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Cysteine protease - Wikipedia
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Available in 16 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-16" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">16 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B3%D9%8A%D8%B3%D8%AA%D9%8A%D9%86_%D8%A8%D8%B1%D9%88%D8%AA%D9%8A%D8%A7%D8%B2" title="سيستين بروتياز – Arabic" lang="ar" hreflang="ar" data-title="سيستين بروتياز" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Cistein_peptidaza" title="Cistein peptidaza – Bosnian" lang="bs" hreflang="bs" data-title="Cistein peptidaza" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Cysteinproteasen" title="Cysteinproteasen – German" lang="de" hreflang="de" data-title="Cysteinproteasen" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Ciste%C3%ADna_proteasas" title="Cisteína proteasas – Spanish" lang="es" hreflang="es" data-title="Cisteína proteasas" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%DB%8C%D8%B3%D8%AA%D8%A6%DB%8C%D9%86_%D9%BE%D8%B1%D9%88%D8%AA%D8%A6%D8%A7%D8%B2" title="سیستئین پروتئاز – Persian" lang="fa" hreflang="fa" data-title="سیستئین پروتئاز" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Prot%C3%A9ase_%C3%A0_cyst%C3%A9ine" title="Protéase à cystéine – French" lang="fr" hreflang="fr" data-title="Protéase à cystéine" 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/Ciste%C3%ADna_protease" title="Cisteína protease – Galician" lang="gl" hreflang="gl" data-title="Cisteína protease" 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/Sisteina_protease" title="Sisteina protease – Indonesian" lang="id" hreflang="id" data-title="Sisteina protease" 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-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A6%D7%99%D7%A1%D7%98%D7%90%D7%99%D7%9F_%D7%A4%D7%A8%D7%95%D7%98%D7%90%D7%96" title="ציסטאין פרוטאז – Hebrew" lang="he" hreflang="he" data-title="ציסטאין פרוטאז" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%B7%E3%82%B9%E3%83%86%E3%82%A4%E3%83%B3%E3%83%97%E3%83%AD%E3%83%86%E3%82%A2%E3%83%BC%E3%82%BC" title="システインプロテアーゼ – Japanese" lang="ja" hreflang="ja" data-title="システインプロテアーゼ" 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/Proteazy_cysteinowe" title="Proteazy cysteinowe – Polish" lang="pl" hreflang="pl" data-title="Proteazy cysteinowe" 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/Ciste%C3%ADno_protease" title="Cisteíno protease – Portuguese" lang="pt" hreflang="pt" data-title="Cisteíno protease" 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%A6%D0%B8%D1%81%D1%82%D0%B5%D0%B8%D0%BD%D0%BE%D0%B2%D1%8B%D0%B5_%D0%BF%D1%80%D0%BE%D1%82%D0%B5%D0%B0%D0%B7%D1%8B" title="Цистеиновые протеазы – Russian" lang="ru" hreflang="ru" data-title="Цистеиновые протеазы" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Cisteinska_peptidaza" title="Cisteinska peptidaza – Serbian" lang="sr" hreflang="sr" data-title="Cisteinska peptidaza" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a 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class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Class of enzymes</div> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Protein family</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above">Cysteine peptidase, CA clan</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Papain_enzyme.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Papain_enzyme.png/220px-Papain_enzyme.png" decoding="async" width="220" height="149" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/22/Papain_enzyme.png/330px-Papain_enzyme.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/22/Papain_enzyme.png/440px-Papain_enzyme.png 2x" data-file-width="640" data-file-height="434" /></a></span><div class="infobox-caption">Crystal structure of the cysteine peptidase <a href="/wiki/Papain" title="Papain">papain</a> in complex with its covalent inhibitor E-64. Rendered from <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/1PE6">1PE6</a></span>​</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #ddd; color:inherit;">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;">Symbol</th><td class="infobox-data" style="background-color: #eee; color:inherit;">Peptidase_C1</td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Pfam" title="Pfam">Pfam</a></th><td class="infobox-data pfam" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/pfam/PF00112">PF00112</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Pfam" title="Pfam">Pfam</a> clan</th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/set/pfam/CL0125/">CL0125</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/InterPro" title="InterPro">InterPro</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/interpro/entry/IPR000668">IPR000668</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Simple_Modular_Architecture_Research_Tool" title="Simple Modular Architecture Research Tool">SMART</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="http://smart.embl-heidelberg.de/smart/do_annotation.pl?DOMAIN=SM00645">SM00645</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/PROSITE" title="PROSITE">PROSITE</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://prosite.expasy.org/PDOC00126">PDOC00126</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/MEROPS" title="MEROPS">MEROPS</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://www.ebi.ac.uk/merops/cgi-bin/famsum?family=C1">C1</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><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; color:inherit;"><a rel="nofollow" class="external text" href="http://scop2.mrc-lmb.cam.ac.uk/search?t=txt;q=1aec">1aec</a> / <a rel="nofollow" class="external text" href="https://scop.berkeley.edu/pdb/code=1aec">SCOPe</a> / <a rel="nofollow" class="external text" href="http://supfam.org/SUPERFAMILY/cgi-bin/search.cgi?search_field=1aec">SUPFAM</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM superfamily</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/protein_superfamilies/355">355</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit;"><a href="/wiki/Orientations_of_Proteins_in_Membranes_database" title="Orientations of Proteins in Membranes database">OPM protein</a></th><td class="infobox-data" style="background-color: #eee; color:inherit;"><a rel="nofollow" class="external text" href="https://opm.phar.umich.edu/proteins?search=1m6d">1m6d</a></td></tr><tr><td colspan="2" class="infobox-full-data" style="background-color: #eee; color:inherit;"><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; color:inherit;">Available protein structures:</th></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; 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; color:inherit; border:#fafafa 2px solid; border-width:3px 0 0 2px;"><a rel="nofollow" class="external text" href="http://pfam.xfam.org/family/PF00112?tab=pdbBlock">structures</a> / <a rel="nofollow" class="external text" href="http://prodata.swmed.edu/ecod/complete/search?kw=PF00112">ECOD</a>  </td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; 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; color:inherit; 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:PF00112%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:PF00112">PDBe</a>; <a rel="nofollow" class="external text" href="https://pdbj.org/search/pdb?other_db_select=PFam&other_db_field=PF00112">PDBj</a></td></tr><tr><th scope="row" class="infobox-label" style="background-color: #e7dcc3; color:inherit; 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; color:inherit; 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=PF00112">structure summary</a></td></tr></tbody></table></td></tr></tbody></table> <p><b>Cysteine proteases</b>, also known as <b>thiol proteases</b>, are <a href="/wiki/Hydrolase" title="Hydrolase">hydrolase</a> <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> that degrade <a href="/wiki/Protein" title="Protein">proteins</a>. These <a href="/wiki/Protease" title="Protease">proteases</a> share a common <a href="/wiki/Catalysis" title="Catalysis">catalytic mechanism</a> that involves a <a href="/wiki/Nucleophile" title="Nucleophile">nucleophilic</a> <a href="/wiki/Cysteine" title="Cysteine">cysteine</a> <a href="/wiki/Thiol" title="Thiol">thiol</a> in a <a href="/wiki/Catalytic_triad" title="Catalytic triad">catalytic triad</a> or dyad.<sup id="cite_ref-:0_1-0" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Discovered by Gopal Chunder Roy in 1873, the first cysteine protease to be isolated and characterized was <a href="/wiki/Papain" title="Papain">papain</a>, obtained from <a href="/wiki/Papaya" title="Papaya"><i>Carica papaya</i></a>.<sup id="cite_ref-:0_1-1" class="reference"><a href="#cite_note-:0-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Cysteine proteases are commonly encountered in <a href="/wiki/Fruits" class="mw-redirect" title="Fruits">fruits</a> including the <a href="/wiki/Papaya" title="Papaya">papaya</a>, <a href="/wiki/Pineapple" title="Pineapple">pineapple</a>, <a href="/wiki/Common_fig" class="mw-redirect" title="Common fig">fig</a> and <a href="/wiki/Kiwifruit" title="Kiwifruit">kiwifruit</a>. The proportion of protease tends to be higher when the fruit is unripe. In fact, the <a href="/wiki/Latex" title="Latex">latex</a> of dozens of different plant <a href="/wiki/Family_(biology)" title="Family (biology)">families</a> are known to contain cysteine proteases.<sup id="cite_ref-pmid18496785_2-0" class="reference"><a href="#cite_note-pmid18496785-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Cysteine proteases are used as an ingredient in meat tenderizers. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=1" title="Edit section: Classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/MEROPS" title="MEROPS">MEROPS</a> protease classification system counts 14 <a href="/wiki/Protein_superfamily" title="Protein superfamily">superfamilies</a> plus several currently unassigned families (as of 2013) each containing many <a href="/wiki/Protein_family" title="Protein family">families</a>. Each superfamily uses the <a href="/wiki/Catalytic_triad" title="Catalytic triad">catalytic triad</a> or dyad in a different <a href="/wiki/Protein_fold" class="mw-redirect" title="Protein fold">protein fold</a> and so represent <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a> of the <a href="/wiki/Catalytic_mechanism" class="mw-redirect" title="Catalytic mechanism">catalytic mechanism</a>. </p><p>For <a href="/wiki/Protein_superfamily" title="Protein superfamily">superfamilies</a>, P indicates a superfamily containing a mixture of <a href="/wiki/Nucleophile" title="Nucleophile">nucleophile</a> class families, and C indicates purely cysteine proteases. superfamily. Within each superfamily, <a href="/wiki/Protein_family" title="Protein family">families</a> are designated by their catalytic nucleophile (C denoting cysteine proteases). </p> <dl><dd><table class="wikitable"> <caption>Families of cysteine proteases </caption> <tbody><tr> <th><a href="/wiki/Protein_superfamily" title="Protein superfamily">Superfamily</a></th> <th><a href="/wiki/Protein_family" title="Protein family">Families</a></th> <th>Examples </th></tr> <tr> <td><a href="/wiki/Clan_CA" class="mw-redirect" title="Clan CA">CA</a></td> <td>C1, C2, C6, C10, C12, C16, C19, C28, C31, C32, C33, C39, C47, C51, C54, C58, C64, <p>C65, C66, C67, C70, C71, C76, C78, C83, C85, C86, C87, C93, C96, C98, C101 </p> </td> <td><a href="/wiki/Papain" title="Papain">Papain</a> (<i><a href="/wiki/Carica_papaya" class="mw-redirect" title="Carica papaya">Carica papaya</a></i>),<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Bromelain" title="Bromelain">bromelain</a> (<i><a href="/wiki/Ananas_comosus" class="mw-redirect" title="Ananas comosus">Ananas comosus</a></i>), <a href="/wiki/Cathepsin_K" title="Cathepsin K">cathepsin K</a> (<i><a href="/wiki/Liverwort" class="mw-redirect" title="Liverwort">liverwort</a></i>)<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Calpain" title="Calpain">calpain</a> (<i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i>)<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>CD</td> <td>C11, C13, C14, C25, C50, C80, C84</td> <td><a href="/wiki/Caspase_1" title="Caspase 1">Caspase 1</a> (<i><a href="/wiki/Rattus_norvegicus" class="mw-redirect" title="Rattus norvegicus">Rattus norvegicus</a></i>) and <a href="/wiki/Separase" title="Separase">separase</a> (<i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>) </td></tr> <tr> <td>CE</td> <td>C5, C48, C55, C57, C63, C79</td> <td><a href="/wiki/Adenain" title="Adenain">Adenain</a> (human <a href="/wiki/Adenovirus" class="mw-redirect" title="Adenovirus">adenovirus</a> type 2) </td></tr> <tr> <td>CF</td> <td>C15</td> <td><a href="/wiki/Pyroglutamyl-peptidase_I" title="Pyroglutamyl-peptidase I">Pyroglutamyl-peptidase I</a> (<i><a href="/wiki/Bacillus_amyloliquefaciens" title="Bacillus amyloliquefaciens">Bacillus amyloliquefaciens</a></i>) </td></tr> <tr> <td>CL</td> <td>C60, C82</td> <td><a href="/wiki/Sortase_A" title="Sortase A">Sortase A</a> (<i><a href="/wiki/Staphylococcus_aureus" title="Staphylococcus aureus">Staphylococcus aureus</a></i>) </td></tr> <tr> <td>CM</td> <td>C18</td> <td><a href="/wiki/NS2-3_protease" title="NS2-3 protease">Hepatitis C virus peptidase 2</a> (<a href="/wiki/Hepatitis_C_virus" title="Hepatitis C virus">hepatitis C virus</a>) </td></tr> <tr> <td>CN</td> <td>C9</td> <td><a href="/wiki/Sindbis_virus_protease" class="mw-redirect" title="Sindbis virus protease">Sindbis virus-type nsP2 peptidase</a> (<a href="/wiki/Sindbis_virus" title="Sindbis virus">sindbis virus</a>) </td></tr> <tr> <td>CO</td> <td>C40</td> <td><a href="/w/index.php?title=Dipeptidyl-peptidase_VI&action=edit&redlink=1" class="new" title="Dipeptidyl-peptidase VI (page does not exist)">Dipeptidyl-peptidase VI</a> (<i><a rel="nofollow" class="external text" href="http://microbewiki.kenyon.edu/index.php/Lysinibacillus_sphaericus_C3-41">Lysinibacillus sphaericus</a></i>) </td></tr> <tr> <td>CP</td> <td>C97</td> <td><a href="/w/index.php?title=DeSI-1_peptidase&action=edit&redlink=1" class="new" title="DeSI-1 peptidase (page does not exist)">DeSI-1 peptidase</a> (<i><a href="/wiki/Mus_musculus" class="mw-redirect" title="Mus musculus">Mus musculus</a></i>) </td></tr> <tr> <td><a href="/wiki/PA_clan" class="mw-redirect" title="PA clan">PA</a></td> <td><a href="/wiki/Picornain_3C" title="Picornain 3C">C3</a>, <a href="/wiki/TEV_protease" title="TEV protease">C4</a>, C24, <a href="/wiki/3C-like_protease" title="3C-like protease">C30</a>, C37, C62, C74, C99</td> <td><a href="/wiki/TEV_protease" title="TEV protease">TEV protease</a> (<a href="/wiki/Tobacco_etch_virus" title="Tobacco etch virus">tobacco etch virus</a>) </td></tr> <tr> <td>PB</td> <td>C44, C45, C59, C69, C89, C95</td> <td><a href="/wiki/Amidophosphoribosyltransferase" title="Amidophosphoribosyltransferase">Amidophosphoribosyltransferase</a> precursor (<i><a href="/wiki/Homo_sapiens" class="mw-redirect" title="Homo sapiens">Homo sapiens</a></i>) </td></tr> <tr> <td>PC</td> <td>C26, C56</td> <td><a href="/wiki/Gamma-glutamyl_hydrolase" title="Gamma-glutamyl hydrolase">Gamma-glutamyl hydrolase</a> (<i><a href="/wiki/Rattus_norvegicus" class="mw-redirect" title="Rattus norvegicus">Rattus norvegicus</a></i>) </td></tr> <tr> <td>PD</td> <td>C46</td> <td><a href="/wiki/Hedgehog_protein" class="mw-redirect" title="Hedgehog protein">Hedgehog protein</a> (<i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></i>) </td></tr> <tr> <td>PE</td> <td>P1</td> <td><a href="/wiki/Beta-peptidyl_aminopeptidase" title="Beta-peptidyl aminopeptidase">DmpA aminopeptidase</a> (<i><a href="/wiki/Brucella_anthropi" title="Brucella anthropi">Brucella anthropi</a></i>) </td></tr> <tr> <td>unassigned</td> <td>C7, C8, C21, C23, C27, C36, C42, C53, C75</td> <td> </td></tr></tbody></table></dd></dl> <div class="mw-heading mw-heading2"><h2 id="Catalytic_mechanism">Catalytic mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=2" title="Edit section: Catalytic mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Cysteinprotease_Reaktionsmechanismus.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Cysteinprotease_Reaktionsmechanismus.svg/300px-Cysteinprotease_Reaktionsmechanismus.svg.png" decoding="async" width="300" height="450" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Cysteinprotease_Reaktionsmechanismus.svg/450px-Cysteinprotease_Reaktionsmechanismus.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5c/Cysteinprotease_Reaktionsmechanismus.svg/600px-Cysteinprotease_Reaktionsmechanismus.svg.png 2x" data-file-width="733" data-file-height="1099" /></a><figcaption>Reaction mechanism of the cysteine protease mediated cleavage of a peptide bond.</figcaption></figure> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Catalytic_triad" title="Catalytic triad">catalytic triad</a></div> <p>The first step in the reaction mechanism by which cysteine proteases catalyze the hydrolysis of peptide bonds is de<a href="/wiki/Protonation" title="Protonation">protonation</a> of a <a href="/wiki/Thiol" title="Thiol">thiol</a> in the <a href="/wiki/Enzyme" title="Enzyme">enzyme</a>'s active site by an adjacent <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> with a basic <a href="/wiki/Side_chain" title="Side chain">side chain</a>, usually a <a href="/wiki/Histidine" title="Histidine">histidine</a> residue. The next step is nucleophilic attack by the <a href="/wiki/Deprotonated" class="mw-redirect" title="Deprotonated">deprotonated</a> cysteine's <a href="/wiki/Anion" class="mw-redirect" title="Anion">anionic</a> <a href="/wiki/Sulfur" title="Sulfur">sulfur</a> on the <a href="/wiki/Substrate_(biochemistry)" class="mw-redirect" title="Substrate (biochemistry)">substrate</a> <a href="/wiki/Carbonyl" class="mw-redirect" title="Carbonyl">carbonyl</a> <a href="/wiki/Carbon" title="Carbon">carbon</a>. In this step, a fragment of the substrate is released with an <a href="/wiki/Amine" title="Amine">amine</a> terminus, the <a href="/wiki/Histidine" title="Histidine">histidine</a> residue in the <a href="/wiki/Protease" title="Protease">protease</a> is restored to its deprotonated form, and a <a href="/wiki/Thioester" title="Thioester">thioester</a> intermediate linking the new <a href="/wiki/Carboxy-terminus" class="mw-redirect" title="Carboxy-terminus">carboxy-terminus</a> of the substrate to the cysteine <a href="/wiki/Thiol" title="Thiol">thiol</a> is formed. Therefore, they are also sometimes referred to as thiol proteases. The <a href="/wiki/Thioester" title="Thioester">thioester</a> bond is subsequently <a href="/wiki/Hydrolysis" title="Hydrolysis">hydrolyzed</a> to generate a <a href="/wiki/Carboxylic_acid" title="Carboxylic acid">carboxylic acid</a> moiety on the remaining substrate fragment, while regenerating the free enzyme.<sup id="cite_ref-:1_6-0" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Biological_importance">Biological importance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=3" title="Edit section: Biological importance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cysteine proteases play multifaceted roles, virtually in every aspect of physiology and development. In plants they are important in growth and development and in accumulation and mobilization of storage proteins such as in seeds. In addition, they are involved in <a href="/wiki/Signalling_pathway" class="mw-redirect" title="Signalling pathway">signalling pathways</a> and in the response to <a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">biotic</a> and <a href="/wiki/Abiotic" class="mw-redirect" title="Abiotic">abiotic</a> stresses.<sup id="cite_ref-pmid15448724_7-0" class="reference"><a href="#cite_note-pmid15448724-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> In humans and other animals, they are responsible for <a href="/wiki/Senescence" title="Senescence">senescence</a> and <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a> (programmed cell death), MHC class II <a href="/wiki/Immune_response" title="Immune response">immune responses</a>, <a href="/wiki/Prohormone" title="Prohormone">prohormone</a> processing, and <a href="/wiki/Extracellular_matrix" title="Extracellular matrix">extracellular matrix</a> remodeling important to bone development. The ability of <a href="/wiki/Macrophages" class="mw-redirect" title="Macrophages">macrophages</a> and other cells to mobilize elastolytic cysteine proteases to their surfaces under specialized conditions may also lead to accelerated <a href="/wiki/Collagen" title="Collagen">collagen</a> and <a href="/wiki/Elastin" title="Elastin">elastin</a> degradation at sites of <a href="/wiki/Inflammation" title="Inflammation">inflammation</a> in <a href="/wiki/Diseases" class="mw-redirect" title="Diseases">diseases</a> such as <a href="/wiki/Atherosclerosis" title="Atherosclerosis">atherosclerosis</a> and <a href="/wiki/Emphysema" title="Emphysema">emphysema</a>.<sup id="cite_ref-pmid9074757_8-0" class="reference"><a href="#cite_note-pmid9074757-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Several viruses (such as <a href="/wiki/Polio" title="Polio">polio</a> and <a href="/wiki/Hepatitis_C_virus" title="Hepatitis C virus">hepatitis C</a>) express their entire <a href="/wiki/Genome" title="Genome">genome</a> as a single massive <a href="/wiki/Polyprotein" class="mw-redirect" title="Polyprotein">polyprotein</a> and use a protease to cleave it into functional units (for example, <a href="/wiki/Tobacco_etch_virus_protease" class="mw-redirect" title="Tobacco etch virus protease">tobacco etch virus protease</a>). </p> <div class="mw-heading mw-heading2"><h2 id="Regulation">Regulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=4" title="Edit section: Regulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The activity of cysteine proteases is regulated by a few general mechanisms, which includes the production of <a href="/wiki/Zymogenous" class="mw-redirect" title="Zymogenous">zymogens</a>, selective expression, pH modification, cellular compartmentalization, and regulation of their enzymatic activity by endogenous <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">inhibitors</a>, which seemingly is the most efficient mechanism associated with the regulation of the activity of cysteine proteases.<sup id="cite_ref-:1_6-1" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Proteases are usually synthesized as large precursor proteins called <a href="/wiki/Zymogens" class="mw-redirect" title="Zymogens">zymogens</a>, such as the <a href="/wiki/Serine_protease" title="Serine protease">serine protease</a> precursors <a href="/wiki/Trypsinogen" title="Trypsinogen">trypsinogen</a> and <a href="/wiki/Chymotrypsinogen" title="Chymotrypsinogen">chymotrypsinogen</a>, and the <a href="/wiki/Aspartic_protease" title="Aspartic protease">aspartic protease</a> precursor <a href="/wiki/Pepsinogen" class="mw-redirect" title="Pepsinogen">pepsinogen</a>. The protease is activated by removal of an inhibitory segment or protein. Activation occurs once the protease is delivered to a specific intracellular compartment (for example the <a href="/wiki/Lysosome" title="Lysosome">lysosome</a>) or extracellular environment (for example the <a href="/wiki/Stomach" title="Stomach">stomach</a>). This system prevents the <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> that produces the protease from being damaged by it. </p><p>Protease <a href="/wiki/Enzyme_inhibitor" title="Enzyme inhibitor">inhibitors</a> are usually proteins with <a href="/wiki/Structural_domain" class="mw-redirect" title="Structural domain">domains</a> that enter or block a protease <a href="/wiki/Active_site" title="Active site">active site</a> to prevent <a href="/wiki/Substrate_(biochemistry)" class="mw-redirect" title="Substrate (biochemistry)">substrate</a> access. In <a href="/wiki/Competitive_inhibition" title="Competitive inhibition">competitive inhibition</a>, the inhibitor binds to the active site, thus preventing enzyme-substrate interaction. In <a href="/wiki/Non-competitive_inhibition" title="Non-competitive inhibition">non-competitive inhibition</a>, the inhibitor binds to an <a href="/wiki/Allosteric_site" class="mw-redirect" title="Allosteric site">allosteric site</a>, which alters the active site and makes it inaccessible to the substrate. </p><p>Examples of protease inhibitors include: </p> <ul><li><a href="/wiki/Serpin" title="Serpin">Serpins</a></li> <li><a href="/w/index.php?title=Stefin&action=edit&redlink=1" class="new" title="Stefin (page does not exist)">Stefins</a></li> <li><a href="/wiki/Inhibitor_of_apoptosis" title="Inhibitor of apoptosis">IAPs</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="Uses">Uses</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=5" title="Edit section: Uses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Papain#Uses" title="Papain">Papain § Uses</a>, <a href="/wiki/Bromelain#Uses" title="Bromelain">Bromelain § Uses</a>, <a href="/wiki/Ficain" title="Ficain">Ficain</a>, and <a href="/wiki/Chymopapain#Medical_applications" title="Chymopapain">Chymopapain § Medical applications</a></div> <div class="mw-heading mw-heading3"><h3 id="Potential_pharmaceuticals">Potential pharmaceuticals</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=6" title="Edit section: Potential pharmaceuticals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Currently there is no widespread use of cysteine proteases as approved and effective <a href="/wiki/Anthelmintic" title="Anthelmintic">anthelmintics</a> but research into the subject is a promising field of study. Plant cysteine proteases isolated from these plants have been found to have high <a href="/wiki/Proteolysis" title="Proteolysis">proteolytic activities</a> that are known to digest <a href="/wiki/Nematode" title="Nematode">nematode</a> <a href="/wiki/Cuticle" title="Cuticle">cuticles</a>, with very low toxicity.<sup id="cite_ref-pmid15193563_9-0" class="reference"><a href="#cite_note-pmid15193563-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Successful results have been reported against nematodes such as <i><a href="/wiki/Heligmosomoides_bakeri" class="mw-redirect" title="Heligmosomoides bakeri">Heligmosomoides bakeri</a></i>, <i><a href="/wiki/Trichinella_spiralis" title="Trichinella spiralis">Trichinella spiralis</a></i>, <i><a href="/wiki/Nippostrongylus_brasiliensis" title="Nippostrongylus brasiliensis">Nippostrongylus brasiliensis</a></i>, <i><a href="/wiki/Trichuris_muris" title="Trichuris muris">Trichuris muris</a></i>, and <i><a href="/wiki/Ancylostoma_ceylanicum" title="Ancylostoma ceylanicum">Ancylostoma ceylanicum</a></i>; the <a href="/wiki/Tapeworm" class="mw-redirect" title="Tapeworm">tapeworm</a> <i><a href="/wiki/Rodentolepis_microstoma" class="mw-redirect" title="Rodentolepis microstoma">Rodentolepis microstoma</a></i>, and the <a href="/wiki/Porcine" class="mw-redirect" title="Porcine">porcine</a> <a href="/wiki/Acanthocephala" title="Acanthocephala">acanthocephalan</a> parasite <i><a href="/wiki/Macracanthorhynchus" title="Macracanthorhynchus">Macracanthorhynchus</a> hirundinaceus</i>.<sup id="cite_ref-pmid18761736_10-0" class="reference"><a href="#cite_note-pmid18761736-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> A useful property of cysteine proteases is the resistance to acid digestion, allowing possible <a href="/wiki/Oral_administration" title="Oral administration">oral administration</a>. They provide an alternative mechanism of action to current anthelmintics and the development of resistance is thought to be unlikely because it would require a complete change of structure of the helminth <a href="/wiki/Cuticle" title="Cuticle">cuticle</a>. </p><p>In several <a href="/wiki/Traditional_medicine" title="Traditional medicine">traditional medicines</a>, the fruits or latex of the papaya, pineapple and fig are widely used for treatment of <a href="/wiki/Intestinal_parasite" class="mw-redirect" title="Intestinal parasite">intestinal worm</a> <a href="/wiki/Infections" class="mw-redirect" title="Infections">infections</a> both in humans and <a href="/wiki/Livestock" title="Livestock">livestock</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Other">Other</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=7" title="Edit section: Other"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cysteine proteases are used as feed additives for livestock to improve the digestibility of proteins and amino acids.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=8" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col"> <ul><li><a href="/wiki/Protease" title="Protease">Protease</a> <ul><li><a href="/wiki/Serine_protease" title="Serine protease">Serine protease</a></li> <li><a href="/wiki/Threonine_protease" title="Threonine protease">Threonine protease</a></li> <li><a href="/wiki/Aspartic_protease" title="Aspartic protease">Aspartic protease</a></li> <li><a href="/wiki/Metalloprotease" class="mw-redirect" title="Metalloprotease">Metalloprotease</a></li></ul></li> <li><a href="/wiki/Enzyme" title="Enzyme">Enzyme</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">Proteolysis</a></li> <li><a href="/wiki/Catalytic_triad" title="Catalytic triad">Catalytic triad</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergent evolution</a></li> <li><a href="/wiki/PA_clan" class="mw-redirect" title="PA clan">PA clan</a></li> <li><a href="/wiki/The_Proteolysis_Map" title="The Proteolysis Map">The Proteolysis Map</a></li> <li><a href="/wiki/Protease_inhibitor_(pharmacology)" title="Protease inhibitor (pharmacology)">Protease inhibitor (pharmacology)</a></li> <li><a href="/wiki/Protease_inhibitor_(biology)" title="Protease inhibitor (biology)">Protease inhibitor (biology)</a></li> <li><a href="/wiki/TopFIND" title="TopFIND">TopFIND</a> – database of protease specificity, substrates, products and inhibitors</li> <li><a href="/wiki/MEROPS" title="MEROPS">MEROPS</a> – database of protease evolutionary groups</li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cysteine_protease&action=edit&section=9" 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-:0-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFRawatRoyJyotiKaushik2021" class="citation journal cs1">Rawat, Aadish; Roy, Mrinalini; Jyoti, Anupam; Kaushik, Sanket; Verma, Kuldeep; Srivastava, Vijay Kumar (August 2021). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.micres.2021.126784">"Cysteine proteases: Battling pathogenic parasitic protozoans with omnipresent enzymes"</a>. <i>Microbiological Research</i>. <b>249</b>: 126784. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" 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Wattagnet<span class="reference-accessdate">. Retrieved <span class="nowrap">6 January</span> 2018</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Protease+enzymes+improve+amino+acid+digestibility&rft.pub=Wattagnet&rft.date=2012-04-06&rft.aulast=O%27Keefe&rft.aufirst=Terrence&rft_id=https%3A%2F%2Fwww.wattagnet.com%2Farticles%2F12390-protease-enzymes-improve-amino-acid-digestibility&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACysteine+protease" 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 href="/w/index.php?title=Cysteine_protease&action=edit&section=10" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>The <a href="/wiki/MEROPS" title="MEROPS">MEROPS</a> online database for peptidases and their inhibitors: <a rel="nofollow" class="external text" href="http://merops.sanger.ac.uk/cgi-bin/family_index?type=P#C">Cysteine Peptidases</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170404112415/http://merops.sanger.ac.uk/cgi-bin/family_index?type=P#C">Archived</a> 2017-04-04 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Cysteine+endopeptidases">Cysteine+endopeptidases</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output 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href="/wiki/Template:Proteases" title="Template:Proteases"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Proteases" title="Template talk:Proteases"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Proteases" title="Special:EditPage/Template:Proteases"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Hydrolase:_proteases_(EC_3.4)336" style="font-size:114%;margin:0 4em"><a href="/wiki/Hydrolase" title="Hydrolase">Hydrolase</a>: <a href="/wiki/Protease" title="Protease">proteases</a> (<a href="/wiki/Enzyme_Commission_number" title="Enzyme Commission number">EC</a> 3.4)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.11_Aminopeptidases" title="List of EC numbers (EC 3)">3.4.11</a>-19: <a href="/wiki/Exopeptidase" title="Exopeptidase">Exopeptidase</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.11_Aminopeptidases" title="List of EC numbers (EC 3)">3.4.11</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/Aminopeptidase" title="Aminopeptidase">Aminopeptidase</a> <ul><li><a href="/wiki/Alanine_aminopeptidase" title="Alanine aminopeptidase">Alanine</a></li> <li><a href="/wiki/RNPEP" title="RNPEP">Arginyl</a></li> <li><a href="/wiki/DNPEP" title="DNPEP">Aspartyl</a></li> <li><a href="/wiki/Cystinyl_aminopeptidase" class="mw-redirect" title="Cystinyl aminopeptidase">Cystinyl</a></li> <li><a href="/wiki/Leucyl_aminopeptidase" title="Leucyl aminopeptidase">Leucyl</a></li> <li><a href="/wiki/Glutamyl_aminopeptidase" title="Glutamyl aminopeptidase">Glutamyl</a></li> <li><i>Methionyl</i> <ul><li><a href="/wiki/METAP1" title="METAP1">1</a></li> <li><a href="/wiki/METAP2" title="METAP2">2</a></li></ul></li> <li><a href="/wiki/C9orf3" title="C9orf3">O</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.13_Dipeptidases" title="List of EC numbers (EC 3)">3.4.13</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/Dipeptidase" title="Dipeptidase">Dipeptidase</a> <ul><li><a href="/wiki/Dipeptidase_1" title="Dipeptidase 1">1</a></li> <li><a href="/wiki/DPEP2" class="mw-redirect" title="DPEP2">2</a></li> <li><a href="/wiki/DPEP3" class="mw-redirect" title="DPEP3">3</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.14_Dipeptidyl_peptidases_and_tripeptidyl_peptidases" title="List of EC numbers (EC 3)">3.4.14</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/Dipeptidyl_peptidase" title="Dipeptidyl peptidase">Dipeptidyl peptidase</a> <ul><li><a href="/wiki/Cathepsin_C" title="Cathepsin C">Cathepsin C</a></li> <li><a href="/wiki/Dipeptidyl_peptidase-4" title="Dipeptidyl peptidase-4">Dipeptidyl peptidase-4</a></li></ul></li> <li><a href="/wiki/Tripeptidyl_peptidase" title="Tripeptidyl peptidase">Tripeptidyl peptidase</a> <ul><li><a href="/wiki/Tripeptidyl_peptidase_I" title="Tripeptidyl peptidase I">Tripeptidyl peptidase I</a></li> <li><a href="/wiki/Tripeptidyl_peptidase_II" title="Tripeptidyl peptidase II">Tripeptidyl peptidase II</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.15_Peptidyl_dipeptidases" title="List of EC numbers (EC 3)">3.4.15</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/Angiotensin-converting_enzyme" title="Angiotensin-converting enzyme">Angiotensin-converting enzyme</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.16_Serine_type_carboxypeptidases" title="List of EC numbers (EC 3)">3.4.16</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/Serine_type_carboxypeptidase" class="mw-redirect" title="Serine type carboxypeptidase">Serine type carboxypeptidases</a>: <a href="/wiki/Cathepsin_A" title="Cathepsin A">Cathepsin A</a></li> <li><a href="/wiki/DD-Transpeptidase" title="DD-Transpeptidase"><small>DD</small>-Transpeptidase</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.17_Metallocarboxypeptidases" title="List of EC numbers (EC 3)">3.4.17</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <dl><dt><span class="nobold"><a href="/wiki/Metalloexopeptidase" title="Metalloexopeptidase">Metalloexopeptidases</a></span></dt> <dd><a href="/wiki/Carboxypeptidase" title="Carboxypeptidase">Carboxypeptidase</a> <dl><dd><a href="/wiki/Carboxypeptidase_A" title="Carboxypeptidase A">A</a></dd> <dd><a href="/wiki/Carboxypeptidase_A2" title="Carboxypeptidase A2">A2</a></dd> <dd><a href="/wiki/Carboxypeptidase_B" title="Carboxypeptidase B">B</a></dd> <dd><a href="/wiki/Cathepsin_A" title="Cathepsin A">C</a></dd> <dd><a href="/wiki/Carboxypeptidase_E" title="Carboxypeptidase E">E</a></dd> <dd><a href="/wiki/Glutamate_carboxypeptidase_II" title="Glutamate carboxypeptidase II">Glutamate II</a></dd></dl></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other/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/Metalloexopeptidase" title="Metalloexopeptidase">Metalloexopeptidase</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/List_of_EC_numbers_(EC_3)#EC_3.4.21:_Serine_proteases" title="List of EC numbers (EC 3)">3.4.21</a>-25: <a href="/wiki/Endopeptidase" title="Endopeptidase">Endopeptidase</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Serine_protease" title="Serine protease">Serine protease</a></li> <li><a class="mw-selflink selflink">Cysteine protease</a></li> <li><a href="/wiki/Aspartate_protease" class="mw-redirect" title="Aspartate protease">Aspartic acid protease</a></li> <li><a href="/wiki/Metalloendopeptidase" title="Metalloendopeptidase">Metalloendopeptidase</a></li> <li><a href="/wiki/Threonine_endopeptidase" class="mw-redirect" title="Threonine endopeptidase">Threonine endopeptidase</a> <ul><li><a href="/wiki/Proteasome_endopeptidase_complex" title="Proteasome endopeptidase complex">Proteasome endopeptidase complex</a></li> <li><a href="/wiki/HslU%E2%80%94HslV_peptidase" class="mw-redirect" title="HslU—HslV peptidase">HslU—HslV peptidase</a></li></ul></li></ul> <ul><li>Other/ungrouped: <a href="/wiki/Amyloid_precursor_protein_secretase" class="mw-redirect" title="Amyloid precursor protein secretase">Amyloid precursor protein secretase</a> <ul><li><a href="/wiki/Alpha_secretase" title="Alpha secretase">Alpha secretase</a></li> <li><a href="/wiki/Beta-secretase_1" title="Beta-secretase 1">Beta-secretase 1</a></li> <li><a href="/wiki/Beta-secretase_2" title="Beta-secretase 2">Beta-secretase 2</a></li> <li><a href="/wiki/Gamma_secretase" title="Gamma secretase">Gamma secretase</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.99_Endopeptidases_of_unknown_catalytic_mechanism" title="List of EC numbers (EC 3)">3.4.99</a>: Unknown</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Staphylokinase" title="Staphylokinase">Staphylokinase</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Proteases:_cysteine_proteases_(EC_3.4.22)190" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Cysteine_proteases" title="Template:Cysteine proteases"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cysteine_proteases" title="Template talk:Cysteine proteases"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cysteine_proteases" title="Special:EditPage/Template:Cysteine proteases"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Proteases:_cysteine_proteases_(EC_3.4.22)190" style="font-size:114%;margin:0 4em"><a href="/wiki/Protease" title="Protease">Proteases</a>: <a class="mw-selflink selflink">cysteine proteases</a> (<a href="/wiki/Enzyme_Commission_number" title="Enzyme Commission number">EC</a> <a href="/wiki/List_of_EC_numbers_(EC_3)#EC_3.4.22_Cysteine_proteases" title="List of EC numbers (EC 3)">3.4.22</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Caspase" title="Caspase">Caspase</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Caspase_1" title="Caspase 1">Caspase 1</a></li> <li><a href="/wiki/Caspase_2" title="Caspase 2">Caspase 2</a></li> <li><a href="/wiki/Caspase_3" title="Caspase 3">Caspase 3</a></li> <li><a href="/wiki/Caspase_4" title="Caspase 4">Caspase 4</a></li> <li><a href="/wiki/Caspase_5" title="Caspase 5">Caspase 5</a></li> <li><a href="/wiki/Caspase_6" title="Caspase 6">Caspase 6</a></li> <li><a href="/wiki/Caspase_7" title="Caspase 7">Caspase 7</a></li> <li><a href="/wiki/Caspase_8" title="Caspase 8">Caspase 8</a></li> <li><a href="/wiki/Caspase-9" title="Caspase-9">Caspase 9</a></li> <li><a href="/wiki/Caspase_10" title="Caspase 10">Caspase 10</a></li> <li><a href="/wiki/Caspase_12" title="Caspase 12">Caspase 12</a></li> <li><a href="/wiki/Caspase_13" title="Caspase 13">Caspase 13</a></li> <li><a href="/wiki/Caspase_14" title="Caspase 14">Caspase 14</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fruit-derived</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Papain" title="Papain">Papain</a></li> <li><a href="/wiki/Ficain" title="Ficain">Ficain</a></li> <li><a href="/wiki/Bromelain" title="Bromelain">Bromelain</a></li> <li><a href="/wiki/Actinidain" title="Actinidain">Actinidain</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Calpain" title="Calpain">Calpain</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Calpain-1_catalytic_subunit" title="Calpain-1 catalytic subunit">CAPN1</a></li> <li><a href="/wiki/Calpain-2_catalytic_subunit" title="Calpain-2 catalytic subunit">CAPN2</a></li> <li><a href="/wiki/Calpain-3" title="Calpain-3">CAPN3</a></li> <li><a href="/wiki/CAPN5" title="CAPN5">CAPN5</a></li> <li><a href="/wiki/Calpain-6" title="Calpain-6">CAPN6</a></li> <li><a href="/w/index.php?title=CAPN7&action=edit&redlink=1" class="new" title="CAPN7 (page does not exist)">CAPN7</a></li> <li><a href="/wiki/Calpain-8" title="Calpain-8">CAPN8</a></li> <li><a href="/wiki/Calpain-9" title="Calpain-9">CAPN9</a></li> <li><a href="/wiki/CAPN10" title="CAPN10">CAPN10</a></li> <li><a href="/w/index.php?title=CAPN11&action=edit&redlink=1" class="new" title="CAPN11 (page does not exist)">CAPN11</a></li> <li><a href="/w/index.php?title=CAPN12&action=edit&redlink=1" class="new" title="CAPN12 (page does not exist)">CAPN12</a></li> <li><a href="/w/index.php?title=CAPN13&action=edit&redlink=1" class="new" title="CAPN13 (page does not exist)">CAPN13</a></li> <li><a href="/w/index.php?title=CAPN14&action=edit&redlink=1" class="new" title="CAPN14 (page does not exist)">CAPN14</a></li> <li><a href="/wiki/Calpain_small_subunit_1" title="Calpain small subunit 1">CAPNS1</a></li> <li><a href="/w/index.php?title=CAPNS2&action=edit&redlink=1" class="new" title="CAPNS2 (page does not exist)">CAPNS2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cathepsin" title="Cathepsin">Cathepsin</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cathepsin_B" title="Cathepsin B">B</a></li> <li><a href="/wiki/Cathepsin_C" title="Cathepsin C">C</a></li> <li><a href="/wiki/Cathepsin_F" title="Cathepsin F">F</a></li> <li><a href="/wiki/Cathepsin_H" title="Cathepsin H">H</a></li> <li><a href="/wiki/Cathepsin_K" title="Cathepsin K">K</a></li> <li><a href="/wiki/Cathepsin_L1" title="Cathepsin L1">L1</a>/<a href="/wiki/Cathepsin_L2" title="Cathepsin L2">L2</a></li> <li><a href="/wiki/Cathepsin_O" title="Cathepsin O">O</a></li> <li><a href="/wiki/Cathepsin_S" title="Cathepsin S">S</a></li> <li><a href="/wiki/Cathepsin_W" title="Cathepsin W">W</a></li> <li><a href="/wiki/Cathepsin_Z" title="Cathepsin Z">Z</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Clostripain" title="Clostripain">Clostripain</a></li> <li><a href="/wiki/Cancer_procoagulant" title="Cancer procoagulant">Cancer procoagulant</a></li> <li><a href="/wiki/Separase" title="Separase">Separase</a></li> <li><a href="/wiki/Autophagin" title="Autophagin">Autophagin</a></li> <li><a href="/wiki/Cruzipain" title="Cruzipain">Cruzipain</a></li> <li><a href="/wiki/3C-like_protease" title="3C-like protease">3C-like protease</a></li> <li><a href="/wiki/Gingipain_R" title="Gingipain R">Gingipain R</a></li> <li><a href="/wiki/Gingipain_K" title="Gingipain K">Gingipain K</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Enzymes56" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link 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href="/wiki/Active_site" title="Active site">Active site</a></li> <li><a href="/wiki/Binding_site" title="Binding site">Binding site</a></li> <li><a href="/wiki/Catalytic_triad" title="Catalytic triad">Catalytic triad</a></li> <li><a href="/wiki/Oxyanion_hole" title="Oxyanion hole">Oxyanion hole</a></li> <li><a href="/wiki/Enzyme_promiscuity" title="Enzyme promiscuity">Enzyme promiscuity</a></li> <li><a href="/wiki/Diffusion-limited_enzyme" title="Diffusion-limited enzyme">Diffusion-limited enzyme</a></li> <li><a href="/wiki/Cofactor_(biochemistry)" title="Cofactor (biochemistry)">Cofactor</a></li> <li><a href="/wiki/Enzyme_catalysis" title="Enzyme catalysis">Enzyme catalysis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Regulation</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allosteric_regulation" title="Allosteric regulation">Allosteric 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navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Enzyme_kinetics" title="Enzyme kinetics">Enzyme kinetics</a></li> <li><a href="/wiki/Eadie%E2%80%93Hofstee_diagram" title="Eadie–Hofstee diagram">Eadie–Hofstee diagram</a></li> <li><a href="/wiki/Hanes%E2%80%93Woolf_plot" title="Hanes–Woolf plot">Hanes–Woolf plot</a></li> <li><a href="/wiki/Lineweaver%E2%80%93Burk_plot" title="Lineweaver–Burk plot">Lineweaver–Burk plot</a></li> <li><a href="/wiki/Michaelis%E2%80%93Menten_kinetics" title="Michaelis–Menten kinetics">Michaelis–Menten kinetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><b>EC1 <a href="/wiki/Oxidoreductase" title="Oxidoreductase">Oxidoreductases</a></b> (<a href="/wiki/List_of_EC_numbers_(EC_1)" title="List of EC numbers (EC 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