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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-Levels_of_protein_structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Levels_of_protein_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>Levels of protein structure</span> </div> </a> <button aria-controls="toc-Levels_of_protein_structure-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 Levels of protein structure subsection</span> </button> <ul id="toc-Levels_of_protein_structure-sublist" class="vector-toc-list"> <li id="toc-Primary_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Primary_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Primary structure</span> </div> </a> <ul id="toc-Primary_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Secondary_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Secondary_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Secondary structure</span> </div> </a> <ul id="toc-Secondary_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tertiary_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tertiary_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Tertiary structure</span> </div> </a> <ul id="toc-Tertiary_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Quaternary_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Quaternary_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Quaternary structure</span> </div> </a> <ul id="toc-Quaternary_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Homomers" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Homomers"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Homomers</span> </div> </a> <ul id="toc-Homomers-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Domains,_motifs,_and_folds_in_protein_structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Domains,_motifs,_and_folds_in_protein_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Domains, motifs, and folds in protein structure</span> </div> </a> <button aria-controls="toc-Domains,_motifs,_and_folds_in_protein_structure-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 Domains, motifs, and folds in protein structure subsection</span> </button> <ul id="toc-Domains,_motifs,_and_folds_in_protein_structure-sublist" class="vector-toc-list"> <li id="toc-Structural_domain" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structural_domain"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Structural domain</span> </div> </a> <ul id="toc-Structural_domain-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structural_and_sequence_motifs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structural_and_sequence_motifs"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Structural and sequence motifs</span> </div> </a> <ul id="toc-Structural_and_sequence_motifs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Supersecondary_structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Supersecondary_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Supersecondary structure</span> </div> </a> <ul id="toc-Supersecondary_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_fold" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Protein_fold"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Protein fold</span> </div> </a> <ul id="toc-Protein_fold-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Protein_dynamics_and_conformational_ensembles" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Protein_dynamics_and_conformational_ensembles"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Protein dynamics and conformational ensembles</span> </div> </a> <ul id="toc-Protein_dynamics_and_conformational_ensembles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_folding" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Protein_folding"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Protein folding</span> </div> </a> <ul id="toc-Protein_folding-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_stability" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Protein_stability"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Protein stability</span> </div> </a> <ul id="toc-Protein_stability-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_structure_determination" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Protein_structure_determination"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Protein structure determination</span> </div> </a> <ul id="toc-Protein_structure_determination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Protein_structure_databases" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Protein_structure_databases"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Protein structure databases</span> </div> </a> <ul id="toc-Protein_structure_databases-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Structural_classifications_of_proteins" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Structural_classifications_of_proteins"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Structural classifications of proteins</span> </div> </a> <ul id="toc-Structural_classifications_of_proteins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Computational_prediction_of_protein_structure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Computational_prediction_of_protein_structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Computational prediction of protein structure</span> </div> </a> <ul id="toc-Computational_prediction_of_protein_structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 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" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Protein structure</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 34 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-34" 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">34 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%A8%D9%86%D9%8A%D8%A9_%D8%A7%D9%84%D8%A8%D8%B1%D9%88%D8%AA%D9%8A%D9%86" title="بنية البروتين – Arabic" lang="ar" hreflang="ar" data-title="بنية البروتين" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ast badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://ast.wikipedia.org/wiki/Estructura_de_les_prote%C3%ADnes" title="Estructura de les proteínes – Asturian" lang="ast" hreflang="ast" data-title="Estructura de les proteínes" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Struktura_proteina" title="Struktura proteina – Bosnian" lang="bs" hreflang="bs" data-title="Struktura proteina" 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/Proteinstruktur" title="Proteinstruktur – German" lang="de" hreflang="de" data-title="Proteinstruktur" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Estructura_de_las_prote%C3%ADnas" title="Estructura de las proteínas – Spanish" lang="es" hreflang="es" data-title="Estructura de las proteínas" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Proteina_strukturo" title="Proteina strukturo – Esperanto" lang="eo" hreflang="eo" data-title="Proteina strukturo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D8%A7%D8%AE%D8%AA%D8%A7%D8%B1_%D9%BE%D8%B1%D9%88%D8%AA%D8%A6%DB%8C%D9%86" title="ساختار پروتئین – Persian" lang="fa" hreflang="fa" data-title="ساختار پروتئین" 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/Structure_des_prot%C3%A9ines" title="Structure des protéines – French" lang="fr" hreflang="fr" data-title="Structure des protéines" 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/Estrutura_das_prote%C3%ADnas" title="Estrutura das proteínas – Galician" lang="gl" hreflang="gl" data-title="Estrutura das proteínas" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%8B%A8%EB%B0%B1%EC%A7%88%EC%9D%98_%EA%B5%AC%EC%A1%B0" title="단백질의 구조 – Korean" lang="ko" hreflang="ko" data-title="단백질의 구조" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8D%D5%BA%D5%AB%D5%BF%D5%A1%D5%AF%D5%B8%D6%82%D6%81%D5%AB_%D5%AF%D5%A1%D5%BC%D5%B8%D6%82%D6%81%D5%BE%D5%A1%D5%AE%D6%84" title="Սպիտակուցի կառուցվածք – Armenian" lang="hy" hreflang="hy" data-title="Սպիտակուցի կառուցվածք" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A5%8D%E0%A4%B0%E0%A5%8B%E0%A4%9F%E0%A5%80%E0%A4%A8_%E0%A4%B8%E0%A4%82%E0%A4%B0%E0%A4%9A%E0%A4%A8%E0%A4%BE" title="प्रोटीन संरचना – Hindi" lang="hi" hreflang="hi" data-title="प्रोटीन संरचना" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Struktur_protein" title="Struktur protein – Indonesian" lang="id" hreflang="id" data-title="Struktur protein" 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/Struttura_proteica" title="Struttura proteica – Italian" lang="it" hreflang="it" data-title="Struttura proteica" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%AA%E1%83%98%E1%83%9A%E1%83%98%E1%83%A1_%E1%83%A1%E1%83%A2%E1%83%A0%E1%83%A3%E1%83%A5%E1%83%A2%E1%83%A3%E1%83%A0%E1%83%90" title="ცილის სტრუქტურა – Georgian" lang="ka" hreflang="ka" data-title="ცილის სტრუქტურა" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%A1%D1%82%D1%80%D1%83%D0%BA%D1%82%D1%83%D1%80%D0%B0_%D0%BD%D0%B0_%D0%B1%D0%B5%D0%BB%D0%BA%D0%BE%D0%B2%D0%B8%D0%BD%D0%B8%D1%82%D0%B5" title="Структура на белковините – Macedonian" lang="mk" hreflang="mk" data-title="Структура на белковините" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%A3%D1%83%D1%80%D0%B3%D0%B8%D0%B9%D0%BD_%D0%B1%D2%AF%D1%82%D1%8D%D1%86" title="Уургийн бүтэц – Mongolian" lang="mn" hreflang="mn" data-title="Уургийн бүтэц" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Eiwitstructuur" title="Eiwitstructuur – Dutch" lang="nl" hreflang="nl" data-title="Eiwitstructuur" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%BF%E3%83%B3%E3%83%91%E3%82%AF%E8%B3%AA%E6%A7%8B%E9%80%A0" title="タンパク質構造 – Japanese" lang="ja" hreflang="ja" data-title="タンパク質構造" 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-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Estructura_dei_prote%C3%AFnas" title="Estructura dei proteïnas – Occitan" lang="oc" hreflang="oc" data-title="Estructura dei proteïnas" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target"><span>Occitan</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Estrutura_da_prote%C3%ADna" title="Estrutura da proteína – Portuguese" lang="pt" hreflang="pt" data-title="Estrutura da proteína" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Structura_proteinelor" title="Structura proteinelor – Romanian" lang="ro" hreflang="ro" data-title="Structura proteinelor" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A1%D1%82%D1%80%D1%83%D0%BA%D1%82%D1%83%D1%80%D0%B0_%D0%B1%D0%B5%D0%BB%D0%BA%D0%BE%D0%B2" title="Структура белков – Russian" lang="ru" hreflang="ru" data-title="Структура белков" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Struktura_e_proteinave" title="Struktura e proteinave – Albanian" lang="sq" hreflang="sq" data-title="Struktura e proteinave" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Protein_structure" title="Protein structure – Simple English" lang="en-simple" hreflang="en-simple" data-title="Protein structure" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A1%D1%82%D1%80%D1%83%D0%BA%D1%82%D1%83%D1%80%D0%B0_%D0%BF%D1%80%D0%BE%D1%82%D0%B5%D0%B8%D0%BD%D0%B0" title="Структура протеина – Serbian" lang="sr" hreflang="sr" data-title="Структура протеина" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Proteinska_struktura" title="Proteinska struktura – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Proteinska struktura" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Proteiinin_rakenne" title="Proteiinin rakenne – Finnish" lang="fi" hreflang="fi" data-title="Proteiinin rakenne" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Proteinstruktur" title="Proteinstruktur – Swedish" lang="sv" hreflang="sv" data-title="Proteinstruktur" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%AA%E0%AF%81%E0%AE%B0%E0%AE%A4%E0%AE%95%E0%AF%8D_%E0%AE%95%E0%AE%9F%E0%AF%8D%E0%AE%9F%E0%AE%AE%E0%AF%88%E0%AE%AA%E0%AF%8D%E0%AE%AA%E0%AF%81" title="புரதக் கட்டமைப்பு – Tamil" lang="ta" hreflang="ta" data-title="புரதக் கட்டமைப்பு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Protein_yap%C4%B1s%C4%B1" title="Protein yapısı – Turkish" lang="tr" hreflang="tr" data-title="Protein yapısı" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D1%82%D1%80%D1%83%D0%BA%D1%82%D1%83%D1%80%D0%B0_%D0%B1%D1%96%D0%BB%D0%BA%D1%96%D0%B2" title="Структура білків – Ukrainian" lang="uk" hreflang="uk" 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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">Three-dimensional arrangement of atoms in an amino acid-chain molecule</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-More_citations_needed plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Protein_structure" title="Special:EditPage/Protein structure">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Protein+structure%22">"Protein structure"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Protein+structure%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Protein+structure%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Protein+structure%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Protein+structure%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Protein+structure%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">May 2018</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure class="noresize" typeof="mw:File/Thumb"><span><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/Protein_structure_%28full%29-en.svg/270px-Protein_structure_%28full%29-en.svg.png" decoding="async" width="270" height="439" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/77/Protein_structure_%28full%29-en.svg/405px-Protein_structure_%28full%29-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/77/Protein_structure_%28full%29-en.svg/540px-Protein_structure_%28full%29-en.svg.png 2x" data-file-width="2950" data-file-height="4800" usemap="#ImageMap_7475d4ee38a50cbd" resource="/wiki/File:Protein_structure_(full)-en.svg" /></span><map name="ImageMap_7475d4ee38a50cbd"><area href="/wiki/Protein_primary_structure" shape="rect" coords="0,0,288,64" alt="Protein primary structure" title="Protein primary structure" /><area href="/wiki/Protein_secondary_structure" shape="rect" coords="0,64,288,130" alt="Protein secondary structure" title="Protein secondary structure" /><area href="/wiki/Protein_tertiary_structure" shape="rect" coords="0,130,288,230" alt="Protein tertiary structure" title="Protein tertiary structure" /><area href="/wiki/Protein_quaternary_structure" shape="rect" coords="0,230,288,470" alt="Protein quaternary structure" title="Protein quaternary structure" /></map><figcaption><figure class="mw-halign-left" typeof="mw:File"><span title="The image above contains clickable links"><img alt="The image above contains clickable links" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/18px-Interactive_icon.svg.png" decoding="async" width="18" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/27px-Interactive_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Interactive_icon.svg/36px-Interactive_icon.svg.png 2x" data-file-width="133" data-file-height="200" /></span><figcaption>The image above contains clickable links</figcaption></figure> <a href="https://commons.wikimedia.org/wiki/File:Protein_structure_(full)-en.svg" class="extiw" title="c:File:Protein structure (full)-en.svg">This diagram</a> (which is interactive) of <a class="mw-selflink selflink">protein structure</a> uses <a href="/wiki/PCNA" class="mw-redirect" title="PCNA">PCNA</a> as an example. (<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/1AXC">1AXC</a></span>)</figcaption></figure> <p><b>Protein structure</b> is the <a href="/wiki/Molecular_geometry" title="Molecular geometry">three-dimensional arrangement of atoms</a> in an <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a>-chain <a href="/wiki/Molecule" title="Molecule">molecule</a>. <a href="/wiki/Protein" title="Protein">Proteins</a> are <a href="/wiki/Polymer" title="Polymer">polymers</a> –  specifically <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptides</a> –  formed from sequences of <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a>, which are the <a href="/wiki/Monomer" title="Monomer">monomers</a> of the polymer. A single amino acid monomer may also be called a <i>residue</i>, which indicates a repeating unit of a polymer. Proteins form by amino acids undergoing <a href="/wiki/Condensation_reaction" title="Condensation reaction">condensation reactions</a>, in which the amino acids lose one <a href="/wiki/Water_molecule" class="mw-redirect" title="Water molecule">water molecule</a> per <a href="/wiki/Chemical_reaction" title="Chemical reaction">reaction</a> in order to attach to one another with a <a href="/wiki/Peptide_bond" title="Peptide bond">peptide bond</a>. By convention, a chain under 30 amino acids is often identified as a <a href="/wiki/Peptide" title="Peptide">peptide</a>, rather than a protein.<sup id="cite_ref-Stoker2015_1-0" class="reference"><a href="#cite_note-Stoker2015-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> To be able to perform their biological function, proteins fold into one or more specific spatial conformations driven by a number of <a href="/wiki/Non-covalent_interaction" title="Non-covalent interaction">non-covalent interactions</a>, such as <a href="/wiki/Hydrogen_bonding" class="mw-redirect" title="Hydrogen bonding">hydrogen bonding</a>, <a href="/wiki/Ionic_interaction" class="mw-redirect" title="Ionic interaction">ionic interactions</a>, <a href="/wiki/Van_der_Waals_forces" class="mw-redirect" title="Van der Waals forces">Van der Waals forces</a>, and <a href="/wiki/Hydrophobic" class="mw-redirect" title="Hydrophobic">hydrophobic</a> packing. To understand the functions of proteins at a molecular level, it is often necessary to determine their <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">three-dimensional structure</a>. This is the topic of the scientific field of <a href="/wiki/Structural_biology" title="Structural biology">structural biology</a>, which employs techniques such as <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>, <a href="/wiki/Protein_NMR" class="mw-redirect" title="Protein NMR">NMR spectroscopy</a>, <a href="/wiki/Cryogenic_electron_microscopy" title="Cryogenic electron microscopy">cryo-electron microscopy (cryo-EM)</a> and <a href="/wiki/Dual_polarisation_interferometry" class="mw-redirect" title="Dual polarisation interferometry">dual polarisation interferometry</a>, to determine the structure of proteins. </p><p>Protein structures range in size from tens to several thousand amino acids.<sup id="cite_ref-Brocchieri2005_2-0" class="reference"><a href="#cite_note-Brocchieri2005-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> By physical size, proteins are classified as <a href="/wiki/Nanoparticle" title="Nanoparticle">nanoparticles</a>, between 1–100 nm. Very large <a href="/wiki/Protein_complexes" class="mw-redirect" title="Protein complexes">protein complexes</a> can be formed from <a href="/wiki/Protein_subunit" title="Protein subunit">protein subunits</a>. For example, many thousands of <a href="/wiki/Actin" title="Actin">actin</a> molecules assemble into a <a href="/wiki/Microfilament" title="Microfilament">microfilament</a>. </p><p>A protein usually undergoes <a href="/wiki/Reversible_process_(thermodynamics)" title="Reversible process (thermodynamics)">reversible</a> <a href="/wiki/Conformational_change" title="Conformational change">structural changes</a> in performing its biological function. The alternative structures of the same protein are referred to as different <a href="/wiki/Conformational_isomerism" title="Conformational isomerism">conformations</a>, and transitions between them are called <a href="/wiki/Conformational_change" title="Conformational change">conformational changes</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Levels_of_protein_structure">Levels of protein structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=1" title="Edit section: Levels of protein structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are four distinct levels of protein structure. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Protein_structure.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Protein_structure.png/400px-Protein_structure.png" decoding="async" width="400" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Protein_structure.png/600px-Protein_structure.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/Protein_structure.png/800px-Protein_structure.png 2x" data-file-width="1344" data-file-height="1008" /></a><figcaption>Four levels of protein structure</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Primary_structure">Primary structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=2" title="Edit section: Primary structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_primary_structure" title="Protein primary structure">Protein primary structure</a></div> <p>The <a href="/wiki/Primary_structure" class="mw-redirect" title="Primary structure">primary structure</a> of a protein refers to the sequence of <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> in the polypeptide chain. The primary structure is held together by <a href="/wiki/Peptide_bonds" class="mw-redirect" title="Peptide bonds">peptide bonds</a> that are made during the process of <a href="/wiki/Protein_biosynthesis" title="Protein biosynthesis">protein biosynthesis</a>. The two ends of the <a href="/wiki/Polypeptide_chain" class="mw-redirect" title="Polypeptide chain">polypeptide chain</a> are referred to as the <a href="/wiki/Carboxyl_terminus" class="mw-redirect" title="Carboxyl terminus">carboxyl terminus</a> (C-terminus) and the <a href="/wiki/Amino_terminus" class="mw-redirect" title="Amino terminus">amino terminus</a> (N-terminus) based on the nature of the free group on each extremity. Counting of residues always starts at the N-terminal end (NH<sub>2</sub>-group), which is the end where the amino group is not involved in a peptide bond. The primary structure of a protein is determined by the <a href="/wiki/Gene" title="Gene">gene</a> corresponding to the protein. A specific sequence of <a href="/wiki/Nucleotide" title="Nucleotide">nucleotides</a> in <a href="/wiki/DNA" title="DNA">DNA</a> is <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a> into <a href="/wiki/MRNA" class="mw-redirect" title="MRNA">mRNA</a>, which is read by the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> in a process called <a href="/wiki/Translation_(biology)" title="Translation (biology)">translation</a>. The sequence of amino acids in insulin was discovered by <a href="/wiki/Frederick_Sanger" title="Frederick Sanger">Frederick Sanger</a>, establishing that proteins have defining amino acid sequences.<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><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> The sequence of a protein is unique to that protein, and defines the structure and function of the protein. The sequence of a protein can be determined by methods such as <a href="/wiki/Edman_degradation" title="Edman degradation">Edman degradation</a> or <a href="/wiki/Mass_spectrometry#Protein_identification" title="Mass spectrometry">tandem mass spectrometry</a>. Often, however, it is read directly from the sequence of the gene using the <a href="/wiki/Genetic_code" title="Genetic code">genetic code</a>. It is strictly recommended to use the words "amino acid residues" when discussing proteins because when a peptide bond is formed, a <a href="/wiki/Water_molecule" class="mw-redirect" title="Water molecule">water molecule</a> is lost, and therefore proteins are made up of amino acid residues. <a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modifications</a> such as <a href="/wiki/Phosphorylation" title="Phosphorylation">phosphorylations</a> and <a href="/wiki/Glycosylation" title="Glycosylation">glycosylations</a> are usually also considered a part of the primary structure, and cannot be read from the gene. For example, <a href="/wiki/Insulin" title="Insulin">insulin</a> is composed of 51 amino acids in 2 chains. One chain has 31 amino acids, and the other has 20 amino acids. </p> <div class="mw-heading mw-heading3"><h3 id="Secondary_structure">Secondary structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=3" title="Edit section: Secondary structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Alpha_helix.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Alpha_helix.png/100px-Alpha_helix.png" decoding="async" width="100" height="232" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/Alpha_helix.png/150px-Alpha_helix.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/75/Alpha_helix.png/200px-Alpha_helix.png 2x" data-file-width="287" data-file-height="666" /></a><figcaption>An α-helix with hydrogen bonds (yellow dots)</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_secondary_structure" title="Protein secondary structure">Protein secondary structure</a></div> <p><a href="/wiki/Secondary_structure" class="mw-redirect" title="Secondary structure">Secondary structure</a> refers to highly regular local sub-structures on the actual polypeptide backbone chain. Two main types of secondary structure, the <a href="/wiki/Alpha_helix" title="Alpha helix">α-helix</a> and the <a href="/wiki/Beta_strand" class="mw-redirect" title="Beta strand">β-strand</a> or <a href="/wiki/Beta_sheet" title="Beta sheet">β-sheets</a>, were suggested in 1951 by <a href="/wiki/Linus_Pauling" title="Linus Pauling">Linus Pauling</a>.<sup id="cite_ref-Pauling1951_5-0" class="reference"><a href="#cite_note-Pauling1951-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> These secondary structures are defined by patterns of <a href="/wiki/Hydrogen_bonds" class="mw-redirect" title="Hydrogen bonds">hydrogen bonds</a> between the main-chain peptide groups. They have a regular geometry, being constrained to specific values of the dihedral angles ψ and φ on the <a href="/wiki/Ramachandran_plot" title="Ramachandran plot">Ramachandran plot</a>. Both the α-helix and the β-sheet represent a way of saturating all the hydrogen bond donors and acceptors in the peptide backbone. Some parts of the protein are ordered but do not form any regular structures. They should not be confused with <a href="/wiki/Random_coil" title="Random coil">random coil</a>, an unfolded polypeptide chain lacking any fixed three-dimensional structure. Several sequential secondary structures may form a "<a href="/wiki/Supersecondary_structure" title="Supersecondary structure">supersecondary unit</a>".<sup id="cite_ref-ChiangYS2007_6-0" class="reference"><a href="#cite_note-ChiangYS2007-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tertiary_structure">Tertiary structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=4" title="Edit section: Tertiary structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">Protein tertiary structure</a></div> <p><a href="/wiki/Tertiary_structure" class="mw-redirect" title="Tertiary structure">Tertiary structure</a> refers to the three-dimensional structure created by a single protein molecule (a single <a href="/wiki/Polypeptide_chain" class="mw-redirect" title="Polypeptide chain">polypeptide chain</a>). It may include <a href="/wiki/Protein_domain" title="Protein domain">one or several domains</a>. The α-helices and β-pleated-sheets are folded into a compact <a href="/wiki/Globular_structure" class="mw-redirect" title="Globular structure">globular structure</a>. The folding is driven by the <i>non-specific</i> <a href="/wiki/Hydrophobic_interactions" class="mw-redirect" title="Hydrophobic interactions">hydrophobic interactions</a>, the burial of <a href="/wiki/Hydrophobic_residues" class="mw-redirect" title="Hydrophobic residues">hydrophobic residues</a> from <a href="/wiki/Water" title="Water">water</a>, but the structure is stable only when the parts of a <a href="/wiki/Protein_domain" title="Protein domain">protein domain</a> are locked into place by <i>specific</i> tertiary interactions, such as <a href="/wiki/Salt_bridge_(protein_and_supramolecular)" title="Salt bridge (protein and supramolecular)">salt bridges</a>, hydrogen bonds, and the tight packing of side chains and <a href="/wiki/Disulfide_bond" class="mw-redirect" title="Disulfide bond">disulfide bonds</a>. The disulfide bonds are extremely rare in cytosolic proteins, since the <a href="/wiki/Cytosol" title="Cytosol">cytosol</a> (intracellular fluid) is generally a <a href="/wiki/Redox" title="Redox">reducing</a> environment. </p> <div class="mw-heading mw-heading3"><h3 id="Quaternary_structure">Quaternary structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=5" title="Edit section: Quaternary structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_quaternary_structure" title="Protein quaternary structure">Protein quaternary structure</a></div> <p>Quaternary structure is the three-dimensional structure consisting of the aggregation of two or more individual polypeptide chains (subunits) that operate as a single functional unit (<a href="/wiki/Multimer" class="mw-redirect" title="Multimer">multimer</a>). The resulting multimer is stabilized by the same <a href="/wiki/Non-covalent_interaction" title="Non-covalent interaction">non-covalent interactions</a> and disulfide bonds as in tertiary structure. There are many possible quaternary structure organisations.<sup id="cite_ref-pmid19059267_7-0" class="reference"><a href="#cite_note-pmid19059267-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Complexes of two or more polypeptides (i.e. multiple subunits) are called <a href="/wiki/Multimer" class="mw-redirect" title="Multimer">multimers</a>. Specifically it would be called a <a href="/wiki/Dimer_(chemistry)" class="mw-redirect" title="Dimer (chemistry)">dimer</a> if it contains two subunits, a <a href="/wiki/Trimer_(chemistry)" title="Trimer (chemistry)">trimer</a> if it contains three subunits, a <a href="/wiki/Tetramer" title="Tetramer">tetramer</a> if it contains four subunits, and a <a href="/wiki/Pentamer" title="Pentamer">pentamer</a> if it contains five subunits, and so forth. The subunits are frequently related to one another by <a href="/wiki/Symmetry_group" title="Symmetry group">symmetry operations</a>, such as a 2-fold axis in a dimer. Multimers made up of identical subunits are referred to with a prefix of "homo-" and those made up of different subunits are referred to with a prefix of "hetero-", for example, a heterotetramer, such as the two alpha and two beta chains of <a href="/wiki/Hemoglobin" title="Hemoglobin">hemoglobin</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Homomers">Homomers</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=6" title="Edit section: Homomers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An assemblage of multiple copies of a particular <a href="/wiki/Polypeptide" class="mw-redirect" title="Polypeptide">polypeptide</a> chain can be described as a <a href="/wiki/Homomeric" title="Homomeric">homomer</a>, <a href="/wiki/Multimer" class="mw-redirect" title="Multimer">multimer</a> or <a href="/wiki/Oligomer" title="Oligomer">oligomer</a>. Bertolini et al. in 2021<sup id="cite_ref-Bertolini2021_8-0" class="reference"><a href="#cite_note-Bertolini2021-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> presented evidence that homomer formation may be driven by interaction between nascent polypeptide chains as they are translated from <a href="/wiki/Messenger_RNA" title="Messenger RNA">mRNA</a> by nearby adjacent <a href="/wiki/Ribosome" title="Ribosome">ribosomes</a>. Hundreds of proteins have been identified as being assembled into homomers in human cells.<sup id="cite_ref-Bertolini2021_8-1" class="reference"><a href="#cite_note-Bertolini2021-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The process of assembly is often initiated by the interaction of the N-terminal region of polypeptide chains. Evidence that numerous gene products form homomers (multimers) in a variety of organisms based on <a href="/wiki/Complementation_(genetics)" title="Complementation (genetics)">intragenic complementation</a> evidence was reviewed in 1965.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Domains,_motifs,_and_folds_in_protein_structure"><span id="Domains.2C_motifs.2C_and_folds_in_protein_structure"></span>Domains, motifs, and folds in protein structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=7" title="Edit section: Domains, motifs, and folds in protein structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Domain_Homology.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Domain_Homology.png/220px-Domain_Homology.png" decoding="async" width="220" height="103" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Domain_Homology.png/330px-Domain_Homology.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/19/Domain_Homology.png/440px-Domain_Homology.png 2x" data-file-width="929" data-file-height="435" /></a><figcaption>Protein domains. The two shown protein structures share a common domain (maroon), the <a href="/wiki/Pleckstrin_homology_domain" title="Pleckstrin homology domain">PH domain</a>, which is involved in <a href="/wiki/Phosphatidylinositol_(3,4,5)-trisphosphate" title="Phosphatidylinositol (3,4,5)-trisphosphate">phosphatidylinositol (3,4,5)-trisphosphate</a> binding</figcaption></figure> <p>Proteins are frequently described as consisting of several structural units. These units include domains, <a href="/wiki/Structural_motif#In_proteins" title="Structural motif">motifs</a>, and folds. Despite the fact that there are about 100,000 different proteins expressed in <a href="/wiki/Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> systems, there are many fewer different domains, structural motifs and folds. </p> <div class="mw-heading mw-heading3"><h3 id="Structural_domain">Structural domain</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=8" title="Edit section: Structural domain"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A <a href="/wiki/Structural_domain" class="mw-redirect" title="Structural domain">structural domain</a> is an element of the protein's overall structure that is self-stabilizing and often <a href="/wiki/Protein_folding" title="Protein folding">folds</a> independently of the rest of the protein chain. Many domains are not unique to the protein products of one <a href="/wiki/Gene" title="Gene">gene</a> or one <a href="/wiki/Gene_family" title="Gene family">gene family</a> but instead appear in a variety of proteins. Domains often are named and singled out because they figure prominently in the biological function of the protein they belong to; for example, the "<a href="/wiki/Calcium" title="Calcium">calcium</a>-binding domain of <a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a>". Because they are independently stable, domains can be "swapped" by <a href="/wiki/Genetic_engineering" title="Genetic engineering">genetic engineering</a> between one protein and another to make <a href="/wiki/Chimera_(protein)" class="mw-redirect" title="Chimera (protein)">chimera</a> proteins. A conservative combination of several domains that occur in different proteins, such as <a href="/wiki/Protein_tyrosine_phosphatase" title="Protein tyrosine phosphatase">protein tyrosine phosphatase</a> domain and <a href="/wiki/C2_domain" title="C2 domain">C2 domain</a> pair, was called "a superdomain" that may evolve as a single unit.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Structural_and_sequence_motifs">Structural and sequence motifs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=9" title="Edit section: Structural and sequence motifs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Structural_motif" title="Structural motif">structural</a> and <a href="/wiki/Sequence_motif" title="Sequence motif">sequence motifs</a> refer to short segments of protein three-dimensional structure or amino acid sequence that were found in a large number of different proteins </p> <div class="mw-heading mw-heading3"><h3 id="Supersecondary_structure">Supersecondary structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=10" title="Edit section: Supersecondary structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Tertiary protein structures can have multiple secondary elements on the same polypeptide chain. The <a href="/wiki/Supersecondary_structure" title="Supersecondary structure">supersecondary structure</a> refers to a specific combination of <a href="/wiki/Secondary_structure" class="mw-redirect" title="Secondary structure">secondary structure</a> elements, such as β-α-β units or a <a href="/wiki/Helix-turn-helix" title="Helix-turn-helix">helix-turn-helix</a> motif. Some of them may be also referred to as structural motifs. </p> <div class="mw-heading mw-heading3"><h3 id="Protein_fold">Protein fold</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=11" title="Edit section: Protein fold"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A protein fold refers to the general protein architecture, like a <a href="/wiki/Helix_bundle" title="Helix bundle">helix bundle</a>, <a href="/wiki/Beta_barrel" title="Beta barrel">β-barrel</a>, <a href="/wiki/Rossmann_fold" title="Rossmann fold">Rossmann fold</a> or different "folds" provided in the <a href="/wiki/Structural_Classification_of_Proteins_database" title="Structural Classification of Proteins database">Structural Classification of Proteins database</a>.<sup id="cite_ref-Govinda_rajan_11-0" class="reference"><a href="#cite_note-Govinda_rajan-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> A related concept is <a href="/wiki/Protein_topology" title="Protein topology">protein topology</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Protein_dynamics_and_conformational_ensembles">Protein dynamics and conformational ensembles</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=12" title="Edit section: Protein dynamics and conformational ensembles"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_dynamics" title="Protein dynamics">Protein dynamics</a></div> <p>Proteins are not static objects, but rather populate ensembles of <a href="/wiki/Conformational_change" title="Conformational change">conformational states</a>. Transitions between these states typically occur on <a href="/wiki/Nanoscopic_scale" class="mw-redirect" title="Nanoscopic scale">nanoscales</a>, and have been linked to functionally relevant phenomena such as <a href="/wiki/Allosteric_regulation" title="Allosteric regulation">allosteric signaling</a><sup id="cite_ref-pmid21570668_12-0" class="reference"><a href="#cite_note-pmid21570668-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Enzyme_catalysis" title="Enzyme catalysis">enzyme catalysis</a>.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Protein_dynamics" title="Protein dynamics">Protein dynamics</a> and <a href="/wiki/Conformational_change" title="Conformational change">conformational changes</a> allow proteins to function as nanoscale <a href="/wiki/Biological_machine" class="mw-redirect" title="Biological machine">biological machines</a> within cells, often in the form of <a href="/wiki/Protein_complex" title="Protein complex">multi-protein complexes</a>.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Examples include <a href="/wiki/Motor_proteins" class="mw-redirect" title="Motor proteins">motor proteins</a>, such as <a href="/wiki/Myosin" title="Myosin">myosin</a>, which is responsible for <a href="/wiki/Muscle" title="Muscle">muscle</a> contraction, <a href="/wiki/Kinesin" title="Kinesin">kinesin</a>, which moves cargo inside cells away from the <a href="/wiki/Cell_nucleus" title="Cell nucleus">nucleus</a> along <a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubules</a>, and <a href="/wiki/Dynein" title="Dynein">dynein</a>, which moves cargo inside cells towards the nucleus and produces the axonemal beating of <a href="/wiki/Cilia#Motile_cilia" class="mw-redirect" title="Cilia">motile cilia</a> and <a href="/wiki/Flagella" class="mw-redirect" title="Flagella">flagella</a>. "[I]n effect, the [motile cilium] is a nanomachine composed of perhaps over 600 proteins in molecular complexes, many of which also function independently as nanomachines...<a href="/wiki/Flexible_linker" class="mw-redirect" title="Flexible linker">Flexible linkers</a> allow the <a href="/wiki/Protein_domain#Domains_and_protein_flexibility" title="Protein domain">mobile protein domains</a> connected by them to recruit their binding partners and induce long-range <a href="/wiki/Allostery" class="mw-redirect" title="Allostery">allostery</a> via <a href="/wiki/Protein_dynamics#Global_flexibility:_multiple_domains" title="Protein dynamics">protein domain dynamics</a>. "<sup id="cite_ref-Satir2008_15-0" class="reference"><a href="#cite_note-Satir2008-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg/500px-Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg" decoding="async" width="500" height="289" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg/750px-Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/0/08/Schematic_view_of_the_two_main_ensemble_modeling_approaches.jpg 2x" data-file-width="851" data-file-height="492" /></a><figcaption>Schematic view of the two main ensemble modeling approaches.<sup id="cite_ref-:2_16-0" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Proteins are often thought of as relatively stable <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">tertiary structures</a> that experience conformational changes after being affected by interactions with other proteins or as a part of enzymatic activity. However, proteins may have varying degrees of stability, and some of the less stable variants are <a href="/wiki/Intrinsically_disordered_proteins" title="Intrinsically disordered proteins">intrinsically disordered proteins</a>. These proteins exist and function in a relatively 'disordered' state lacking a stable <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">tertiary structure</a>. As a result, they are difficult to describe by a single fixed <a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">tertiary structure</a>. <a href="/wiki/Conformational_ensembles" title="Conformational ensembles">Conformational ensembles</a> have been devised as a way to provide a more accurate and 'dynamic' representation of the conformational state of <a href="/wiki/Intrinsically_disordered_proteins" title="Intrinsically disordered proteins">intrinsically disordered proteins</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_16-1" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Protein <a href="/wiki/Conformational_ensembles" title="Conformational ensembles">ensemble</a> files are a representation of a protein that can be considered to have a flexible structure. Creating these files requires determining which of the various theoretically possible protein conformations actually exist. One approach is to apply computational algorithms to the protein data in order to try to determine the most likely set of conformations for an <a href="/wiki/Conformational_ensembles" title="Conformational ensembles">ensemble</a> file. There are multiple methods for preparing data for the <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180310010556/http://pedb.vib.be/">Protein Ensemble Database</a> that fall into two general methodologies – pool and molecular dynamics (MD) approaches (diagrammed in the figure). The pool based approach uses the protein's amino acid sequence to create a massive pool of random conformations. This pool is then subjected to more computational processing that creates a set of theoretical parameters for each conformation based on the structure. Conformational subsets from this pool whose average theoretical parameters closely match known experimental data for this protein are selected. The alternative molecular dynamics approach takes multiple random conformations at a time and subjects all of them to experimental data. Here the experimental data is serving as limitations to be placed on the conformations (e.g. known distances between atoms). Only conformations that manage to remain within the limits set by the experimental data are accepted. This approach often applies large amounts of experimental data to the conformations which is a very computationally demanding task.<sup id="cite_ref-:2_16-2" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>The conformational ensembles were generated for a number of highly dynamic and partially unfolded proteins, such as <a href="/wiki/Sic1" title="Sic1">Sic1</a>/<a href="/wiki/Cell_division_control_protein_4" title="Cell division control protein 4">Cdc4</a>,<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> <a href="/wiki/KIAA0101" title="KIAA0101">p15 PAF</a>,<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> <a href="/wiki/MAP2K7" title="MAP2K7">MKK7</a>,<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Beta-synuclein" title="Beta-synuclein">Beta-synuclein</a><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/CDKN1B" title="CDKN1B">P27</a><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Protein_folding">Protein folding</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=13" title="Edit section: Protein folding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-Expand_section plainlinks metadata ambox mbox-small-left ambox-content" role="presentation"><tbody><tr><td class="mbox-image"><span typeof="mw:File"><a href="/wiki/File:Wiki_letter_w_cropped.svg" class="mw-file-description"><img alt="[icon]" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1c/Wiki_letter_w_cropped.svg/20px-Wiki_letter_w_cropped.svg.png" decoding="async" width="20" height="14" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1c/Wiki_letter_w_cropped.svg/30px-Wiki_letter_w_cropped.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1c/Wiki_letter_w_cropped.svg/40px-Wiki_letter_w_cropped.svg.png 2x" data-file-width="44" data-file-height="31" /></a></span></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs expansion</b>. You can help by <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Protein_structure&action=edit&section=">adding to it</a>. <span class="date-container"><i>(<span class="date">April 2019</span>)</i></span></div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_folding" title="Protein folding">Protein folding</a></div> <p>As it is translated, polypeptides exit the <a href="/wiki/Ribosome" title="Ribosome">ribosome</a> mostly as a <a href="/wiki/Random_coil" title="Random coil">random coil</a> and folds into its <a href="/wiki/Native_state" title="Native state">native state</a>.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Alberts_24-0" class="reference"><a href="#cite_note-Alberts-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> The final structure of the protein chain is generally assumed to be determined by its amino acid sequence (<a href="/wiki/Anfinsen%27s_dogma" title="Anfinsen's dogma">Anfinsen's dogma</a>).<sup id="cite_ref-Anfinsen_25-0" class="reference"><a href="#cite_note-Anfinsen-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Protein_stability">Protein stability</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=14" title="Edit section: Protein stability"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Equilibrium_unfolding" title="Equilibrium unfolding">Equilibrium unfolding</a></div> <p>Thermodynamic stability of proteins represents the <a href="/wiki/Gibbs_free_energy" title="Gibbs free energy">free energy difference</a> between the folded and <a href="/wiki/Denaturation_(biochemistry)" title="Denaturation (biochemistry)">unfolded</a> protein states. This free energy difference is very sensitive to temperature, hence a change in temperature may result in unfolding or denaturation. <a href="/wiki/Denaturation_(biochemistry)" title="Denaturation (biochemistry)">Protein denaturation</a> may result in loss of function, and loss of native state. The free energy of stabilization of soluble globular proteins typically does not exceed 50 kJ/mol.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2018)">citation needed</span></a></i>]</sup> Taking into consideration the large number of hydrogen bonds that take place for the stabilization of secondary structures, and the stabilization of the inner core through hydrophobic interactions, the free energy of stabilization emerges as small difference between large numbers.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Protein_structure_determination">Protein structure determination</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=15" title="Edit section: Protein structure determination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Protein_structure_examples.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Protein_structure_examples.png/220px-Protein_structure_examples.png" decoding="async" width="220" height="271" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/24/Protein_structure_examples.png/330px-Protein_structure_examples.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/24/Protein_structure_examples.png/440px-Protein_structure_examples.png 2x" data-file-width="1571" data-file-height="1938" /></a><figcaption>Examples of protein structures from the <a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">PDB</a> </figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Rate_of_Protein_Structure_Determination-2014.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Rate_of_Protein_Structure_Determination-2014.png/400px-Rate_of_Protein_Structure_Determination-2014.png" decoding="async" width="400" height="286" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Rate_of_Protein_Structure_Determination-2014.png/600px-Rate_of_Protein_Structure_Determination-2014.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/10/Rate_of_Protein_Structure_Determination-2014.png/800px-Rate_of_Protein_Structure_Determination-2014.png 2x" data-file-width="3500" data-file-height="2500" /></a><figcaption>Rate of Protein Structure Determination by Method and Year</figcaption></figure> <p>Around 90% of the protein structures available in the <a href="/wiki/Protein_Data_Bank" title="Protein Data Bank">Protein Data Bank</a> have been determined by <a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a>.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> This method allows one to measure the three-dimensional (3-D) density distribution of <a href="/wiki/Electron" title="Electron">electrons</a> in the protein, in the <a href="/wiki/Crystallized" class="mw-redirect" title="Crystallized">crystallized</a> state, and thereby <a href="/wiki/Infer" class="mw-redirect" title="Infer">infer</a> the 3-D coordinates of all the <a href="/wiki/Atom" title="Atom">atoms</a> to be determined to a certain resolution. Roughly 7% of the known protein structures have been obtained by <a href="/wiki/Protein_NMR" class="mw-redirect" title="Protein NMR">nuclear magnetic resonance</a> (NMR) techniques.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> For larger protein complexes, <a href="/wiki/Cryo-electron_microscopy" class="mw-redirect" title="Cryo-electron microscopy">cryo-electron microscopy</a> can determine protein structures. The resolution is typically lower than that of X-ray crystallography, or NMR, but the maximum resolution is steadily increasing. This technique is still a particularly valuable for very large protein complexes such as <a href="/wiki/Virus_coat_protein" class="mw-redirect" title="Virus coat protein">virus coat proteins</a> and <a href="/wiki/Amyloid" title="Amyloid">amyloid</a> fibers. </p><p>General secondary structure composition can be determined via <a href="/wiki/Circular_dichroism" title="Circular dichroism">circular dichroism</a>. <a href="/wiki/Vibrational_spectroscopy" class="mw-redirect" title="Vibrational spectroscopy">Vibrational spectroscopy</a> can also be used to characterize the conformation of peptides, polypeptides, and proteins.<sup id="cite_ref-pmid3541539_29-0" class="reference"><a href="#cite_note-pmid3541539-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Two-dimensional_infrared_spectroscopy" title="Two-dimensional infrared spectroscopy">Two-dimensional infrared spectroscopy</a> has become a valuable method to investigate the structures of flexible peptides and proteins that cannot be studied with other methods.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> A more qualitative picture of protein structure is often obtained by <a href="/wiki/Proteolysis" title="Proteolysis">proteolysis</a>, which is also useful to screen for more crystallizable protein samples. Novel implementations of this approach, including <a href="/wiki/Fast_parallel_proteolysis_(FASTpp)" class="mw-redirect" title="Fast parallel proteolysis (FASTpp)">fast parallel proteolysis (FASTpp)</a>, can probe the structured fraction and its stability without the need for purification.<sup id="cite_ref-pmid23056252_32-0" class="reference"><a href="#cite_note-pmid23056252-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Once a protein's structure has been experimentally determined, further detailed studies can be done computationally, using <a href="/wiki/Molecular_dynamics" title="Molecular dynamics">molecular dynamic</a> simulations of that structure.<sup id="cite_ref-pmid28637405_33-0" class="reference"><a href="#cite_note-pmid28637405-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Protein_structure_databases">Protein structure databases</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=16" title="Edit section: Protein structure databases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A <a href="/wiki/Protein_structure_database" title="Protein structure database">protein structure database</a> is a database that is <a href="/wiki/Data_modeling" title="Data modeling">modeled</a> around the various <a href="#Protein_structure_determination">experimentally determined</a> protein structures. The aim of most protein structure databases is to organize and annotate the protein structures, providing the biological community access to the experimental data in a useful way. Data included in protein structure databases often includes 3D coordinates as well as experimental information, such as unit cell dimensions and angles for <a href="/wiki/X-ray_crystallography#Biological_macromolecular_crystallography" title="X-ray crystallography">x-ray crystallography</a> determined structures. Though most instances, in this case either proteins or a specific structure determinations of a protein, also contain sequence information and some databases even provide means for performing sequence based queries, the primary attribute of a structure database is structural information, whereas <a href="/wiki/Sequence_database" title="Sequence database">sequence databases</a> focus on sequence information, and contain no structural information for the majority of entries. Protein structure databases are critical for many efforts in <a href="/wiki/Computational_biology" title="Computational biology">computational biology</a> such as <a href="/wiki/Drug_design#Structure-based" title="Drug design">structure based drug design</a>, both in developing the computational methods used and in providing a large experimental dataset used by some methods to provide insights about the function of a protein.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Structural_classifications_of_proteins">Structural classifications of proteins</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=17" title="Edit section: Structural classifications of proteins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Protein structures can be grouped based on their structural similarity, <a href="/wiki/Circuit_topology" title="Circuit topology">topological class</a> or a common <a href="/wiki/Evolution" title="Evolution">evolutionary</a> origin. The <a href="/wiki/Structural_Classification_of_Proteins_database" title="Structural Classification of Proteins database">Structural Classification of Proteins database</a><sup id="cite_ref-pmid7723011_35-0" class="reference"><a href="#cite_note-pmid7723011-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/CATH" class="mw-redirect" title="CATH">CATH</a> database<sup id="cite_ref-pmid9309224_36-0" class="reference"><a href="#cite_note-pmid9309224-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> provide two different structural classifications of proteins. When the structural similarity is large the two proteins have possibly diverged from a common ancestor,<sup id="cite_ref-Pascual2009_37-0" class="reference"><a href="#cite_note-Pascual2009-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> and shared structure between proteins is considered evidence of <a href="/wiki/Homology_(biology)" title="Homology (biology)">homology</a>. Structure similarity can then be used to group proteins together into <a href="/wiki/Protein_superfamilies" class="mw-redirect" title="Protein superfamilies">protein superfamilies</a>.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> If shared structure is significant but the fraction shared is small, the fragment shared may be the consequence of a more dramatic evolutionary event such as <a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>, and joining proteins sharing these fragments into protein superfamilies is no longer justified.<sup id="cite_ref-Pascual2009_37-1" class="reference"><a href="#cite_note-Pascual2009-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Topology of a protein can be used to classify proteins as well. <a href="/wiki/Knot_theory" title="Knot theory">Knot theory</a> and <a href="/wiki/Circuit_topology" title="Circuit topology">circuit topology</a> are two topology frameworks developed for classification of protein folds based on chain crossing and intrachain contacts respectively. </p> <div class="mw-heading mw-heading2"><h2 id="Computational_prediction_of_protein_structure">Computational prediction of protein structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=18" title="Edit section: Computational prediction of protein structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Protein_structure_prediction" title="Protein structure prediction">Protein structure prediction</a></div> <p>The generation of a <a href="/wiki/Protein_sequence" class="mw-redirect" title="Protein sequence">protein sequence</a> is much easier than the determination of a protein structure. However, the structure of a protein gives much more insight in the function of the protein than its sequence. Therefore, a number of methods for the computational prediction of protein structure from its sequence have been developed.<sup id="cite_ref-zhang2008_39-0" class="reference"><a href="#cite_note-zhang2008-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> <i>Ab initio</i> prediction methods use just the sequence of the protein. <a href="/wiki/Threading_(protein_sequence)" title="Threading (protein sequence)">Threading</a> and <a href="/wiki/Homology_modeling" title="Homology modeling">homology modeling</a> methods can build a 3-D model for a protein of unknown structure from experimental structures of evolutionarily-related proteins, called a <a href="/wiki/Protein_family" title="Protein family">protein family</a>. </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=Protein_structure&action=edit&section=19" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Biomolecular_structure" title="Biomolecular structure">Biomolecular structure</a></li> <li><a href="/wiki/Gene_structure" title="Gene structure">Gene structure</a></li> <li><a href="/wiki/Nucleic_acid_structure" title="Nucleic acid structure">Nucleic acid structure</a></li> <li><a href="/wiki/PCRPi-DB" title="PCRPi-DB">PCRPi-DB</a></li> <li><a href="/wiki/Ribbon_diagram" title="Ribbon diagram">Ribbon diagram</a> 3D schematic representation of proteins</li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=20" 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 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href="https://pubmed.ncbi.nlm.nih.gov/20457744">20457744</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nucleic+Acids+Research&rft.atitle=Dali+server%3A+conservation+mapping+in+3D&rft.volume=38&rft.issue=Web+Server+issue&rft.pages=W545-W549&rft.date=2010-07&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2896194%23id-name%3DPMC&rft_id=info%3Apmid%2F20457744&rft_id=info%3Adoi%2F10.1093%2Fnar%2Fgkq366&rft.aulast=Holm&rft.aufirst=L&rft.au=Rosenstr%C3%B6m%2C+P&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2896194&rfr_id=info%3Asid%2Fen.wikipedia.org%3AProtein+structure" class="Z3988"></span></span> </li> <li id="cite_note-zhang2008-39"><span class="mw-cite-backlink"><b><a href="#cite_ref-zhang2008_39-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZhang2008" class="citation journal 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Opinion+in+Structural+Biology&rft.atitle=Progress+and+challenges+in+protein+structure+prediction&rft.volume=18&rft.issue=3&rft.pages=342-348&rft.date=2008-06&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2680823%23id-name%3DPMC&rft_id=info%3Apmid%2F18436442&rft_id=info%3Adoi%2F10.1016%2Fj.sbi.2008.02.004&rft.aulast=Zhang&rft.aufirst=Y&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2680823&rfr_id=info%3Asid%2Fen.wikipedia.org%3AProtein+structure" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Protein_structure&action=edit&section=21" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> 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modification</a></li> <li><a href="/wiki/Protein_folding" title="Protein folding">Protein folding</a></li> <li><a href="/wiki/Protein_targeting" title="Protein targeting">Protein targeting</a></li> <li><a href="/wiki/Proteome" title="Proteome">Proteome</a></li> <li><a href="/wiki/Protein_methods" title="Protein methods">Protein methods</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Structures</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 class="mw-selflink selflink">Protein structure</a></li> <li><a href="/wiki/Protein_domain" title="Protein domain">Protein structural domains</a></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</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" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/List_of_proteins" title="List of proteins">List of proteins</a></li> <li><a href="/wiki/Membrane_protein" title="Membrane protein">Membrane protein</a></li> <li><a href="/wiki/Globular_protein" title="Globular protein">Globular protein</a> <ul><li><a href="/wiki/Globulin" title="Globulin">Globulin</a></li> <li><a href="/wiki/Edestin" title="Edestin">Edestin</a></li> <li><a href="/wiki/Albumin" title="Albumin">Albumin</a></li></ul></li> <li><a href="/wiki/Scleroprotein" class="mw-redirect" title="Scleroprotein">Fibrous protein</a></li> <li><a href="/wiki/Chromoprotein" title="Chromoprotein">Chromoprotein</a></li> <li><a href="/wiki/Photoreceptor_protein" title="Photoreceptor protein">Photoreceptor protein</a></li> <li><a href="/wiki/Biliprotein" title="Biliprotein">Biliprotein</a> <ul><li><a href="/wiki/Phycobiliprotein" title="Phycobiliprotein">Phycobiliprotein</a></li> <li><a href="/wiki/Phytochrome" title="Phytochrome">Phytochrome</a></li> <li><a href="/wiki/Lipocalin" title="Lipocalin">Lipocalin</a></li></ul></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="Protein_domains" 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:Protein_domains" title="Template:Protein domains"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Protein_domains" title="Template talk:Protein domains"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Protein_domains" title="Special:EditPage/Template:Protein domains"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Protein_domains" style="font-size:114%;margin:0 4em"><a href="/wiki/Protein_domain" title="Protein domain">Protein domains</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/3H_domain" title="3H domain">3H</a></li> <li><a href="/wiki/ABM_domain" title="ABM domain">ABM</a></li> <li><a href="/wiki/ACDC_domain" title="ACDC domain">ACDC</a></li> <li><a href="/wiki/ACT_domain" title="ACT domain">ACT</a></li> <li><a href="/wiki/ADF-H_domain" title="ADF-H domain">ADF-H</a></li> <li><a href="/wiki/ANTH_domain" title="ANTH domain">ANTH</a></li> <li><a href="/wiki/ARID_domain" title="ARID domain">ARID</a></li> <li><a href="/wiki/BAR_domain" title="BAR domain">BAR</a></li> <li><a href="/wiki/BEN_domain" title="BEN domain">BEN</a></li> <li><a href="/wiki/BESS_domain" title="BESS domain">BESS</a></li> <li><a href="/wiki/Inhibitor_of_apoptosis_domain" title="Inhibitor of apoptosis domain">BIR</a></li> <li><a href="/wiki/BMC_domain" title="BMC domain">BMC</a></li> <li><a href="/wiki/BPS_domain" title="BPS domain">BPS</a></li> <li><a href="/wiki/BTB/POZ_domain" title="BTB/POZ domain">BTB/POZ</a></li> <li><a href="/wiki/BZIP_domain" title="BZIP domain">BZIP</a></li> <li><a href="/wiki/C1_domain" title="C1 domain">C1</a></li> <li><a href="/wiki/C2_domain" title="C2 domain">C2</a></li> <li><a href="/wiki/Cache_domain" title="Cache domain">Cache</a></li> <li><a href="/wiki/CBS_domain" title="CBS domain">CBS</a></li> <li><a href="/wiki/Kinase_binding_protein_CGI-121" title="Kinase binding protein CGI-121">CGI-121</a></li> <li><a href="/wiki/CRM_domain" title="CRM domain">CRM</a></li> <li><a href="/wiki/CUB_domain" title="CUB domain">CUB</a></li> <li><a href="/wiki/CUT_domain" title="CUT domain">CUT</a></li> <li><a href="/wiki/CVNH_domain" title="CVNH domain">CVHN</a></li> <li><a href="/wiki/Death_fold" title="Death fold">Death</a> <ul><li><a href="/wiki/Death_domain" title="Death domain">DD</a></li> <li><a href="/wiki/Death_effector_domain" title="Death effector domain">DED</a></li> <li><a href="/wiki/CARD_(domain)" title="CARD (domain)">CARD</a></li> <li><a href="/wiki/Pyrin_domain" title="Pyrin domain">Pyrin</a></li></ul></li> <li><a href="/wiki/DEP_domain" title="DEP domain">DEP</a></li> <li><a href="/wiki/DHHC_domain" title="DHHC domain">DHHC</a></li> <li><a href="/wiki/DHR1_domain" title="DHR1 domain">DHR1</a></li> <li><a href="/wiki/DHR2_domain" title="DHR2 domain">DHR2</a></li> <li><a href="/wiki/DM_domain" title="DM domain">DM</a></li> <li><a href="/wiki/EcoEI_R_protein_C-terminal_domain" title="EcoEI R protein C-terminal domain">EcoEI_R_C</a></li> <li><a href="/wiki/EF1_guanine_nucleotide_exchange_domain" title="EF1 guanine nucleotide exchange domain">EF1</a></li> <li><a href="/wiki/ENTH_domain" title="ENTH domain">ENTH</a></li> <li><a href="/wiki/FGGY_carbohydrate_kinase_family" title="FGGY carbohydrate kinase family">FGGY</a></li> <li><a href="/wiki/FYVE_domain" title="FYVE domain">FYVE</a></li> <li><a href="/wiki/HEAT_repeat_domain" class="mw-redirect" title="HEAT repeat domain">HEAT</a></li> <li><a href="/wiki/Kringle_domain" title="Kringle domain">Kringle</a></li> <li><a href="/wiki/LIM_domain" title="LIM domain">LIM</a></li> <li><a href="/wiki/Leucine-rich_repeat" title="Leucine-rich repeat">LRR</a></li> <li><a href="/wiki/NACHT_domain" title="NACHT domain">NACHT</a></li> <li><a href="/wiki/PAS_domain" title="PAS domain">PAS</a> <ul><li><a href="/wiki/Light-oxygen-voltage-sensing_domain" title="Light-oxygen-voltage-sensing domain">LOV</a></li></ul></li> <li><a href="/wiki/PDZ_domain" title="PDZ domain">PDZ</a></li> <li><a href="/wiki/Pleckstrin_homology_domain" title="Pleckstrin homology domain">PH</a></li> <li><a href="/wiki/PX_domain" title="PX domain">PX</a></li> <li><a href="/wiki/SH2_domain" title="SH2 domain">SH2</a></li> <li><a href="/wiki/SH3_domain" title="SH3 domain">SH3</a></li> <li><a href="/wiki/SUN_domain" title="SUN domain">SUN</a></li> <li><a href="/wiki/CRAL-TRIO_domain" title="CRAL-TRIO domain">TRIO</a></li> <li><a href="/wiki/WD40_repeat" title="WD40 repeat">WD40</a></li> <li><a href="/wiki/X8_protein_domain" title="X8 protein domain">X8</a></li> <li><a href="/wiki/YTH_protein_domain" title="YTH protein domain">YTH</a></li> <li><a href="/wiki/Zinc_finger" title="Zinc finger">zinc finger</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="Protein_structural_analysis" 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" style="background:#eee8aa;"><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:Protein_structural_analysis" title="Template:Protein structural analysis"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Protein_structural_analysis" title="Template talk:Protein structural analysis"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Protein_structural_analysis" title="Special:EditPage/Template:Protein structural analysis"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Protein_structural_analysis" style="font-size:114%;margin:0 4em"><a class="mw-selflink selflink">Protein structural analysis</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">High resolution</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/Cryogenic_electron_microscopy" title="Cryogenic electron microscopy">Cryogenic electron microscopy</a></li> <li><a href="/wiki/X-ray_crystallography" title="X-ray crystallography">X-ray crystallography</a></li> <li><a href="/wiki/Nuclear_magnetic_resonance_spectroscopy_of_proteins" title="Nuclear magnetic resonance spectroscopy of proteins">NMR</a></li> <li><a href="/wiki/Electron_crystallography" title="Electron crystallography">Electron crystallography</a></li> <li><a href="/wiki/Electron_paramagnetic_resonance" title="Electron paramagnetic resonance">EPR</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Medium resolution</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/Fiber_diffraction" title="Fiber diffraction">Fiber diffraction</a></li> <li><a href="/wiki/Mass_spectrometry" title="Mass spectrometry">Mass spectrometry</a></li> <li><a href="/wiki/Biological_small-angle_scattering" title="Biological small-angle scattering">SAXS</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Spectroscopic</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nuclear_magnetic_resonance_spectroscopy_of_proteins" title="Nuclear magnetic resonance spectroscopy of proteins">NMR</a></li> <li><a href="/wiki/Circular_dichroism" title="Circular dichroism">Circular dichroism</a></li> <li><a href="/wiki/Dual-polarization_interferometry" title="Dual-polarization interferometry">Dual-polarization interferometry</a></li> <li><a href="/wiki/Absorbance" title="Absorbance">Absorbance</a></li> <li><a href="/wiki/Fluorescence" title="Fluorescence">Fluorescence</a></li> <li><a href="/wiki/Fluorescence_anisotropy" title="Fluorescence anisotropy">Fluorescence anisotropy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Translational diffusion</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/Ultracentrifuge#Analytical_ultracentrifuge" title="Ultracentrifuge">Analytical ultracentrifugation</a></li> <li><a href="/wiki/Size_exclusion_chromatography" class="mw-redirect" title="Size exclusion chromatography">Size exclusion chromatography</a></li> <li><a href="/wiki/Dynamic_light_scattering" title="Dynamic light scattering">Light scattering</a></li> <li><a href="/wiki/Nuclear_magnetic_resonance_spectroscopy_of_proteins" title="Nuclear magnetic resonance spectroscopy of proteins">NMR</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Rotational diffusion</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/Fluorescence_anisotropy" title="Fluorescence anisotropy">Fluorescence anisotropy</a></li> <li><a href="/wiki/Flow_birefringence" title="Flow birefringence">Flow birefringence</a></li> <li><a href="/wiki/Dielectric_relaxation" class="mw-redirect" title="Dielectric relaxation">Dielectric relaxation</a></li> <li><a href="/wiki/Nuclear_magnetic_resonance_spectroscopy_of_proteins" title="Nuclear magnetic resonance spectroscopy of proteins">NMR</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Chemical</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/Hydrogen%E2%80%93deuterium_exchange" title="Hydrogen–deuterium exchange">Hydrogen–deuterium exchange</a></li> <li><a href="/wiki/Site-directed_mutagenesis" title="Site-directed mutagenesis">Site-directed mutagenesis</a></li> <li><a href="/wiki/Chemical_modification" title="Chemical modification">Chemical modification</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Thermodynamic</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/Equilibrium_unfolding" title="Equilibrium unfolding">Equilibrium unfolding</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background:#eee8aa;">Computational</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/Protein_structure_prediction" title="Protein structure prediction">Protein structure prediction</a></li> <li><a href="/wiki/Docking_(molecular)" title="Docking (molecular)">Molecular docking</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background:#eee8aa;"><div><div style="float:left;"><a href="/wiki/Protein_tertiary_structure" title="Protein tertiary structure">←Tertiary structure</a></div><div style="float:right;"><a href="/wiki/Protein_quaternary_structure" title="Protein quaternary structure">Quaternary structure→</a></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="Biomolecular_structure" 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" style="background:lightblue"><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:Biomolecular_structure" title="Template:Biomolecular structure"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Biomolecular_structure" title="Template talk:Biomolecular 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structure">Tertiary</a></li> <li><a href="/wiki/Protein_quaternary_structure" title="Protein quaternary structure">Quaternary</a></li> <li><a class="mw-selflink-fragment" href="#Protein_structure_determination">Determination</a></li> <li><a href="/wiki/Protein_structure_prediction" title="Protein structure prediction">Prediction</a></li> <li><a href="/wiki/Protein_design" title="Protein design">Design</a></li> <li><a href="/wiki/Protein_thermodynamics" class="mw-redirect" title="Protein thermodynamics">Thermodynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:lightblue;width:1%"><a href="/wiki/Nucleic_acid_structure" title="Nucleic acid structure">Nucleic acid</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/Nucleic_acid_sequence" title="Nucleic acid sequence">Primary</a></li> <li><a href="/wiki/Nucleic_acid_secondary_structure" title="Nucleic acid secondary structure">Secondary</a></li> <li><a href="/wiki/Nucleic_acid_tertiary_structure" title="Nucleic acid tertiary structure">Tertiary</a></li> <li><a href="/wiki/Nucleic_acid_quaternary_structure" title="Nucleic acid quaternary structure">Quaternary</a></li> <li><a href="/wiki/Nucleic_acid_structure_determination" title="Nucleic acid structure determination">Determination</a></li> <li><a href="/wiki/Nucleic_acid_structure_prediction" title="Nucleic acid structure prediction">Prediction</a></li> <li><a href="/wiki/Nucleic_acid_design" title="Nucleic acid design">Design</a></li> <li><a href="/wiki/Nucleic_acid_thermodynamics" title="Nucleic acid thermodynamics">Thermodynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:lightblue;width:1%">See also</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/Protein" title="Protein">Protein</a></li> <li><a href="/wiki/Protein_domain" title="Protein domain">Protein domain</a></li> <li><a href="/wiki/Protein_engineering" title="Protein engineering">Protein engineering</a></li> <li><a href="/wiki/Proteasome" title="Proteasome">Proteasome</a></li> <li><a href="/wiki/Nucleic_acid" title="Nucleic acid">Nucleic acid</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Structural_motif" title="Structural motif">Structural motif</a></li> <li><a href="/wiki/Nucleic_acid_double_helix" title="Nucleic acid double helix">Nucleic acid double helix</a></li></ul> </div></td></tr></tbody></table></div> <ul><li><a rel="nofollow" class="external text" href="https://www.drugdesign.org/chapters/protein-structure/">Protein Structure</a> drugdesign.org</li></ul>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" 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