are sorted to different intracellular locations, often associating with actin filament populations that are involved in specific processes.<sup id="cite_ref-Heimann_15-0" class="reference"><a href="#cite_note-Heimann-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Percival_16-0" class="reference"><a href="#cite_note-Percival-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Percival1_17-0" class="reference"><a href="#cite_note-Percival1-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Lin_18-0" class="reference"><a href="#cite_note-Lin-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Direct visualization of spatial segregation of isoforms was initially observed by Burgoyne and Norman and soon after by Lin and co-workers.<sup id="cite_ref-Lin_18-1" class="reference"><a href="#cite_note-Lin-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Burgoyne_19-0" class="reference"><a href="#cite_note-Burgoyne-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Burgoyne1_20-0" class="reference"><a href="#cite_note-Burgoyne1-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> They observed that specific isoforms were associated with distinct cellular structures.<sup id="cite_ref-Lin_18-2" class="reference"><a href="#cite_note-Lin-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Using specific antibodies, they were able to identify the presence of both HMW and the LMW isoforms of the γ-gene in stress fibers; however, only LMW isoforms were detected in <a href="/wiki/Membrane_ruffling" title="Membrane ruffling">ruffling membranes</a>.<sup id="cite_ref-Lin_18-3" class="reference"><a href="#cite_note-Lin-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>These studies have been extended to a number of cell types with similar results. Extensive studies in neuronal cells,<sup id="cite_ref-Gunning3_21-0" class="reference"><a href="#cite_note-Gunning3-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Fibroblast" title="Fibroblast">fibroblasts</a>,<sup id="cite_ref-Percival_16-1" class="reference"><a href="#cite_note-Percival-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Percival1_17-1" class="reference"><a href="#cite_note-Percival1-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Schevzov_22-0" class="reference"><a href="#cite_note-Schevzov-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> skeletal muscle<sup id="cite_ref-Lin_and_lin_23-0" class="reference"><a href="#cite_note-Lin_and_lin-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kee_24-0" class="reference"><a href="#cite_note-Kee-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Osteoclast" title="Osteoclast">osteoclast</a> cells has further highlighted the complex association tropomyosin isoforms have with cellular structures. These studies have led to the realization that the regulation of isoform sorting is extremely complex and highly regulated. </p> <div class="mw-heading mw-heading3"><h3 id="Regulation_of_sorting">Regulation of sorting</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=7" title="Edit section: Regulation of sorting"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Sorting of tropomyosin isoforms in discrete intracellular locations is developmentally regulated. Initial studies reported that the sorting of isoforms changed through development, where Tropomyosin 4 was initially localized to the <a href="/wiki/Growth_cone" title="Growth cone">growth cone</a> of growing neurons, but in mature neurons it was relocated to the somatodendritic compartment.<sup id="cite_ref-Had_25-0" class="reference"><a href="#cite_note-Had-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> These observations have been supported by studies on different tropomyosin isoforms, showing how tropomyosin populations were relocated during neuron maturation. This evidence is supportive of the notion that tropomyosin isoforms are subject to temporal regulation. </p><p>Additional studies have identified the role the cell cycle plays in isoform sorting. A study that screened a range HMW products from the α- and β-genes and compared localisation with LMW products from the γ-gene found that the HMW and LMW products are mutually exclusively segregated during the early <a href="/wiki/G1_phase" title="G1 phase">G1 phase</a> of the cell cycle.<sup id="cite_ref-Percival1_17-2" class="reference"><a href="#cite_note-Percival1-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mechanism_of_sorting">Mechanism of sorting</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=8" title="Edit section: Mechanism of sorting"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While studies suggest that tropomyosin sorting may be influenced by the sorting of mRNAs,<sup id="cite_ref-Gunning3_21-1" class="reference"><a href="#cite_note-Gunning3-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> there is no absolute correlation between mRNA and protein location. In neurons, Tropomyosin 5NM1 mRNA was found to sort to the pole of the neuron elaborating an axon prior to morphological differentiation.<sup id="cite_ref-Hannan_26-0" class="reference"><a href="#cite_note-Hannan-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The sorting of Tropomyosin 5NM1/2 mRNA to this location correlated with the expression of the Tropomyosin 5NM1/2 protein. In contrast, the mRNA encoding the Tropomyosin Br2 protein was excluded from the pole of the neuron.<sup id="cite_ref-Hannan_26-1" class="reference"><a href="#cite_note-Hannan-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>The link between mRNA sorting and protein location has been tested in transgenic mice models. The models were created so that the coding regions of Tropomyosin 5NM1/2 and Tropomyosin 3 were expressed under the control of the β-actin promoter with the a β-actin 3'-untranslated region lacking targeting information.<sup id="cite_ref-schevzov1_27-0" class="reference"><a href="#cite_note-schevzov1-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The study found that Tropomyosin 3, an isoform that is not normally expressed in neuronal cells, was broadly distributed throughout the neuron, while exogenous expression of the neuronal isoform Tropomyosin 5NM1/2 was found to sort to the growth cone of neurons as does the endogenous Tropomyosin 5NM1/2. As these two transgenes differ only in the tropomyosin coding region yet are localized in two distinct areas, the findings suggest that, in addition to mRNA sorting, the proteins themselves contain sorting information. </p><p>Studies suggest that tropomyosin isoform sorting may also be influenced by the actin isoform composition of microfilaments.<sup id="cite_ref-schevzov1_27-1" class="reference"><a href="#cite_note-schevzov1-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> In myoblasts, overexpression of γ-actin resulted in the down-regulation of β–actin and the removal of Tropomyosin 2 but not Tropomyosin 5 from stress fibers.<sup id="cite_ref-schevzov2_28-0" class="reference"><a href="#cite_note-schevzov2-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> It was later found that, when cells were exposed to cytochalasin D, a chemical that results in the disorganization of actin filaments, tropomyosin isoform sorting was disrupted. Upon the washing out of cytochalasin D, tropomyosin isoform sorting was re-established.<sup id="cite_ref-schevzov3_29-0" class="reference"><a href="#cite_note-schevzov3-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This is suggestive of a strong relationship between the process of tropomyosin isoform sorting and the incorporation of tropomyosin isoforms into organized arrays of actin filaments. There is no evidence for active transport of tropomyosin isoforms to specific locations. Rather, it appears that sorting is the result of local assembly of preferred isoforms at specific intracellular site. The mechanisms that underlie tropomyosin isoform sorting appear to be inherently flexible and dynamic in nature. </p> <div class="mw-heading mw-heading2"><h2 id="Isoforms_are_not_functionally_redundant">Isoforms are not functionally redundant</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=9" title="Edit section: Isoforms are not functionally redundant"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many studies have led to the understanding that tropomyosins perform essential functions and are required in a diverse range of species from yeast, worms, and flies to complex mammals. </p><p>The essential role of tropomyosins was discovered in the Bretscher laboratory, where researchers found that, by eliminating the TPM1 gene of budding yeasts, growth rates were reduced, the presence of actin cables disappeared, defects in vesicular transport were observed, and mating of the yeast was poor.<sup id="cite_ref-Lui_30-0" class="reference"><a href="#cite_note-Lui-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> When a second yeast gene, TPM2, was deleted, no observable changes in the phenotype were recorded; however, when deleted in combination with TPM1, it resulted in lethality. This suggests that TPM1 and -2 genes have overlapping function; however, TPM2 cannot fully compensate of the loss of TPM1, indicating that some functions of TPM1 are unique. Similar results have been observed in flies, worms, amphibians, and mammals, confirming previous results and suggestive of tropomyosin's being involved in a wide range of cellular functions. However, the three co-expressed TMP1, 2, and 4 genes cannot compensate for deletion of the TPM3 gene in embryonic stem cells and preimplantation mouse embryos. </p><p>Results from gene knockout experiments can be ambiguous and must be carefully examined. In studies in which the deletion of a gene leads to lethality, it can at first appear that the gene product had a truly unique role. However, lethality can also be the result of the inability of the compromised cell to express other isoforms to rescue the phenotype because the required isoform is not naturally expressed in the cell. </p> <div class="mw-heading mw-heading2"><h2 id="Specific_roles_and_functions">Specific roles and functions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=10" title="Edit section: Specific roles and functions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Influencing_binding_of_actin_binding_proteins_to_actin_filaments">Influencing binding of actin binding proteins to actin filaments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=11" title="Edit section: Influencing binding of actin binding proteins to actin filaments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Actin_microfilament" class="mw-redirect" title="Actin microfilament">actin microfilament</a> system is the fundamental cytoskeletal system involved in the development and maintenance of cell morphology. The ability of this system to readily respond to cellular cues and undergo structural re-organisation has led to the belief that this system regulates specific structural changes within different cellular regions. </p><p>Within humans, there are only six actin isoforms, and these isoforms are responsible for an array of unique and complex cellular structures and key cellular interactions. It is thought that the function and form of the actin cytoskeleton is controlled largely by <a href="/wiki/Actin-binding_protein" title="Actin-binding protein">actin-binding proteins</a> (ABP) that are associated with the actin polymer. ABP are a group of proteins that bind to actin. Although tropomyosin is sometimes included as an ABP, it is not a true ABP. The tropomyosin dimer has very low affinity for an actin filament and forms no van der waals contacts with actin. It is only the formation of a tropomyosin polymer's winding around the actin filament that provides stability to the tropomyosin-actin filament interaction. </p><p>Many studies suggest that the binding of tropomyosin isoforms to an actin filament may influence the binding of other ABPs, which together alter the structure and convey specific properties and ultimately specific functions to an actin filament. This is demonstrated in neuroepithelial cells, where increased expression of Tropomyosin 5NM1 increases the recruitment of myosin IIB, a myosin motor protein to the growth cone area.<sup id="cite_ref-Bryce_31-0" class="reference"><a href="#cite_note-Bryce-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> However, the over-expression of Tropomyosin Br3 had the opposite effect, decreasing myosin activity in the same region. </p><p>In a pioneering study by Bernstein and Bamburg, it was observed that the actin-binding protein <i>actin depolymerisation factor (ADF)/<a href="/wiki/Cofilin" class="mw-redirect" title="Cofilin">cofilin</a></i>, a factor that promotes actin filament depolymerisation, competed with tropomyosin for binding to the actin filament.<sup id="cite_ref-Bernstein_32-0" class="reference"><a href="#cite_note-Bernstein-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> The expression of Tropomyosin 5NM1 in neuronal cells eliminated ADF/cofilin from the growth cone region, leading to more stable actin filaments.<sup id="cite_ref-Bryce_31-1" class="reference"><a href="#cite_note-Bryce-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> However, the increased expression of Tropomyosin Br3 was observed to recruit ADF/cofilin to actin filaments bound to the Tropomyosin Br3 isoform within the lamellipodium, which led to the disassembly of actin filaments.<sup id="cite_ref-Bryce_31-2" class="reference"><a href="#cite_note-Bryce-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> This phenomenon, whereby a specific tropomyosin isoform directs specific interactions between actin-binding proteins, and the actin filament has been observed in a variety of model systems with a range of different binding proteins (reviewed in Gunning et al., 2008<sup id="cite_ref-Gunning1_10-2" class="reference"><a href="#cite_note-Gunning1-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>). These interactions, under the influence of tropomyosin isoforms, allow actin filaments to be involved in a diverse range of cellular functions. </p> <div class="mw-heading mw-heading3"><h3 id="Function_in_skeletal_muscle_contraction">Function in skeletal muscle contraction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=12" title="Edit section: Function in skeletal muscle contraction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Skeletal_muscle" title="Skeletal muscle">Skeletal muscle</a> is composed of large, multi-nucleated cells (<a href="/wiki/Muscle_fiber" class="mw-redirect" title="Muscle fiber">muscle fibers</a>). Each muscle fiber is packed with longitudinal arrays of <a href="/wiki/Myofibril" title="Myofibril">myofibrils</a>. Myofibrils are composed of repeating protein structures or <a href="/wiki/Sarcomere" title="Sarcomere">sarcomeres</a>, the basic functional unit of skeletal muscle. The sarcomere is a highly structured protein array, consisting of interdigitating thick and thin filaments, where the thin filaments are tethered to a protein structure, the <a href="/wiki/Sarcomere#Bands" title="Sarcomere">Z-line</a>. The dynamic interaction between the thick and thin filaments results in muscle contraction. </p><p>Myosin belongs to a family of motor proteins, and the muscle isoforms of this family comprise the thick filament. The thin filament is made of the skeletal muscle isoforms of actin. Each myosin protein 'paddles' along the thin actin filament, repeatedly binding to myosin-binding sites along the actin filament, ratcheting and letting go. In effect, the thick filament moves or slides along the thin filament, resulting in <a href="/wiki/Muscle_contraction" title="Muscle contraction">muscle contraction</a>. This process is known as the <a href="/wiki/Sliding_filament_model" class="mw-redirect" title="Sliding filament model">sliding filament model</a>. </p><p>The binding of the myosin heads to the muscle actin is a highly regulated process. The thin filament is made of actin, tropomyosin, and troponin. The contraction of skeletal muscle is triggered by nerve impulses that in turn stimulate the release of Ca<sup>2+</sup>. The release of Ca<sup>2+</sup> from the <a href="/wiki/Sarcoplasmic_reticulum" title="Sarcoplasmic reticulum">sarcoplasmic reticulum</a> causes an increase in the concentration of Ca<sup>2+</sup> in the cytosol. Calcium ions then bind to troponin, which is associated with tropomyosin. Binding causes changes in the shape of troponin and subsequently causes the tropomyosin isoform to shift its position on the actin filament. This shifting in position exposes the myosin-binding sites on the actin filament, allowing the myosin heads of the thick filament to bind to the thin filament. </p><p>Structural and biochemical studies suggest that the position of tropomyosin and troponin on the thin filament regulates the interactions between the myosin heads of the thick filament and the binding sites on the actin of the thin filament. <a href="/wiki/X-ray_diffraction" title="X-ray diffraction">X-ray diffraction</a> and <a href="/wiki/Cryoelectron_microscopy" class="mw-redirect" title="Cryoelectron microscopy">cryoelectron microscopy</a> suggest that tropomyosin <a href="/wiki/Steric" class="mw-redirect" title="Steric">sterically</a> blocks the access of myosin to the actin filament. </p><p>Although this model is well-established, it is unclear as to whether the movement of tropomyosin directly causes the myosin head to engage the actin filament. As such, an alternative model has emerged, whereby the movement of the tropomyosin in the filament functions as an <a href="/wiki/Allosteric" class="mw-redirect" title="Allosteric">allosteric</a> switch that is modulated by activating myosin binding but does not function solely by regulating myosin binding. </p> <div class="mw-heading mw-heading3"><h3 id="Regulation_of_contraction_in_smooth_muscle">Regulation of contraction in smooth muscle</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=13" title="Edit section: Regulation of contraction in smooth muscle"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Smooth_muscle" title="Smooth muscle">Smooth muscle</a> is a type of non-striated muscle, and, unlike striated muscle, contraction of smooth muscle is not under conscious control. Smooth muscle may contract spontaneously or rhythmically and be induced by a number of physiochemical agents (hormones, drugs, neurotransmitters). Smooth muscle is found within the walls of various organs and tubes in the body such as the esophagus, stomach, intestines, bronchi, urethra, bladder, and blood vessels. </p><p>Although smooth muscles do not form regular arrays of thick and thin filaments like the sarcomeres of striated muscles, contraction is still due to the same sliding filament mechanism controlled by myosin crossbridges interacting with actin filaments. The thin filament of smooth muscle is made of actin, tropomyosin, <a href="/wiki/Caldesmon" title="Caldesmon">caldesmon</a>, and <a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a>. Within this type of muscle, caldesmon and calmodulin control the tropomyosin-mediated transition between on and off activity states. Caldesmon binds to actin, tropomyosin, calmodulin, and myosin, of which its interactions with actin are most important. The binding of caldesmon is strongly influenced by tropomyosin. Caldesmon is an inhibitor of actinomyosin ATPase and motility, and both actin binding and caldesmon inhibition are greatly enhanced in the presence of tropomyosin. </p><p>Smooth muscle contraction is initiated by the release of Ca<sup>2+</sup>. Ca<sup>2+</sup> binds to and activates calmodulin, which then binds to caldesmon. This binding causes the caldesmon protein to disengage from the actin filament, exposing the myosin-binding sites on the actin filament. Myosin motor heads are phosphorylated by <a href="/wiki/Myosin_light-chain_kinase" title="Myosin light-chain kinase">myosin light-chain kinase</a>, allowing the myosin head to interact with the actin filament and cause contraction. </p> <div class="mw-heading mw-heading3"><h3 id="Role_in_cytoskeleton_function">Role in cytoskeleton function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=14" title="Edit section: Role in cytoskeleton function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The cytoskeleton is an elaborate network of filaments required for the proper functioning of a range of cellular processes including cell motility, cell division, intracellular trafficking, and the maintenance of cell shape. The cytoskeleton is composed of three distinct filament systems: microtubules, intermediate filaments, and microfilaments (also known as the actin cytoskeleton). It is the dynamic interactions between these filaments that provide cells with unique structures and functions. </p><p>A number of regulatory mechanisms, employing many actin-binding proteins, have evolved to control the dynamics of the actin filament system. It is believed that tropomyosins play a pivotal role in this regulatory system, influencing the associations the actin filament has with other ABPs. Together, these associations confer specific properties on the filament, allowing these structures to be involved in a wide range of cellular processes, but also to rapidly respond to cellular stimuli. </p> <div class="mw-heading mw-heading2"><h2 id="Role_in_diseases">Role in diseases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=15" title="Edit section: Role in diseases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Cancer">Cancer</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=16" title="Edit section: Cancer"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many studies have shown that there are specific changes to the repertoire of tropomyosins expressed in cells that are undergoing cellular transformation. These highly reproducible results suggest that, during the process of cellular transformation, a process whereby a normal cell becomes malignant, there is a decreased synthesis of HMW tropomyosin isoforms. In the initial studies, transformation of rat embryo fibroblast cell line REF-52 and of normal rat kidney cells led to decreased synthesis of HMW tropomyosins.<sup id="cite_ref-hendricks1_33-0" class="reference"><a href="#cite_note-hendricks1-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-hendricks2_34-0" class="reference"><a href="#cite_note-hendricks2-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-hLin2_35-0" class="reference"><a href="#cite_note-hLin2-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> In both of these systems, the down regulation was contributed to a decrease in the mRNA levels. These early results suggested that tropomyosins played a critical role in facilitating certain processes that occurred during cell transformation, such as actin filament re-organisation and changes in cell shape. These studies have been reproduced in other laboratories and in other cell lines, with similar results (reviewed in Gunning et al., 2008<sup id="cite_ref-Gunning1_10-3" class="reference"><a href="#cite_note-Gunning1-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>). </p><p>Furthermore, studies have highlighted a link between tropomyosin isoform expression and the acquisition of metastatic properties. A study compared isoform expression between a low- and highly-metastatic Lewis lung carcinoma cell line.<sup id="cite_ref-Takenaga_36-0" class="reference"><a href="#cite_note-Takenaga-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Takenaga1_37-0" class="reference"><a href="#cite_note-Takenaga1-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The study found that as cells become more metastatic, there is a marked decrease in the expression of HMW tropomyosin 2 protein and mRNA levels. </p><p>These results have been confirmed in primary tumors and human models. Studies in colon and bladder cancer found increased expression of the LMW tropomyosin <i>Tropomyosin 5NM1</i>.<sup id="cite_ref-Lin3_38-0" class="reference"><a href="#cite_note-Lin3-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pawlak_39-0" class="reference"><a href="#cite_note-Pawlak-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> The elevated expression of this isoform has also been seen in transformed rat fibroblasts, and it is thought that this isoform is required for the motility of highly metastatic melanoma.<sup id="cite_ref-Miyado_40-0" class="reference"><a href="#cite_note-Miyado-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> In addition, elevated expression of Tropomyosin 4 has been linked with lymph node metastasis in breast cancer. </p><p>All of these studies suggest that changes in the expression and complement of tropomyosin isoforms are integral to cancer and cancer progression. The consensus is that, in general, cancer cells become more reliant on LMW tropomyosins as HMW tropomyosins disappear with increasing malignancy.<sup id="cite_ref-Gunning1_10-4" class="reference"><a href="#cite_note-Gunning1-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> This discovery has led to the development of novel anti-tropomyosin compounds as potential anti-cancer agents. </p> <div class="mw-heading mw-heading3"><h3 id="Autoimmunity">Autoimmunity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=17" title="Edit section: Autoimmunity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Tropomyosins have been implicated in the autoimmune disease <a href="/wiki/Ulcerative_colitis" title="Ulcerative colitis">ulcerative colitis</a>, a disease of the colon that is characterised by ulcers or open sores. The link between this disease and tropomyosin was first acknowledged in a study that found blood serum taken from 95% of patients with ulcerative colitis contained antibodies that reacted positively to tropomyosin.<sup id="cite_ref-Das_41-0" class="reference"><a href="#cite_note-Das-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Additional studies have confirmed these results, but also identify Tropomyosin 5 and Tropomyosin 1 as the primary tropomyosins involved in the pathogenesis of ulcerative colitis.<sup id="cite_ref-Biancone_42-0" class="reference"><a href="#cite_note-Biancone-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Geng_43-0" class="reference"><a href="#cite_note-Geng-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Tropomyosin 5 has been related to the development of pouchitis in the ileal pouch following surgery for ulcerative colitis. The elevated number of IgG-producing cells in the colonic mucosa of ulcerative colitis patients is largely committed to producing IgG against Tropomyosin 5-related epitopes. Tropomyosin 5 is, therefore, capable of inducing a significant T-cell response.<sup id="cite_ref-Biancone2_44-0" class="reference"><a href="#cite_note-Biancone2-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> A physicochemical analysis of common structural motifs present in 109 human autoantigens revealed that tropomyosins have the highest number of such motifs, and thus a very high propensity to act as autoantigens.<sup id="cite_ref-Dohlam_45-0" class="reference"><a href="#cite_note-Dohlam-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>In addition to the role tropomyosins play in ulcerative colitis, tropomyosin antibodies have also been reported in acute <a href="/wiki/Rheumatic_fever" title="Rheumatic fever">rheumatic fever</a><sup id="cite_ref-khanna_46-0" class="reference"><a href="#cite_note-khanna-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> and the inflammatory disorder <a href="/wiki/Behcet%27s_syndrome" class="mw-redirect" title="Behcet's syndrome">Behcet's syndrome</a>.<sup id="cite_ref-Mor_47-0" class="reference"><a href="#cite_note-Mor-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> In both instances, it is unclear as to whether these antibodies play a direct role in the pathogenesis of these human conditions or reflect the high antigenicity of tropomyosins released from compromised cells. </p> <div class="mw-heading mw-heading3"><h3 id="Muscle_diseases">Muscle diseases</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=18" title="Edit section: Muscle diseases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Nemaline myopathy is a muscle disease that is characterised by the presence of electron-dense rod bodies in skeletal muscle fibers. These electron-dense rod bodies are composed mainly of α-actinin and actin. The disorder is often clinically categorized into several groups, including mild (typical), intermediate, severe, and adult-onset; however, these distinctions are somewhat ambiguous, as the categories frequently overlap. Causative mutations have been detected in skeletal α-actinin, tropomyosin, <a href="/wiki/Nebulin" title="Nebulin">nebulin</a>, and troponin. Within humans, mutations in both the γ-Tropomyosin and β-Tropomyosin genes have been identified. No mutations in the α-Tropomyosin gene have been identified in this condition for humans. </p> <div class="mw-heading mw-heading3"><h3 id="Allergies">Allergies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=19" title="Edit section: Allergies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Tropomyosin is the primary protein responsible for <a href="/wiki/Shellfish_allergy" title="Shellfish allergy">shellfish allergies</a>, including those to <a href="/wiki/Crustacean" title="Crustacean">crustaceans</a> and <a href="/wiki/Mollusca" title="Mollusca">molluscs</a>.<sup id="cite_ref-Tong2018_48-0" class="reference"><a href="#cite_note-Tong2018-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Ruethers2018_49-0" class="reference"><a href="#cite_note-Ruethers2018-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Thal2015_50-0" class="reference"><a href="#cite_note-Thal2015-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> </p><p>Tropomyosin also causes some cases of <a href="/wiki/Cockroach" title="Cockroach">cockroach</a> allergy.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Tools_and_technologies_to_study_tropomyosins">Tools and technologies to study tropomyosins</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=20" title="Edit section: Tools and technologies to study tropomyosins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Antibodies">Antibodies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=21" title="Edit section: Antibodies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Given the vast array of processes that this protein has been reported to be involved with, there is great interest in tropomyosin isoforms within the scientific community. </p><p>One way in which specific isoforms of this protein can be studied in detail is through the use of antibodies. These specific antibodies can be used in protein-blotting experiments and applied to cells or tissue sections and observed under a microscope. This allows researchers not only to determine the level or concentration of an isoform or a group of isoforms, but to also identify the cellular location of a particular isoform and associations with other cellular structures or proteins. </p><p>At the present time, there are many commercially available antibodies; however, many of these antibodies are sold with minimal information regarding the antigen used to raise the antibody and therefore, the isoform specificity; as such, some research groups develop their own antibodies. Before these antibodies can be used, they must be extensively characterised, a process whereby the specificity of the antibody is examined to ensure that the antibody does not cross-react with other tropomyosins or other proteins. </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Tropomyosin&action=edit&section=22" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Pittenger-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Pittenger_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Pittenger_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Pittenger_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Pittenger_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFPittengerKazzazHelfman1994" class="citation journal cs1">Pittenger, MF; Kazzaz, JA; Helfman, DM (1994). "Functional properties of non-muscle tropomyosin". <i>Curr Opin Cell Biol</i>. <b>6</b> (1): 96–104. <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%2F0955-0674%2894%2990122-8">10.1016/0955-0674(94)90122-8</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/8167032">8167032</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Curr+Opin+Cell+Biol&rft.atitle=Functional+properties+of+non-muscle+tropomyosin&rft.volume=6&rft.issue=1&rft.pages=96-104&rft.date=1994&rft_id=info%3Adoi%2F10.1016%2F0955-0674%2894%2990122-8&rft_id=info%3Apmid%2F8167032&rft.aulast=Pittenger&rft.aufirst=MF&rft.au=Kazzaz%2C+JA&rft.au=Helfman%2C+DM&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATropomyosin" class="Z3988"></span></span> </li> <li id="cite_note-Gunning-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Gunning_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Gunning_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Gunning_2-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGunningSchevzovKeeHardeman2005" class="citation journal cs1">Gunning, PW; Schevzov, G; Kee, AJ; Hardeman, EC (2005). 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Allergy+and+Clinical+Immunology&rft.atitle=Cockroach+allergens+and+asthma+in+Brazil%3A+identification+of+tropomyosin+as+a+major+allergen+with+potential+cross-reactivity+with+mite+and+shrimp+allergens&rft.volume=104&rft.issue=2&rft.pages=329-337&rft.date=1999&rft_id=info%3Adoi%2F10.1016%2FS0091-6749%2899%2970375-1&rft_id=info%3Apmid%2F10452753&rft.aulast=Santos&rft.aufirst=A.+B.&rft.au=Chapman%2C+M.+D.&rft.au=Aalberse%2C+R.+C.&rft.au=Vailes%2C+L.+D.&rft.au=Ferriani%2C+V.+P.&rft.au=Oliver%2C+C.&rft.au=Rizzo%2C+M.+C.&rft.au=Naspitz%2C+C.+K.&rft.au=Arruda%2C+L.+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ATropomyosin" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a 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class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Skeletal_striated_muscle" class="mw-redirect" title="Skeletal striated muscle">Skeletal<br /> muscle</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Costamere" title="Costamere">Costamere</a>/<br /><a href="/wiki/Dystrophin-associated_protein_complex" title="Dystrophin-associated protein complex">DAPC</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Membrane/<br />extracellular</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="DAP:" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dystrophin-associated_protein" title="Dystrophin-associated protein">DAP</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/Sarcoglycan" title="Sarcoglycan">Sarcoglycan</a> <ul><li><a href="/wiki/SGCA" title="SGCA">SGCA</a></li> <li><a href="/wiki/SGCB" title="SGCB">SGCB</a></li> <li><a href="/wiki/Delta-sarcoglycan" title="Delta-sarcoglycan">SGCD</a></li> <li><a href="/wiki/SGCE" title="SGCE">SGCE</a></li> <li><a href="/wiki/SGCG" title="SGCG">SGCG</a></li> <li><a href="/wiki/SGCZ" title="SGCZ">SGCZ</a></li></ul></li> <li><a href="/wiki/Dystroglycan" title="Dystroglycan">Dystroglycan</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Sarcospan" title="Sarcospan">Sarcospan</a></li> <li><a href="/wiki/Laminin,_alpha_2" class="mw-redirect" title="Laminin, alpha 2">Laminin, alpha 2</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Intracellular</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dystrophin" title="Dystrophin">Dystrophin</a></li> <li><a href="/wiki/Dystrobrevin" title="Dystrobrevin">Dystrobrevin</a> <ul><li><a href="/wiki/DTNA" class="mw-redirect" title="DTNA">A</a></li> <li><a href="/wiki/DTNB" class="mw-redirect" title="DTNB">B</a></li></ul></li> <li><a href="/wiki/Syntrophin" title="Syntrophin">Syntrophin</a> <ul><li><a href="/wiki/Syntrophin,_alpha_1" title="Syntrophin, alpha 1">A</a></li> <li><a href="/wiki/SNTB1" title="SNTB1">B1</a></li> <li><a href="/wiki/SNTB2" title="SNTB2">B2</a></li> <li><a href="/wiki/SNTG1" title="SNTG1">G1</a></li> <li><a href="/wiki/SNTG2" title="SNTG2">G2</a></li></ul></li> <li><a href="/wiki/Syncoilin" title="Syncoilin">Syncoilin</a></li> <li><a href="/wiki/Dysbindin" title="Dysbindin">Dysbindin</a></li> <li><a href="/wiki/Synemin" title="Synemin">Synemin/desmuslin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>related:</i></span></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/NOS1" title="NOS1">NOS1</a></li> <li><a href="/wiki/Caveolin_3" title="Caveolin 3">Caveolin 3</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sarcomere" title="Sarcomere">Sarcomere</a>/<br />(a, i, and h bands;<br />z and m lines)</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/Myofilament" title="Myofilament">Myofilament</a> <ul><li><a href="/wiki/Actin" title="Actin">thin filament/actin</a></li> <li><a href="/wiki/Myosin" title="Myosin">thick filament/myosin</a></li> <li><a href="/wiki/Titin" title="Titin">elastic filament/titin</a></li> <li><a href="/wiki/Nebulin" title="Nebulin">nebulin</a></li></ul></li></ul> <ul><li><a class="mw-selflink selflink">Tropomyosin</a></li></ul> <ul><li><a href="/wiki/Troponin" title="Troponin">Troponin</a> <ul><li><a href="/wiki/Troponin_T" title="Troponin T">T</a></li> <li><a href="/wiki/Troponin_C" title="Troponin C">C</a></li> <li><a href="/wiki/Troponin_I" title="Troponin I">I</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissue</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/Epimysium" title="Epimysium">Epimysium</a></li> <li><a href="/wiki/Muscle_fascicle" title="Muscle fascicle">Fascicle</a></li> <li><a href="/wiki/Perimysium" title="Perimysium">Perimysium</a></li> <li><a href="/wiki/Endomysium" title="Endomysium">Endomysium</a></li> <li><a href="/wiki/Connective_tissue_in_skeletal_muscle" class="mw-redirect" title="Connective tissue in skeletal muscle">Connective tissue in skeletal muscle</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General</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/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></li> <li><a href="/wiki/Motor_unit" title="Motor unit">Motor unit</a></li> <li><a href="/wiki/Muscle_spindle" title="Muscle spindle">Muscle spindle</a></li> <li><a href="/wiki/Excitation%E2%80%93contraction_coupling" class="mw-redirect" title="Excitation–contraction coupling">Excitation–contraction coupling</a></li> <li><a href="/wiki/Sliding_filament_mechanism" class="mw-redirect" title="Sliding filament mechanism">Sliding filament mechanism</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cardiac_muscle" title="Cardiac muscle">Cardiac<br /> muscle</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/Myocardium" class="mw-redirect" title="Myocardium">Myocardium</a></li> <li><a href="/wiki/Intercalated_disc" title="Intercalated disc">Intercalated disc</a></li> <li><a href="/wiki/Nebulette" title="Nebulette">Nebulette</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Both</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Fiber</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/Skeletal_muscle" title="Skeletal muscle">Muscle fiber</a> <ul><li><a href="/wiki/Intrafusal_muscle_fiber" title="Intrafusal muscle fiber">intrafusal</a></li> <li><a href="/wiki/Extrafusal_muscle_fiber" title="Extrafusal muscle fiber">extrafusal</a></li></ul></li> <li><a href="/wiki/Myofibril" title="Myofibril">Myofibril</a></li> <li><a href="/wiki/Microfilament" title="Microfilament">Microfilament</a>/<a href="/wiki/Myofilament" title="Myofilament">Myofilament</a></li> <li><a href="/wiki/Sarcomere" title="Sarcomere">Sarcomere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cells</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/Myoblast" class="mw-redirect" title="Myoblast">Myoblast</a>/<a href="/wiki/Muscle_cell" title="Muscle cell">Muscle cell</a></li> <li><a href="/wiki/Myosatellite_cell" title="Myosatellite cell">Myosatellite cell</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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Desmin" title="Desmin">Desmin</a></li> <li><a href="/wiki/Sarcoplasm" title="Sarcoplasm">Sarcoplasm</a></li> <li><a href="/wiki/Sarcolemma" title="Sarcolemma">Sarcolemma</a> <ul><li><a href="/wiki/T-tubule" title="T-tubule">T-tubule</a></li></ul></li> <li><a href="/wiki/Sarcoplasmic_reticulum" title="Sarcoplasmic reticulum">Sarcoplasmic reticulum</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/<br />ungrouped</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/Myotilin" class="mw-redirect" title="Myotilin">Myotilin</a></li> <li><a href="/wiki/Telethonin" title="Telethonin">Telethonin</a></li> <li><a href="/wiki/Dysferlin" title="Dysferlin">Dysferlin</a></li> <li><a href="/wiki/Fukutin" title="Fukutin">Fukutin</a></li> <li><a href="/wiki/Fukutin-related_protein" title="Fukutin-related protein">Fukutin-related protein</a></li></ul> </div></td></tr></tbody></table><div></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="Proteins_of_the_cytoskeleton" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed 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:Cytoskeletal_proteins" title="Template:Cytoskeletal proteins"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cytoskeletal_proteins" title="Template talk:Cytoskeletal proteins"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cytoskeletal_proteins" title="Special:EditPage/Template:Cytoskeletal proteins"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Proteins_of_the_cytoskeleton" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein" title="Protein">Proteins</a> of the <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeleton</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Human</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microfilament" title="Microfilament">Microfilaments</a><br />and <a href="/wiki/Actin-binding_protein" title="Actin-binding protein">ABPs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Myofilament" title="Myofilament">Myofilament</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Actin" title="Actin">Actins</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/Actin,_alpha_1" class="mw-redirect" title="Actin, alpha 1">A1</a></li> <li><a href="/wiki/ACTA2" title="ACTA2">A2</a></li> <li><a href="/wiki/Beta-actin" title="Beta-actin">B</a></li> <li><a href="/wiki/ACTC1" title="ACTC1">C1</a></li> <li><a href="/wiki/ACTG1" class="mw-redirect" title="ACTG1">G1</a></li> <li><a href="/wiki/ACTG2" title="ACTG2">G2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Myosin" title="Myosin">Myosins</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>I <ul><li><a href="/wiki/MYO1A" class="mw-redirect" title="MYO1A">MYO1A</a></li> <li><a href="/wiki/MYO1B" title="MYO1B">MYO1B</a></li> <li><a href="/wiki/MYO1C" title="MYO1C">MYO1C</a></li> <li><a href="/w/index.php?title=MYO1D&action=edit&redlink=1" class="new" title="MYO1D (page does not exist)">MYO1D</a></li> <li><a href="/wiki/MYO1E" title="MYO1E">MYO1E</a></li> <li><a href="/wiki/MYO1F" title="MYO1F">MYO1F</a></li> <li><a href="/wiki/MYO1G" title="MYO1G">MYO1G</a></li> <li><a href="/w/index.php?title=MYO1H&action=edit&redlink=1" class="new" title="MYO1H (page does not exist)">MYO1H</a>)</li></ul></li> <li>II <ul><li><a href="/wiki/MYH1" title="MYH1">MYH1</a></li> <li><a href="/wiki/MYH2" class="mw-redirect" title="MYH2">MYH2</a></li> <li><a href="/wiki/MYH3" title="MYH3">MYH3</a></li> <li><a href="/wiki/MYH4" title="MYH4">MYH4</a></li> <li><a href="/wiki/MYH6" title="MYH6">MYH6</a></li> <li><a href="/wiki/MYH7" title="MYH7">MYH7</a></li> <li><a href="/wiki/MYH7B" title="MYH7B">MYH7B</a></li> <li><a href="/wiki/MYH8" title="MYH8">MYH8</a></li> <li><a href="/wiki/MYH9" title="MYH9">MYH9</a></li> <li><a href="/wiki/MYH10" title="MYH10">MYH10</a></li> <li><a href="/wiki/MYH11" class="mw-redirect" title="MYH11">MYH11</a></li> <li><a href="/wiki/MYH13" title="MYH13">MYH13</a></li> <li><a href="/wiki/MYH14" title="MYH14">MYH14</a></li> <li><a href="/wiki/MYH15" title="MYH15">MYH15</a></li> <li><a href="/wiki/MYH16" class="mw-redirect" title="MYH16">MYH16</a></li></ul></li> <li>III <ul><li><a href="/wiki/MYO3A" class="mw-redirect" title="MYO3A">MYO3A</a></li> <li><a href="/w/index.php?title=MYO3B&action=edit&redlink=1" class="new" title="MYO3B (page does not exist)">MYO3B</a></li></ul></li> <li>V <ul><li><a href="/wiki/MYO5A" class="mw-redirect" title="MYO5A">MYO5A</a></li> <li><a href="/wiki/MYO5B" title="MYO5B">MYO5B</a></li> <li><a href="/w/index.php?title=MYO5C&action=edit&redlink=1" class="new" title="MYO5C (page does not exist)">MYO5C</a></li></ul></li> <li>VI <ul><li><a href="/wiki/MYO6" class="mw-redirect" title="MYO6">MYO6</a></li></ul></li> <li>VII <ul><li><a href="/wiki/MYO7A" title="MYO7A">MYO7A</a></li> <li><a href="/w/index.php?title=MYO7B&action=edit&redlink=1" class="new" title="MYO7B (page does not exist)">MYO7B</a></li></ul></li> <li>IX <ul><li><a href="/w/index.php?title=MYO9A&action=edit&redlink=1" class="new" title="MYO9A (page does not exist)">MYO9A</a></li> <li><a href="/wiki/MYO9B" title="MYO9B">MYO9B</a></li></ul></li> <li>X <ul><li><a href="/wiki/MYO10" title="MYO10">MYO10</a></li></ul></li> <li>XV <ul><li><a href="/wiki/MYO15A" title="MYO15A">MYO15A</a></li></ul></li> <li>XVIII <ul><li><a href="/wiki/MYO18A" title="MYO18A">MYO18A</a></li> <li><a href="/wiki/MYO18B" title="MYO18B">MYO18B</a></li></ul></li> <li><a href="/wiki/Myosin_light_chain" title="Myosin light chain">LC</a> <ul><li><a href="/wiki/MYL1" title="MYL1">MYL1</a></li> <li><a href="/wiki/MYL2" title="MYL2">MYL2</a></li> <li><a href="/wiki/MYL3" title="MYL3">MYL3</a></li> <li><a href="/wiki/MYL4" title="MYL4">MYL4</a></li> <li><a href="/w/index.php?title=MYL5&action=edit&redlink=1" class="new" title="MYL5 (page does not exist)">MYL5</a></li> <li><a href="/wiki/MYL6" title="MYL6">MYL6</a></li> <li><a href="/wiki/MYL6B" title="MYL6B">MYL6B</a></li> <li><a href="/wiki/MYL7" title="MYL7">MYL7</a></li> <li><a href="/wiki/MYL9" title="MYL9">MYL9</a></li> <li><a href="/wiki/MYLIP" title="MYLIP">MYLIP</a></li> <li><a href="/wiki/MYLK" title="MYLK">MYLK</a></li> <li><a href="/wiki/MYLK2" title="MYLK2">MYLK2</a></li> <li><a href="/w/index.php?title=MYLL1&action=edit&redlink=1" class="new" title="MYLL1 (page does not exist)">MYLL1</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Tropomodulin" title="Tropomodulin">Tropomodulin</a> <ul><li><a href="/wiki/TMOD1" class="mw-redirect" title="TMOD1">1</a></li> <li><a href="/wiki/TMOD2" class="mw-redirect" title="TMOD2">2</a></li> <li><a href="/wiki/TMOD3" class="mw-redirect" title="TMOD3">3</a></li> <li><a href="/wiki/TMOD4" class="mw-redirect" title="TMOD4">4</a></li></ul></li> <li><a href="/wiki/Troponin" title="Troponin">Troponin</a> <ul><li><a href="/wiki/Troponin_T" title="Troponin T">T</a> <a href="/wiki/TNNT1" title="TNNT1">1</a> <a href="/wiki/TNNT2" title="TNNT2">2</a> <a href="/wiki/TNNT3" title="TNNT3">3</a></li> <li><a href="/wiki/Troponin_C" title="Troponin C">C</a> <a href="/wiki/Troponin_C_type_1" title="Troponin C type 1">1</a> <a href="/wiki/TNNC2" class="mw-redirect" title="TNNC2">2</a></li> <li><a href="/wiki/Troponin_I" title="Troponin I">I</a> <a href="/wiki/TNNI1" title="TNNI1">1</a> <a href="/wiki/TNNI2" title="TNNI2">2</a> <a href="/wiki/TNNI3" title="TNNI3">3</a></li></ul></li> <li><a class="mw-selflink selflink">Tropomyosin</a> <ul><li><a href="/wiki/TPM1" title="TPM1">1</a></li> <li><a href="/wiki/TPM2" title="TPM2">2</a></li> <li><a href="/wiki/Tropomyosin_3" title="Tropomyosin 3">3</a></li> <li><a href="/wiki/TPM4" title="TPM4">4</a></li></ul></li></ul> <ul><li><a href="/wiki/Actinin" title="Actinin">Actinin</a> <ul><li><a href="/wiki/Actinin,_alpha_1" class="mw-redirect" title="Actinin, alpha 1">1</a></li> <li><a href="/wiki/Actinin,_alpha_2" class="mw-redirect" title="Actinin, alpha 2">2</a></li> <li><a href="/wiki/ACTN3" class="mw-redirect" title="ACTN3">3</a></li> <li><a href="/wiki/Actinin_alpha_4" class="mw-redirect" title="Actinin alpha 4">4</a></li></ul></li> <li><a href="/wiki/Arp2/3_complex" title="Arp2/3 complex">Arp2/3 complex</a></li> <li><i><a href="/wiki/Actin_depolymerizing_factor" class="mw-redirect" title="Actin depolymerizing factor">actin depolymerizing factors</a></i> <ul><li><a href="/wiki/Cofilin" class="mw-redirect" title="Cofilin">Cofilin</a> <ul><li><a href="/wiki/Cofilin_1" title="Cofilin 1">1</a></li> <li><a href="/wiki/CFL2_(gene)" class="mw-redirect" title="CFL2 (gene)">2</a></li></ul></li> <li><a href="/wiki/Destrin" title="Destrin">Destrin</a></li></ul></li> <li><a href="/wiki/Gelsolin" title="Gelsolin">Gelsolin</a></li> <li><a href="/wiki/Profilin" title="Profilin">Profilin</a> <ul><li><a href="/wiki/Profilin_1" title="Profilin 1">1</a></li> <li><a href="/wiki/PFN2" title="PFN2">2</a></li></ul></li> <li><a href="/wiki/Titin" title="Titin">Titin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</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/Wiskott%E2%80%93Aldrich_syndrome_protein" title="Wiskott–Aldrich syndrome protein">Wiskott–Aldrich syndrome protein</a></li> <li><a href="/wiki/Fibrillin" title="Fibrillin">Fibrillin</a></li> <li><a href="/wiki/Filamin" title="Filamin">Filamin</a> <ul><li><a href="/wiki/FLNA" title="FLNA">FLNA</a></li> <li><a href="/wiki/FLNB" title="FLNB">FLNB</a></li> <li><a href="/wiki/FLNC_(gene)" title="FLNC (gene)">FLNC</a></li></ul></li> <li><a href="/wiki/Espin_(protein)" title="Espin (protein)">Espin</a></li> <li><a href="/wiki/TRIOBP" title="TRIOBP">TRIOBP</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Intermediate_filament" title="Intermediate filament">Intermediate <br />filaments</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Type 1/2<br />(<a href="/wiki/Keratin" title="Keratin">Keratin</a>,<br /><a href="/wiki/Cytokeratin" title="Cytokeratin">Cytokeratin</a>)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Epithelial_keratin" class="mw-redirect" title="Epithelial keratin">Epithelial keratins</a><br />(soft alpha-keratins)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Type_I_keratin" title="Type I keratin">type I</a>/<a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/Keratin_9" title="Keratin 9">9</a></li> <li><a href="/wiki/Keratin_10" title="Keratin 10">10</a></li> <li><a href="/wiki/Keratin_12" title="Keratin 12">12</a></li> <li><a href="/wiki/Keratin_13" title="Keratin 13">13</a></li> <li><a href="/wiki/Keratin_14" title="Keratin 14">14</a></li> <li><a href="/wiki/Keratin_15" title="Keratin 15">15</a></li> <li><a href="/wiki/Keratin_16" title="Keratin 16">16</a></li> <li><a href="/wiki/Keratin_17" title="Keratin 17">17</a></li> <li><a href="/wiki/Keratin_19" title="Keratin 19">19</a></li> <li><a href="/wiki/Keratin_20" title="Keratin 20">20</a></li></ul></li> <li><i><a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/Keratin_18" title="Keratin 18">18</a></li></ul></li> <li><i>none</i> <ul><li><a href="/wiki/Keratin_21" title="Keratin 21">21</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Type_II_keratin" title="Type II keratin">type II</a>/<a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/Keratin_1" title="Keratin 1">1</a></li> <li><a href="/wiki/Keratin_2A" title="Keratin 2A">2A</a></li> <li><a href="/wiki/Keratin_3" title="Keratin 3">3</a></li> <li><a href="/wiki/Keratin_4" title="Keratin 4">4</a></li> <li><a href="/wiki/Keratin_5" title="Keratin 5">5</a></li> <li><a href="/wiki/Keratin_6A" title="Keratin 6A">6A</a></li> <li><a href="/wiki/Keratin_6B" title="Keratin 6B">6B</a></li> <li><a href="/wiki/Keratin_6C" title="Keratin 6C">6C</a></li> <li><a href="/wiki/Keratin_7" title="Keratin 7">7</a></li> <li><a href="/wiki/Keratin_8" title="Keratin 8">8</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hair_keratin" title="Hair keratin">Hair keratins</a><br />(hard alpha-keratins)</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Type_I_hair_keratin" class="mw-redirect" title="Type I hair keratin">type I</a>/<a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/KRT31" title="KRT31">31</a></li> <li><a href="/wiki/KRT32" title="KRT32">32</a></li> <li><a href="/wiki/KRT33A" title="KRT33A">33A</a></li> <li><a href="/wiki/KRT33B" title="KRT33B">33B</a></li> <li><a href="/wiki/Keratin_34" title="Keratin 34">34</a></li> <li><a href="/wiki/KRT35" title="KRT35">35</a></li> <li><a href="/wiki/KRT36" title="KRT36">36</a></li> <li><a href="/wiki/KRT37" title="KRT37">37</a></li> <li><a href="/wiki/KRT38" title="KRT38">38</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Type_II_hair_keratin" class="mw-redirect" title="Type II hair keratin">type II</a>/<a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/KRT81" title="KRT81">81</a></li> <li><a href="/wiki/KRT82" title="KRT82">82</a></li> <li><a href="/wiki/KRT83" title="KRT83">83</a></li> <li><a href="/wiki/KRT84" title="KRT84">84</a></li> <li><a href="/wiki/KRT85" title="KRT85">85</a></li> <li><a href="/wiki/KRT86" title="KRT86">86</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ungrouped alpha</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Chromosome_17" title="Chromosome 17">chromosome 17</a></i> <ul><li><a href="/wiki/KRT23" title="KRT23">23</a></li> <li><a href="/wiki/KRT24" title="KRT24">24</a></li> <li><a href="/wiki/KRT25" title="KRT25">25</a></li> <li><a href="/wiki/KRT26" title="KRT26">26</a></li> <li><a href="/wiki/KRT27" title="KRT27">27</a></li> <li><a href="/wiki/KRT28" title="KRT28">28</a></li> <li><a href="/wiki/KRT39" title="KRT39">39</a></li> <li><a href="/wiki/KRT40" title="KRT40">40</a></li></ul></li></ul> <ul><li><i><a href="/wiki/Chromosome_12" title="Chromosome 12">chromosome 12</a></i> <ul><li><a href="/wiki/KRT71" title="KRT71">71</a></li> <li><a href="/wiki/KRT72" title="KRT72">72</a></li> <li><a href="/wiki/KRT73" title="KRT73">73</a></li> <li><a href="/wiki/KRT74" title="KRT74">74</a></li> <li><a href="/wiki/KRT75" title="KRT75">75</a></li> <li><a href="/wiki/KRT76" title="KRT76">76</a></li> <li><a href="/wiki/KRT77" title="KRT77">77</a></li> <li><a href="/wiki/KRT78" title="KRT78">78</a></li> <li><a href="/wiki/KRT79" title="KRT79">79</a></li> <li><a href="/wiki/KRT80" title="KRT80">80</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Not alpha</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/Beta-keratin" title="Beta-keratin">Beta-keratin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 3</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/Desmin" title="Desmin">Desmin</a></li> <li><a href="/wiki/Glial_fibrillary_acidic_protein" title="Glial fibrillary acidic protein">GFAP</a></li> <li><a href="/wiki/Peripherin" title="Peripherin">Peripherin</a></li> <li><a href="/wiki/Vimentin" title="Vimentin">Vimentin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 4</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/Internexin" title="Internexin">Internexin</a></li> <li><a href="/wiki/Nestin_(protein)" title="Nestin (protein)">Nestin</a></li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofilament</a> <ul><li><a href="/wiki/NEFL" class="mw-redirect" title="NEFL">NEFL</a></li> <li><a href="/wiki/NEFM" title="NEFM">NEFM</a></li> <li><a href="/wiki/NEFH" title="NEFH">NEFH</a></li></ul></li> <li><a href="/wiki/Synemin" title="Synemin">Synemin</a></li> <li><a href="/wiki/Syncoilin" title="Syncoilin">Syncoilin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Type 5</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/Lamin" title="Lamin">Nuclear lamins</a>: <a href="/wiki/Lamin_A/C" class="mw-redirect" title="Lamin A/C">A/C</a></li> <li><a href="/wiki/Lamin_B1" title="Lamin B1">B1</a></li> <li><a href="/wiki/Lamin_B2" title="Lamin B2">B2</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microtubule" title="Microtubule">Microtubules</a><br />and <a href="/wiki/Microtubule-associated_protein" title="Microtubule-associated protein">MAPs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tubulin" title="Tubulin">Tubulins</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/TUBA1A" class="mw-redirect" title="TUBA1A">TUBA1A</a></li> <li><a href="/wiki/TUBA1B" title="TUBA1B">TUBA1B</a></li> <li><a href="/wiki/TUBA1C" title="TUBA1C">TUBA1C</a></li> <li><a href="/wiki/TUBA3C" class="mw-redirect" title="TUBA3C">TUBA3C</a></li> <li><a href="/w/index.php?title=TUBA3D&action=edit&redlink=1" class="new" title="TUBA3D (page does not exist)">TUBA3D</a></li> <li><a href="/w/index.php?title=TUBA3E&action=edit&redlink=1" class="new" title="TUBA3E (page does not exist)">TUBA3E</a></li> <li><a href="/wiki/TUBA4A" title="TUBA4A">TUBA4A</a></li> <li><a href="/wiki/TUBA8" title="TUBA8">TUBA8</a></li> <li><a href="/wiki/TUBB" title="TUBB">TUBB</a></li> <li><a href="/wiki/TUBB1" title="TUBB1">TUBB1</a></li> <li><a href="/wiki/TUBB2A" class="mw-redirect" title="TUBB2A">TUBB2A</a></li> <li><a href="/w/index.php?title=TUBB2B&action=edit&redlink=1" class="new" title="TUBB2B (page does not exist)">TUBB2B</a></li> <li><a href="/wiki/TUBB2C" class="mw-redirect" title="TUBB2C">TUBB2C</a></li> <li><a href="/wiki/TUBB3" class="mw-redirect" title="TUBB3">TUBB3</a></li> <li><a href="/wiki/TUBB4" class="mw-redirect" title="TUBB4">TUBB4</a></li> <li><a href="/w/index.php?title=TUBB4Q&action=edit&redlink=1" class="new" title="TUBB4Q (page does not exist)">TUBB4Q</a></li> <li><a href="/w/index.php?title=TUBB6&action=edit&redlink=1" class="new" title="TUBB6 (page does not exist)">TUBB6</a></li> <li><a href="/w/index.php?title=TUBB8&action=edit&redlink=1" class="new" title="TUBB8 (page does not exist)">TUBB8</a></li> <li><a href="/wiki/TUBG1" title="TUBG1">TUBG1</a></li> <li><a href="/wiki/TUBG2" title="TUBG2">TUBG2</a></li> <li><a href="/wiki/TUBGCP2" title="TUBGCP2">TUBGCP2</a></li> <li><a href="/wiki/TUBGCP3" title="TUBGCP3">TUBGCP3</a></li> <li><a href="/wiki/TUBGCP4" title="TUBGCP4">TUBGCP4</a></li> <li><a href="/wiki/TUBGCP5" title="TUBGCP5">TUBGCP5</a></li> <li><a href="/wiki/TUBGCP6" title="TUBGCP6">TUBGCP6</a></li> <li><a href="/wiki/TUBD1" title="TUBD1">TUBD1</a></li> <li><a href="/wiki/TUBE1" title="TUBE1">TUBE1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microtubule-associated_protein" title="Microtubule-associated protein">MAPs</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/MAPRE1" title="MAPRE1">EB1</a></li> <li><a href="/wiki/MAPRE2" title="MAPRE2">EB2</a></li> <li><a href="/wiki/MAPRE3" title="MAPRE3">EB3</a></li> <li><a href="/wiki/MAP1A" title="MAP1A">MAP1A</a></li> <li><a href="/wiki/MAP1B" title="MAP1B">MAP1B</a></li> <li><a href="/wiki/MAP2" class="mw-redirect" title="MAP2">MAP2</a></li> <li><a href="/wiki/MAP4" title="MAP4">MAP4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Kinesin" title="Kinesin">Kinesins</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/KIF1A" title="KIF1A">KIF1A</a></li> <li><a href="/wiki/KIF1B" title="KIF1B">KIF1B</a></li> <li><a href="/wiki/KIF2A" title="KIF2A">KIF2A</a></li> <li><a href="/wiki/KIF2C" title="KIF2C">KIF2C</a></li> <li><a href="/wiki/KIF3B" title="KIF3B">KIF3B</a></li> <li><a href="/wiki/KIF3C" title="KIF3C">KIF3C</a></li> <li><a href="/wiki/KIF4A" title="KIF4A">KIF4A</a></li> <li><a href="/wiki/KIF4B" title="KIF4B">KIF4B</a></li> <li><a href="/wiki/KIF5A" title="KIF5A">KIF5A</a></li> <li><a href="/wiki/KIF5B" title="KIF5B">KIF5B</a></li> <li><a href="/wiki/KIF5C" title="KIF5C">KIF5C</a></li> <li><a href="/wiki/KIF6" title="KIF6">KIF6</a></li> <li><a href="/wiki/KIF7" title="KIF7">KIF7</a></li> <li><a href="/wiki/KIF9" title="KIF9">KIF9</a></li> <li><a href="/wiki/KIF11" class="mw-redirect" title="KIF11">KIF11</a></li> <li><a href="/wiki/KIF12" title="KIF12">KIF12</a></li> <li><a href="/wiki/KIF13A" title="KIF13A">KIF13A</a></li> <li><a href="/wiki/KIF13B" title="KIF13B">KIF13B</a></li> <li><a href="/wiki/KIF14" title="KIF14">KIF14</a></li> <li><a href="/wiki/KIF15" title="KIF15">KIF15</a></li> <li><a href="/wiki/KIF16B" title="KIF16B">KIF16B</a></li> <li><a href="/wiki/KIF17" title="KIF17">KIF17</a></li> <li><a href="/wiki/KIF18A" title="KIF18A">KIF18A</a></li> <li><a href="/wiki/KIF18B" title="KIF18B">KIF18B</a></li> <li><a href="/wiki/KIF19" title="KIF19">KIF19</a></li> <li><a href="/wiki/KIF20A" title="KIF20A">KIF20A</a></li> <li><a href="/wiki/KIF20B" title="KIF20B">KIF20B</a></li> <li><a href="/wiki/KIF21A" title="KIF21A">KIF21A</a></li> <li><a href="/wiki/KIF21B" title="KIF21B">KIF21B</a></li> <li><a href="/wiki/KIF22" title="KIF22">KIF22</a></li> <li><a href="/wiki/KIF23" title="KIF23">KIF23</a></li> <li><a href="/wiki/KIF24" title="KIF24">KIF24</a></li> <li><a href="/wiki/KIF25" title="KIF25">KIF25</a></li> <li><a href="/wiki/KIF26A" title="KIF26A">KIF26A</a></li> <li><a href="/w/index.php?title=KIF26B&action=edit&redlink=1" class="new" title="KIF26B (page does not exist)">KIF26B</a></li> <li><a href="/wiki/KIF27" title="KIF27">KIF27</a></li> <li><a href="/wiki/KIFC1" title="KIFC1">KIFC1</a></li> <li><a href="/w/index.php?title=KIFC2&action=edit&redlink=1" class="new" title="KIFC2 (page does not exist)">KIFC2</a></li> <li><a href="/wiki/KIFC3" title="KIFC3">KIFC3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dynein" title="Dynein">Dyneins</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>axonemal: <a href="/wiki/DNAH1" title="DNAH1">DNAH1</a></li> <li><a href="/w/index.php?title=DNAH2&action=edit&redlink=1" class="new" title="DNAH2 (page does not exist)">DNAH2</a></li> <li><a href="/w/index.php?title=DNAH3&action=edit&redlink=1" class="new" title="DNAH3 (page does not exist)">DNAH3</a></li> <li><a href="/wiki/DNAH5" title="DNAH5">DNAH5</a></li> <li><a href="/w/index.php?title=DNAH6&action=edit&redlink=1" class="new" title="DNAH6 (page does not exist)">DNAH6</a></li> <li><a href="/wiki/DNAH7" title="DNAH7">DNAH7</a></li> <li><a href="/w/index.php?title=DNAH8&action=edit&redlink=1" class="new" title="DNAH8 (page does not exist)">DNAH8</a></li> <li><a href="/wiki/DNAH9" title="DNAH9">DNAH9</a></li> <li><a href="/w/index.php?title=DNAH10&action=edit&redlink=1" class="new" title="DNAH10 (page does not exist)">DNAH10</a></li> <li><a href="/wiki/DNAH11" title="DNAH11">DNAH11</a></li> <li><a href="/w/index.php?title=DNAH12&action=edit&redlink=1" class="new" title="DNAH12 (page does not exist)">DNAH12</a></li> <li><a href="/w/index.php?title=DNAH13&action=edit&redlink=1" class="new" title="DNAH13 (page does not exist)">DNAH13</a></li> <li><a href="/w/index.php?title=DNAH14&action=edit&redlink=1" class="new" title="DNAH14 (page does not exist)">DNAH14</a></li> <li><a href="/w/index.php?title=DNAH17&action=edit&redlink=1" class="new" title="DNAH17 (page does not exist)">DNAH17</a></li> <li><a href="/wiki/DNAI1" title="DNAI1">DNAI1</a></li> <li><a href="/wiki/DNAI2" title="DNAI2">DNAI2</a></li> <li><a href="/w/index.php?title=DNALI1&action=edit&redlink=1" class="new" title="DNALI1 (page does not exist)">DNALI1</a></li> <li><a href="/wiki/DNAL1" class="mw-redirect" title="DNAL1">DNAL1</a></li> <li><a href="/wiki/DNAL4" title="DNAL4">DNAL4</a></li></ul> <ul><li>cytoplasmic: <a href="/wiki/DYNC1H1" title="DYNC1H1">DYNC1H1</a></li> <li><a href="/wiki/DYNC2H1" title="DYNC2H1">DYNC2H1</a></li> <li><a href="/wiki/DYNC1I1" title="DYNC1I1">DYNC1I1</a></li> <li><a href="/wiki/DYNC1I2" title="DYNC1I2">DYNC1I2</a></li> <li><a href="/wiki/DYNC1LI1" title="DYNC1LI1">DYNC1LI1</a></li> <li><a href="/wiki/DYNC1LI2" title="DYNC1LI2">DYNC1LI2</a></li> <li><a href="/w/index.php?title=DYNC2LI1&action=edit&redlink=1" class="new" title="DYNC2LI1 (page does not exist)">DYNC2LI1</a></li> <li><a href="/wiki/DYNLL1" title="DYNLL1">DYNLL1</a></li> <li><a href="/wiki/DYNLL2" title="DYNLL2">DYNLL2</a></li> <li><a href="/wiki/DYNLRB1" title="DYNLRB1">DYNLRB1</a></li> <li><a href="/w/index.php?title=DYNLRB2&action=edit&redlink=1" class="new" title="DYNLRB2 (page does not exist)">DYNLRB2</a></li> <li><a href="/wiki/DYNLT1" title="DYNLT1">DYNLT1</a></li> <li><a href="/wiki/DYNLT3" title="DYNLT3">DYNLT3</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Microtubule_organizing_center" title="Microtubule organizing center">Microtubule organising proteins</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/CAMSAP1" title="CAMSAP1">CAMSAP1</a></li> <li><a href="/wiki/CAMSAP2" title="CAMSAP2">CAMSAP2</a></li> <li><a href="/wiki/CAMSAP3" title="CAMSAP3">CAMSAP3</a></li> <li><a href="/wiki/Centrin_1" title="Centrin 1">Centrin 1</a></li> <li><a href="/wiki/Centrin_2" title="Centrin 2">Centrin 2</a></li> <li><a href="/wiki/Centrin_3" title="Centrin 3">Centrin 3</a></li> <li><a href="/wiki/PCM1" title="PCM1">PCM1</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Microtubule severing proteins</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/Katanin" title="Katanin">Katanin</a></li> <li><a href="/wiki/Spastin" title="Spastin">Spastin</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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Tau_protein" title="Tau protein">Tau protein</a></li> <li><a href="/wiki/Dynactin" title="Dynactin">Dynactin</a> <ul><li><a href="/wiki/DCTN1" title="DCTN1">DCTN1</a></li></ul></li> <li><a href="/wiki/Stathmin" title="Stathmin">Stathmin</a></li> <li><a href="/wiki/Tektin" title="Tektin">Tektin</a> <ul><li><a href="/wiki/TEKT1" class="mw-redirect" title="TEKT1">TEKT1</a></li> <li><a href="/w/index.php?title=TEKT2&action=edit&redlink=1" class="new" title="TEKT2 (page does not exist)">TEKT2</a></li> <li><a href="/w/index.php?title=TEKT3&action=edit&redlink=1" class="new" title="TEKT3 (page does not exist)">TEKT3</a></li> <li><a href="/w/index.php?title=TEKT4&action=edit&redlink=1" class="new" title="TEKT4 (page does not exist)">TEKT4</a></li> <li><a href="/w/index.php?title=TEKT5&action=edit&redlink=1" class="new" title="TEKT5 (page does not exist)">TEKT5</a></li></ul></li> <li><a href="/wiki/Dynamin" title="Dynamin">Dynamin</a> <ul><li><a href="/wiki/DNM1" title="DNM1">DNM1</a></li> <li><a href="/wiki/DNM2" title="DNM2">DNM2</a></li> <li><a href="/wiki/DNM3" title="DNM3">DNM3</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Catenin" title="Catenin">Catenins</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/%CE%91-Catenin" title="Α-Catenin">Alpha catenin</a></li> <li><a href="/wiki/Catenin_beta-1" title="Catenin beta-1">Beta catenin</a> <ul><li><a href="/wiki/Adenomatous_polyposis_coli" title="Adenomatous polyposis coli">APC</a></li></ul></li> <li><a href="/wiki/Plakoglobin" title="Plakoglobin">Plakoglobin (gamma catenin)</a></li> <li><a href="/wiki/%CE%94-Catenin" title="Δ-Catenin">Delta catenin</a></li> <li><a href="/wiki/Gigaxonin" title="Gigaxonin">GAN</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Membrane_protein" title="Membrane protein">Membrane</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/Dystrophin" title="Dystrophin">Dystrophin</a> <ul><li><a href="/wiki/Dystroglycan" title="Dystroglycan">Dystroglycan</a></li></ul></li> <li><a href="/wiki/Utrophin" title="Utrophin">Utrophin</a></li> <li><a href="/wiki/Ankyrin" title="Ankyrin">Ankyrin</a> <ul><li><a href="/wiki/Ankyrin-1" title="Ankyrin-1">ANK1</a></li> <li><a href="/wiki/Ankyrin-2" title="Ankyrin-2">ANK2</a></li> <li><a href="/wiki/Ankyrin-3" title="Ankyrin-3">ANK3</a></li></ul></li> <li><a href="/wiki/Spectrin" title="Spectrin">Spectrin</a> <ul><li><a href="/wiki/Spectrin,_alpha_1" title="Spectrin, alpha 1">SPTA1</a></li> <li><a href="/wiki/SPTAN1" title="SPTAN1">SPTAN1</a></li> <li><a href="/wiki/SPTB" title="SPTB">SPTB</a></li> <li><a href="/wiki/SPTBN1" title="SPTBN1">SPTBN1</a></li> <li><a href="/wiki/SPTBN2" title="SPTBN2">SPTBN2</a></li> <li><a href="/wiki/SPTBN4" title="SPTBN4">SPTBN4</a></li> <li><a href="/wiki/SPTBN5" title="SPTBN5">SPTBN5</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Plakin" title="Plakin">Plakins</a> <ul><li><a href="/wiki/Corneodesmosin" title="Corneodesmosin">Corneodesmosin</a></li> <li><a href="/wiki/Desmoplakin" title="Desmoplakin">Desmoplakin</a></li> <li><a href="/wiki/Dystonin" title="Dystonin">Dystonin</a></li> <li><a href="/wiki/Envoplakin" title="Envoplakin">Envoplakin</a></li> <li><a href="/wiki/MACF1" title="MACF1">MACF1</a></li> <li><a href="/wiki/Periplakin" title="Periplakin">Periplakin</a></li> <li><a href="/wiki/Plectin" title="Plectin">Plectin</a></li></ul></li> <li><a href="/wiki/Talin_(protein)" title="Talin (protein)">Talin</a> <ul><li><a href="/wiki/TLN1" title="TLN1">TLN1</a></li></ul></li> <li><a href="/wiki/Vinculin" title="Vinculin">Vinculin</a></li> <li><a href="/wiki/Plakophilin" title="Plakophilin">Plakophilin</a> <ul><li><a href="/wiki/Plakophilin-1" title="Plakophilin-1">PKP1</a></li> <li><a href="/wiki/Plakophilin-2" title="Plakophilin-2">PKP2</a></li></ul></li> <li><a href="/wiki/ACF7" class="mw-redirect" title="ACF7">ACF7</a></li> <li><a href="/wiki/PLEKHA7" title="PLEKHA7">PLEKHA7</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Nonhuman</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/Major_sperm_protein" title="Major sperm protein">Major sperm proteins</a></li> <li><a href="/wiki/Prokaryotic_cytoskeleton" title="Prokaryotic cytoskeleton">Prokaryotic cytoskeleton</a> <ul><li><a href="/wiki/Crescentin" title="Crescentin">Crescentin</a></li> <li><a href="/wiki/FtsZ" title="FtsZ">FtsZ</a></li> <li><a href="/wiki/MreB" title="MreB">MreB</a></li> <li><a href="/wiki/ParM" title="ParM">ParM</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="padding:0;"><div><i>See also: <a href="/wiki/Template:Cytoskeletal_defects" title="Template:Cytoskeletal defects">cytoskeletal defects</a></i></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" 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8.83% 85.459 1 Template:External_media"," 7.97% 77.103 1 Template:Short_description"," 6.80% 65.785 1 Template:Harvnb"," 5.36% 51.904 1 Template:Lead_too_short"]},"scribunto":{"limitreport-timeusage":{"value":"0.684","limit":"10.000"},"limitreport-memusage":{"value":7547623,"limit":52428800},"limitreport-logs":"anchor_id_list = table#1 {\n [\"CITEREFBeiselKennedy1994\"] = 1,\n [\"CITEREFBernsteinBamburg1982\"] = 1,\n [\"CITEREFBianconePalmieriLombardiColantoni2003\"] = 1,\n [\"CITEREFBryceSchevzovFergusonPercival2003\"] = 1,\n [\"CITEREFBurgoyneNorman1985\"] = 2,\n [\"CITEREFDasDasguptaMandalGeng1993\"] = 1,\n [\"CITEREFDohlamLupasCarson1993\"] = 1,\n [\"CITEREFEyreAkkariWiltonCallen1995\"] = 1,\n [\"CITEREFGengBianconeDaiLin1998\"] = 1,\n [\"CITEREFGunningGhoshdastiderWhitakerPopp2015\"] = 1,\n [\"CITEREFGunningHardemanJeffreyWeinberger1998\"] = 1,\n [\"CITEREFGunningO\u0026#039;neillHardeman2008\"] = 1,\n [\"CITEREFGunningSchevzovKeeHardeman2005\"] = 1,\n [\"CITEREFHadFaivre-SarrailhLegrandMery2005\"] = 1,\n [\"CITEREFHannanGunningJeffreyWeinberger1995\"] = 1,\n [\"CITEREFHeimannPercivalWeinbergerGunning1999\"] = 1,\n [\"CITEREFHendricksWeintraub1981\"] = 1,\n [\"CITEREFHendricksWeintraub1984\"] = 1,\n [\"CITEREFHuntEyreAkkariMeredith1995\"] = 1,\n [\"CITEREFKeeSchevzovNair-ShallikerRobinson2004\"] = 1,\n [\"CITEREFKhannaNomuraFischettiZabriskie1997\"] = 1,\n [\"CITEREFLees-MillerGoodwinHelfman1990\"] = 1,\n [\"CITEREFLees-MillerHelfman1991\"] = 1,\n [\"CITEREFLinGengBhattacharyaYu2002\"] = 1,\n [\"CITEREFLinHegmannLin1988\"] = 1,\n [\"CITEREFLinHelfmanHughesChou1985\"] = 1,\n [\"CITEREFLinLin1986\"] = 1,\n [\"CITEREFLopataKleine-TebbeKamath2016\"] = 1,\n [\"CITEREFLuiBretscher1992\"] = 1,\n [\"CITEREFMartinSchevzovGunning2009\"] = 1,\n [\"CITEREFMiyadoKimuraTaniguchi1996\"] = 1,\n [\"CITEREFMorWeinbergerCohen2002\"] = 1,\n [\"CITEREFPawlakMcGarveyNguyenTomaszewski2004\"] = 1,\n [\"CITEREFPercivalHughesBrownSchevzov2004\"] = 1,\n [\"CITEREFPercivalThomasCockGardiner2000\"] = 1,\n [\"CITEREFPittengerKazzazHelfman1994\"] = 1,\n [\"CITEREFRuethersTakiJohnstonNugraha2018\"] = 1,\n [\"CITEREFRuiz-OpazoNadal-Ginard1987\"] = 1,\n [\"CITEREFRuiz-OpazoWeinbergerNadal-Ginard1985\"] = 1,\n [\"CITEREFSantosChapmanAalberseVailes1999\"] = 1,\n [\"CITEREFSchevzovBryceAlmonte-BaldonadoJoya2005\"] = 1,\n [\"CITEREFSchevzovGunningJeffreyTemm-Grove1997\"] = 1,\n [\"CITEREFSchevzovLloydHailstonesGunning1993\"] = 1,\n [\"CITEREFSchevzovVrhovskiBryceElmir2005\"] = 1,\n [\"CITEREFTakenagaNakamuraSakiyama1988\"] = 1,\n [\"CITEREFTakenagaNakamuraTokunagaKageyama1988\"] = 1,\n [\"CITEREFThalayasingamLee2015\"] = 1,\n [\"CITEREFTongYuenWaiLeung2018\"] = 1,\n [\"CITEREFWiltonEyreAkkariWatkins1995\"] = 1,\n [\"CITEREFWiltonLimDoroszGunn1996\"] = 1,\n [\"CITEREFbianconeMonteleoneMarascoPallone1998\"] = 1,\n}\ntemplate_list = table#1 {\n [\"Cite journal\"] = 52,\n [\"Cytoskeletal Proteins\"] = 1,\n [\"External media\"] = 1,\n [\"Harvnb\"] = 1,\n [\"Lead too short\"] = 1,\n [\"Muscle tissue\"] = 1,\n [\"PDB\"] = 1,\n [\"Pb\"] = 1,\n [\"Pfam box\"] = 1,\n [\"Reflist\"] = 1,\n [\"Short description\"] = 1,\n}\narticle_whitelist = table#1 {\n}\n"},"cachereport":{"origin":"mw-web.codfw.main-6b7f745dd4-7lg6z","timestamp":"20241125135345","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Tropomyosin","url":"https:\/\/en.wikipedia.org\/wiki\/Tropomyosin","sameAs":"http:\/\/www.wikidata.org\/entity\/Q418454","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q418454","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, 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