CINXE.COM

<!DOCTYPE html> <html class="client-nojs vector-feature-language-in-header-enabled vector-feature-language-in-main-page-header-disabled vector-feature-sticky-header-disabled vector-feature-page-tools-pinned-disabled vector-feature-toc-pinned-clientpref-1 vector-feature-main-menu-pinned-disabled vector-feature-limited-width-clientpref-1 vector-feature-limited-width-content-enabled vector-feature-custom-font-size-clientpref-1 vector-feature-appearance-pinned-clientpref-1 vector-feature-night-mode-enabled skin-theme-clientpref-day vector-toc-available" lang="en" dir="ltr"> <head> <meta charset="UTF-8"> <title>Visual phototransduction - Wikipedia</title> <script>(function(){var className="client-js vector-feature-language-in-header-enabled vector-feature-language-in-main-page-header-disabled vector-feature-sticky-header-disabled vector-feature-page-tools-pinned-disabled vector-feature-toc-pinned-clientpref-1 vector-feature-main-menu-pinned-disabled vector-feature-limited-width-clientpref-1 vector-feature-limited-width-content-enabled vector-feature-custom-font-size-clientpref-1 vector-feature-appearance-pinned-clientpref-1 vector-feature-night-mode-enabled skin-theme-clientpref-day vector-toc-available";var cookie=document.cookie.match(/(?:^|; )enwikimwclientpreferences=([^;]+)/);if(cookie){cookie[1].split('%2C').forEach(function(pref){className=className.replace(new RegExp('(^| )'+pref.replace(/-clientpref-\w+$|[^\w-]+/g,'')+'-clientpref-\\w+( |$)'),'$1'+pref+'$2');});}document.documentElement.className=className;}());RLCONF={"wgBreakFrames":false,"wgSeparatorTransformTable":["",""],"wgDigitTransformTable":["",""],"wgDefaultDateFormat":"dmy", "wgMonthNames":["","January","February","March","April","May","June","July","August","September","October","November","December"],"wgRequestId":"ef03035f-bd3c-4c98-9a0b-1e69298c4746","wgCanonicalNamespace":"","wgCanonicalSpecialPageName":false,"wgNamespaceNumber":0,"wgPageName":"Visual_phototransduction","wgTitle":"Visual phototransduction","wgCurRevisionId":1244956549,"wgRevisionId":1244956549,"wgArticleId":1887433,"wgIsArticle":true,"wgIsRedirect":false,"wgAction":"view","wgUserName":null,"wgUserGroups":["*"],"wgCategories":["CS1 maint: multiple names: authors list","Articles with short description","Short description is different from Wikidata","Wikipedia articles that are too technical from January 2024","All articles that are too technical","All articles with unsourced statements","Articles with unsourced statements from January 2024","All pages needing factual verification","Wikipedia articles needing factual verification from January 2024","Articles needing attention", "Visual system","Nervous system","Sensory receptors","Metabolism"],"wgPageViewLanguage":"en","wgPageContentLanguage":"en","wgPageContentModel":"wikitext","wgRelevantPageName":"Visual_phototransduction","wgRelevantArticleId":1887433,"wgIsProbablyEditable":true,"wgRelevantPageIsProbablyEditable":true,"wgRestrictionEdit":[],"wgRestrictionMove":[],"wgNoticeProject":"wikipedia","wgCiteReferencePreviewsActive":false,"wgFlaggedRevsParams":{"tags":{"status":{"levels":1}}},"wgMediaViewerOnClick":true,"wgMediaViewerEnabledByDefault":true,"wgPopupsFlags":0,"wgVisualEditor":{"pageLanguageCode":"en","pageLanguageDir":"ltr","pageVariantFallbacks":"en"},"wgMFDisplayWikibaseDescriptions":{"search":true,"watchlist":true,"tagline":false,"nearby":true},"wgWMESchemaEditAttemptStepOversample":false,"wgWMEPageLength":20000,"wgRelatedArticlesCompat":[],"wgCentralAuthMobileDomain":false,"wgEditSubmitButtonLabelPublish":true,"wgULSPosition":"interlanguage","wgULSisCompactLinksEnabled":false, "wgVector2022LanguageInHeader":true,"wgULSisLanguageSelectorEmpty":false,"wgWikibaseItemId":"Q445081","wgCheckUserClientHintsHeadersJsApi":["brands","architecture","bitness","fullVersionList","mobile","model","platform","platformVersion"],"GEHomepageSuggestedEditsEnableTopics":true,"wgGETopicsMatchModeEnabled":false,"wgGEStructuredTaskRejectionReasonTextInputEnabled":false,"wgGELevelingUpEnabledForUser":false};RLSTATE={"ext.globalCssJs.user.styles":"ready","site.styles":"ready","user.styles":"ready","ext.globalCssJs.user":"ready","user":"ready","user.options":"loading","ext.cite.styles":"ready","skins.vector.search.codex.styles":"ready","skins.vector.styles":"ready","skins.vector.icons":"ready","jquery.makeCollapsible.styles":"ready","ext.wikimediamessages.styles":"ready","ext.visualEditor.desktopArticleTarget.noscript":"ready","ext.uls.interlanguage":"ready","wikibase.client.init":"ready","ext.wikimediaBadges":"ready"};RLPAGEMODULES=["ext.cite.ux-enhancements","mediawiki.page.media", "site","mediawiki.page.ready","jquery.makeCollapsible","mediawiki.toc","skins.vector.js","ext.centralNotice.geoIP","ext.centralNotice.startUp","ext.gadget.ReferenceTooltips","ext.gadget.switcher","ext.urlShortener.toolbar","ext.centralauth.centralautologin","mmv.bootstrap","ext.popups","ext.visualEditor.desktopArticleTarget.init","ext.visualEditor.targetLoader","ext.echo.centralauth","ext.eventLogging","ext.wikimediaEvents","ext.navigationTiming","ext.uls.interface","ext.cx.eventlogging.campaigns","ext.cx.uls.quick.actions","wikibase.client.vector-2022","ext.checkUser.clientHints","ext.growthExperiments.SuggestedEditSession","wikibase.sidebar.tracking"];</script> <script>(RLQ=window.RLQ||[]).push(function(){mw.loader.impl(function(){return["user.options@12s5i",function($,jQuery,require,module){mw.user.tokens.set({"patrolToken":"+\\","watchToken":"+\\","csrfToken":"+\\"}); }];});});</script> <link rel="stylesheet" href="/w/load.php?lang=en&modules=ext.cite.styles%7Cext.uls.interlanguage%7Cext.visualEditor.desktopArticleTarget.noscript%7Cext.wikimediaBadges%7Cext.wikimediamessages.styles%7Cjquery.makeCollapsible.styles%7Cskins.vector.icons%2Cstyles%7Cskins.vector.search.codex.styles%7Cwikibase.client.init&only=styles&skin=vector-2022"> <script async="" src="/w/load.php?lang=en&modules=startup&only=scripts&raw=1&skin=vector-2022"></script> <meta name="ResourceLoaderDynamicStyles" content=""> <link rel="stylesheet" href="/w/load.php?lang=en&modules=site.styles&only=styles&skin=vector-2022"> <meta name="generator" content="MediaWiki 1.44.0-wmf.4"> <meta name="referrer" content="origin"> <meta name="referrer" content="origin-when-cross-origin"> <meta name="robots" content="max-image-preview:standard"> <meta name="format-detection" content="telephone=no"> <meta name="viewport" content="width=1120"> <meta property="og:title" content="Visual phototransduction - Wikipedia"> <meta property="og:type" content="website"> <link rel="preconnect" href="//upload.wikimedia.org"> <link rel="alternate" media="only screen and (max-width: 640px)" href="//en.m.wikipedia.org/wiki/Visual_phototransduction"> <link rel="alternate" type="application/x-wiki" title="Edit this page" href="/w/index.php?title=Visual_phototransduction&action=edit"> <link rel="apple-touch-icon" href="/static/apple-touch/wikipedia.png"> <link rel="icon" href="/static/favicon/wikipedia.ico"> <link rel="search" type="application/opensearchdescription+xml" href="/w/rest.php/v1/search" title="Wikipedia (en)"> <link rel="EditURI" type="application/rsd+xml" href="//en.wikipedia.org/w/api.php?action=rsd"> <link rel="canonical" href="https://en.wikipedia.org/wiki/Visual_phototransduction"> <link rel="license" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en"> <link rel="alternate" type="application/atom+xml" title="Wikipedia Atom feed" href="/w/index.php?title=Special:RecentChanges&feed=atom"> <link rel="dns-prefetch" href="//meta.wikimedia.org" /> <link rel="dns-prefetch" href="//login.wikimedia.org"> </head> <body class="skin--responsive skin-vector skin-vector-search-vue mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject mw-editable page-Visual_phototransduction rootpage-Visual_phototransduction skin-vector-2022 action-view"><a class="mw-jump-link" href="#bodyContent">Jump to content</a> <div class="vector-header-container"> <header class="vector-header mw-header"> <div class="vector-header-start"> <nav class="vector-main-menu-landmark" aria-label="Site"> <div id="vector-main-menu-dropdown" class="vector-dropdown vector-main-menu-dropdown vector-button-flush-left vector-button-flush-right" > <input type="checkbox" id="vector-main-menu-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-main-menu-dropdown" class="vector-dropdown-checkbox " aria-label="Main menu" > <label id="vector-main-menu-dropdown-label" for="vector-main-menu-dropdown-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-menu mw-ui-icon-wikimedia-menu"></span> <span class="vector-dropdown-label-text">Main menu</span> </label> <div class="vector-dropdown-content"> <div id="vector-main-menu-unpinned-container" class="vector-unpinned-container"> <div id="vector-main-menu" class="vector-main-menu vector-pinnable-element"> <div class="vector-pinnable-header vector-main-menu-pinnable-header vector-pinnable-header-unpinned" data-feature-name="main-menu-pinned" data-pinnable-element-id="vector-main-menu" data-pinned-container-id="vector-main-menu-pinned-container" data-unpinned-container-id="vector-main-menu-unpinned-container" > <div class="vector-pinnable-header-label">Main menu</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-main-menu.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-main-menu.unpin">hide</button> </div> <div id="p-navigation" class="vector-menu mw-portlet mw-portlet-navigation" > <div class="vector-menu-heading"> Navigation </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="n-mainpage-description" class="mw-list-item"><a href="/wiki/Main_Page" title="Visit the main page [z]" accesskey="z"><span>Main page</span></a></li><li id="n-contents" class="mw-list-item"><a href="/wiki/Wikipedia:Contents" title="Guides to browsing Wikipedia"><span>Contents</span></a></li><li id="n-currentevents" class="mw-list-item"><a href="/wiki/Portal:Current_events" title="Articles related to current events"><span>Current events</span></a></li><li id="n-randompage" class="mw-list-item"><a href="/wiki/Special:Random" title="Visit a randomly selected article [x]" accesskey="x"><span>Random article</span></a></li><li id="n-aboutsite" class="mw-list-item"><a href="/wiki/Wikipedia:About" title="Learn about Wikipedia and how it works"><span>About Wikipedia</span></a></li><li id="n-contactpage" class="mw-list-item"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us" title="How to contact Wikipedia"><span>Contact us</span></a></li> </ul> </div> </div> <div id="p-interaction" class="vector-menu mw-portlet mw-portlet-interaction" > <div class="vector-menu-heading"> Contribute </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="n-help" class="mw-list-item"><a href="/wiki/Help:Contents" title="Guidance on how to use and edit Wikipedia"><span>Help</span></a></li><li id="n-introduction" class="mw-list-item"><a href="/wiki/Help:Introduction" title="Learn how to edit Wikipedia"><span>Learn to edit</span></a></li><li id="n-portal" class="mw-list-item"><a href="/wiki/Wikipedia:Community_portal" title="The hub for editors"><span>Community portal</span></a></li><li id="n-recentchanges" class="mw-list-item"><a href="/wiki/Special:RecentChanges" title="A list of recent changes to Wikipedia [r]" accesskey="r"><span>Recent changes</span></a></li><li id="n-upload" class="mw-list-item"><a href="/wiki/Wikipedia:File_upload_wizard" title="Add images or other media for use on Wikipedia"><span>Upload file</span></a></li> </ul> </div> </div> </div> </div> </div> </div> </nav> <a href="/wiki/Main_Page" class="mw-logo"> <img class="mw-logo-icon" src="/static/images/icons/wikipedia.png" alt="" aria-hidden="true" height="50" width="50"> <span class="mw-logo-container skin-invert"> <img class="mw-logo-wordmark" alt="Wikipedia" src="/static/images/mobile/copyright/wikipedia-wordmark-en.svg" style="width: 7.5em; height: 1.125em;"> <img class="mw-logo-tagline" alt="The Free Encyclopedia" src="/static/images/mobile/copyright/wikipedia-tagline-en.svg" width="117" height="13" style="width: 7.3125em; height: 0.8125em;"> </span> </a> </div> <div class="vector-header-end"> <div id="p-search" role="search" class="vector-search-box-vue vector-search-box-collapses vector-search-box-show-thumbnail vector-search-box-auto-expand-width vector-search-box"> <a href="/wiki/Special:Search" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only search-toggle" title="Search Wikipedia [f]" accesskey="f"><span class="vector-icon mw-ui-icon-search mw-ui-icon-wikimedia-search"></span> <span>Search</span> </a> <div class="vector-typeahead-search-container"> <div class="cdx-typeahead-search cdx-typeahead-search--show-thumbnail cdx-typeahead-search--auto-expand-width"> <form action="/w/index.php" id="searchform" class="cdx-search-input cdx-search-input--has-end-button"> <div id="simpleSearch" class="cdx-search-input__input-wrapper" data-search-loc="header-moved"> <div class="cdx-text-input cdx-text-input--has-start-icon"> <input class="cdx-text-input__input" type="search" name="search" placeholder="Search Wikipedia" aria-label="Search Wikipedia" autocapitalize="sentences" title="Search Wikipedia [f]" accesskey="f" id="searchInput" > <span class="cdx-text-input__icon cdx-text-input__start-icon"></span> </div> <input type="hidden" name="title" value="Special:Search"> </div> <button class="cdx-button cdx-search-input__end-button">Search</button> </form> </div> </div> </div> <nav class="vector-user-links vector-user-links-wide" aria-label="Personal tools"> <div class="vector-user-links-main"> <div id="p-vector-user-menu-preferences" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <div id="p-vector-user-menu-userpage" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <nav class="vector-appearance-landmark" aria-label="Appearance"> <div id="vector-appearance-dropdown" class="vector-dropdown " title="Change the appearance of the page's font size, width, and color" > <input type="checkbox" id="vector-appearance-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-appearance-dropdown" class="vector-dropdown-checkbox " aria-label="Appearance" > <label id="vector-appearance-dropdown-label" for="vector-appearance-dropdown-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-appearance mw-ui-icon-wikimedia-appearance"></span> <span class="vector-dropdown-label-text">Appearance</span> </label> <div class="vector-dropdown-content"> <div id="vector-appearance-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <div id="p-vector-user-menu-notifications" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <div id="p-vector-user-menu-overflow" class="vector-menu mw-portlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-sitesupport-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="https://donate.wikimedia.org/wiki/Special:FundraiserRedirector?utm_source=donate&utm_medium=sidebar&utm_campaign=C13_en.wikipedia.org&uselang=en" class=""><span>Donate</span></a> </li> <li id="pt-createaccount-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="/w/index.php?title=Special:CreateAccount&returnto=Visual+phototransduction" title="You are encouraged to create an account and log in; however, it is not mandatory" class=""><span>Create account</span></a> </li> <li id="pt-login-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="/w/index.php?title=Special:UserLogin&returnto=Visual+phototransduction" title="You're encouraged to log in; however, it's not mandatory. [o]" accesskey="o" class=""><span>Log in</span></a> </li> </ul> </div> </div> </div> <div id="vector-user-links-dropdown" class="vector-dropdown vector-user-menu vector-button-flush-right vector-user-menu-logged-out" title="Log in and more options" > <input type="checkbox" id="vector-user-links-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-user-links-dropdown" class="vector-dropdown-checkbox " aria-label="Personal tools" > <label id="vector-user-links-dropdown-label" for="vector-user-links-dropdown-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-ellipsis mw-ui-icon-wikimedia-ellipsis"></span> <span class="vector-dropdown-label-text">Personal tools</span> </label> <div class="vector-dropdown-content"> <div id="p-personal" class="vector-menu mw-portlet mw-portlet-personal user-links-collapsible-item" title="User menu" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-sitesupport" class="user-links-collapsible-item mw-list-item"><a href="https://donate.wikimedia.org/wiki/Special:FundraiserRedirector?utm_source=donate&utm_medium=sidebar&utm_campaign=C13_en.wikipedia.org&uselang=en"><span>Donate</span></a></li><li id="pt-createaccount" class="user-links-collapsible-item mw-list-item"><a href="/w/index.php?title=Special:CreateAccount&returnto=Visual+phototransduction" title="You are encouraged to create an account and log in; however, it is not mandatory"><span class="vector-icon mw-ui-icon-userAdd mw-ui-icon-wikimedia-userAdd"></span> <span>Create account</span></a></li><li id="pt-login" class="user-links-collapsible-item mw-list-item"><a href="/w/index.php?title=Special:UserLogin&returnto=Visual+phototransduction" title="You're encouraged to log in; however, it's not mandatory. [o]" accesskey="o"><span class="vector-icon mw-ui-icon-logIn mw-ui-icon-wikimedia-logIn"></span> <span>Log in</span></a></li> </ul> </div> </div> <div id="p-user-menu-anon-editor" class="vector-menu mw-portlet mw-portlet-user-menu-anon-editor" > <div class="vector-menu-heading"> Pages for logged out editors <a href="/wiki/Help:Introduction" aria-label="Learn more about editing"><span>learn more</span></a> </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-anoncontribs" class="mw-list-item"><a href="/wiki/Special:MyContributions" title="A list of edits made from this IP address [y]" accesskey="y"><span>Contributions</span></a></li><li id="pt-anontalk" class="mw-list-item"><a href="/wiki/Special:MyTalk" title="Discussion about edits from this IP address [n]" accesskey="n"><span>Talk</span></a></li> </ul> </div> </div> </div> </div> </nav> </div> </header> </div> <div class="mw-page-container"> <div class="mw-page-container-inner"> <div class="vector-sitenotice-container"> <div id="siteNotice"></div> </div> <div class="vector-column-start"> <div class="vector-main-menu-container"> <div id="mw-navigation"> <nav id="mw-panel" class="vector-main-menu-landmark" aria-label="Site"> <div id="vector-main-menu-pinned-container" class="vector-pinned-container"> </div> </nav> </div> </div> <div class="vector-sticky-pinned-container"> <nav id="mw-panel-toc" aria-label="Contents" data-event-name="ui.sidebar-toc" class="mw-table-of-contents-container vector-toc-landmark"> <div id="vector-toc-pinned-container" class="vector-pinned-container"> <div id="vector-toc" class="vector-toc vector-pinnable-element"> <div class="vector-pinnable-header vector-toc-pinnable-header vector-pinnable-header-pinned" data-feature-name="toc-pinned" data-pinnable-element-id="vector-toc" > <h2 class="vector-pinnable-header-label">Contents</h2> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" 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-Overview" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Overview"> <div class="vector-toc-text"> <span class="vector-toc-numb">1</span> <span>Overview</span> </div> </a> <ul id="toc-Overview-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Photoreceptors" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Photoreceptors"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Photoreceptors</span> </div> </a> <ul id="toc-Photoreceptors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Transduction_process" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Transduction_process"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Transduction process</span> </div> </a> <button aria-controls="toc-Transduction_process-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 Transduction process subsection</span> </button> <ul id="toc-Transduction_process-sublist" class="vector-toc-list"> <li id="toc-In_the_dark" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_the_dark"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>In the dark</span> </div> </a> <ul id="toc-In_the_dark-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_the_light" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_the_light"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>In the light</span> </div> </a> <ul id="toc-In_the_light-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Deactivation_of_the_phototransduction_cascade" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Deactivation_of_the_phototransduction_cascade"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Deactivation of the phototransduction cascade</span> </div> </a> <ul id="toc-Deactivation_of_the_phototransduction_cascade-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Visual_cycle" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Visual_cycle"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Visual cycle</span> </div> </a> <ul id="toc-Visual_cycle-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_invertebrates" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#In_invertebrates"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>In invertebrates</span> </div> </a> <ul id="toc-In_invertebrates-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">6</span> <span>References</span> </div> </a> <ul id="toc-References-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">7</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">Visual phototransduction</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 12 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-12" 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">12 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%AA%D8%AD%D9%88%D9%84_%D8%B6%D9%88%D8%A6%D9%8A_%D8%A8%D8%B5%D8%B1%D9%8A" title="تحول ضوئي بصري – Arabic" lang="ar" hreflang="ar" data-title="تحول ضوئي بصري" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Cicle_visual" title="Cicle visual – Catalan" lang="ca" hreflang="ca" data-title="Cicle visual" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Phototransduktion" title="Phototransduktion – German" lang="de" hreflang="de" data-title="Phototransduktion" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Fototransduktsioon" title="Fototransduktsioon – Estonian" lang="et" hreflang="et" data-title="Fototransduktsioon" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%D8%B1%D8%A7%D8%B1%D8%B3%D8%A7%D9%86%DB%8C_%D9%86%D9%88%D8%B1%DB%8C_%D8%A8%DB%8C%D9%86%D8%A7%DB%8C%DB%8C" title="ترارسانی نوری بینایی – Persian" lang="fa" hreflang="fa" data-title="ترارسانی نوری بینایی" 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/Cycle_visuel" title="Cycle visuel – French" lang="fr" hreflang="fr" data-title="Cycle visuel" 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/Fototransduci%C3%B3n_visual" title="Fototransdución visual – Galician" lang="gl" hreflang="gl" data-title="Fototransdución visual" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%8B%9C%EA%B0%81_%EA%B4%91%EB%B3%80%ED%99%98" title="시각 광변환 – Korean" lang="ko" hreflang="ko" data-title="시각 광변환" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B5%E0%B4%BF%E0%B4%B7%E0%B5%8D%E0%B4%B5%E0%B5%BD_%E0%B4%AB%E0%B5%8B%E0%B4%9F%E0%B5%8D%E0%B4%9F%E0%B5%8B%E0%B4%9F%E0%B5%8D%E0%B4%B0%E0%B4%BE%E0%B5%BB%E0%B4%B8%E0%B5%8D%E0%B4%A1%E0%B4%95%E0%B5%8D%E0%B4%B7%E0%B5%BB" title="വിഷ്വൽ ഫോട്ടോട്രാൻസ്ഡക്ഷൻ – Malayalam" lang="ml" hreflang="ml" data-title="വിഷ്വൽ ഫോട്ടോട്രാൻസ്ഡക്ഷൻ" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%93%D1%8D%D1%80%D1%8D%D0%BB_%D0%B4%D0%B0%D0%BC%D0%B6%D1%83%D1%83%D0%BB%D0%B0%D0%BB%D1%82" title="Гэрэл дамжуулалт – Mongolian" lang="mn" hreflang="mn" data-title="Гэрэл дамжуулалт" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%A4%D0%BE%D1%82%D0%BE%D1%80%D0%B5%D1%86%D0%B5%D0%BF%D1%86%D0%B8%D1%98%D0%B0" title="Фоторецепција – Serbian" lang="sr" hreflang="sr" data-title="Фоторецепција" 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-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%A6%96%E8%A6%BA%E5%85%89%E8%BD%89%E5%B0%8E" title="視覺光轉導 – Chinese" lang="zh" hreflang="zh" data-title="視覺光轉導" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q445081#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-associated-pages" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-nstab-main" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Visual_phototransduction" title="View the content page [c]" accesskey="c"><span>Article</span></a></li><li id="ca-talk" class="vector-tab-noicon mw-list-item"><a href="/wiki/Talk:Visual_phototransduction" rel="discussion" title="Discuss improvements to the content page [t]" accesskey="t"><span>Talk</span></a></li> </ul> </div> </div> <div id="vector-variants-dropdown" class="vector-dropdown emptyPortlet" > <input type="checkbox" id="vector-variants-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-variants-dropdown" class="vector-dropdown-checkbox " aria-label="Change language variant" > <label id="vector-variants-dropdown-label" for="vector-variants-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet" aria-hidden="true" ><span class="vector-dropdown-label-text">English</span> </label> <div class="vector-dropdown-content"> <div id="p-variants" class="vector-menu mw-portlet mw-portlet-variants emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> </div> </div> </nav> </div> <div id="right-navigation" class="vector-collapsible"> <nav aria-label="Views"> <div id="p-views" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-views" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-view" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Visual_phototransduction"><span>Read</span></a></li><li id="ca-edit" class="vector-tab-noicon mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&action=edit" title="Edit this page [e]" accesskey="e"><span>Edit</span></a></li><li id="ca-history" class="vector-tab-noicon mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&action=history" title="Past revisions of this page [h]" accesskey="h"><span>View history</span></a></li> </ul> </div> </div> </nav> <nav class="vector-page-tools-landmark" aria-label="Page tools"> <div id="vector-page-tools-dropdown" class="vector-dropdown vector-page-tools-dropdown" > <input type="checkbox" id="vector-page-tools-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-tools-dropdown" class="vector-dropdown-checkbox " aria-label="Tools" > <label id="vector-page-tools-dropdown-label" for="vector-page-tools-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet" aria-hidden="true" ><span class="vector-dropdown-label-text">Tools</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-tools-unpinned-container" class="vector-unpinned-container"> <div id="vector-page-tools" class="vector-page-tools vector-pinnable-element"> <div class="vector-pinnable-header vector-page-tools-pinnable-header vector-pinnable-header-unpinned" data-feature-name="page-tools-pinned" data-pinnable-element-id="vector-page-tools" data-pinned-container-id="vector-page-tools-pinned-container" data-unpinned-container-id="vector-page-tools-unpinned-container" > <div class="vector-pinnable-header-label">Tools</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-page-tools.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-page-tools.unpin">hide</button> </div> <div id="p-cactions" class="vector-menu mw-portlet mw-portlet-cactions emptyPortlet vector-has-collapsible-items" title="More options" > <div class="vector-menu-heading"> Actions </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-more-view" class="selected vector-more-collapsible-item mw-list-item"><a href="/wiki/Visual_phototransduction"><span>Read</span></a></li><li id="ca-more-edit" class="vector-more-collapsible-item mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&action=edit" title="Edit this page [e]" accesskey="e"><span>Edit</span></a></li><li id="ca-more-history" class="vector-more-collapsible-item mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&action=history"><span>View history</span></a></li> </ul> </div> </div> <div id="p-tb" class="vector-menu mw-portlet mw-portlet-tb" > <div class="vector-menu-heading"> General </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="t-whatlinkshere" class="mw-list-item"><a href="/wiki/Special:WhatLinksHere/Visual_phototransduction" title="List of all English Wikipedia pages containing links to this page [j]" accesskey="j"><span>What links here</span></a></li><li id="t-recentchangeslinked" class="mw-list-item"><a href="/wiki/Special:RecentChangesLinked/Visual_phototransduction" rel="nofollow" title="Recent changes in pages linked from this page [k]" accesskey="k"><span>Related changes</span></a></li><li id="t-upload" class="mw-list-item"><a href="/wiki/Wikipedia:File_Upload_Wizard" title="Upload files [u]" accesskey="u"><span>Upload file</span></a></li><li id="t-specialpages" class="mw-list-item"><a href="/wiki/Special:SpecialPages" title="A list of all special pages [q]" accesskey="q"><span>Special pages</span></a></li><li id="t-permalink" class="mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&oldid=1244956549" title="Permanent link to this revision of this page"><span>Permanent link</span></a></li><li id="t-info" class="mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&action=info" title="More information about this page"><span>Page information</span></a></li><li id="t-cite" class="mw-list-item"><a href="/w/index.php?title=Special:CiteThisPage&page=Visual_phototransduction&id=1244956549&wpFormIdentifier=titleform" title="Information on how to cite this page"><span>Cite this page</span></a></li><li id="t-urlshortener" class="mw-list-item"><a href="/w/index.php?title=Special:UrlShortener&url=https%3A%2F%2Fen.wikipedia.org%2Fwiki%2FVisual_phototransduction"><span>Get shortened URL</span></a></li><li id="t-urlshortener-qrcode" class="mw-list-item"><a href="/w/index.php?title=Special:QrCode&url=https%3A%2F%2Fen.wikipedia.org%2Fwiki%2FVisual_phototransduction"><span>Download QR code</span></a></li> </ul> </div> </div> <div id="p-coll-print_export" class="vector-menu mw-portlet mw-portlet-coll-print_export" > <div class="vector-menu-heading"> Print/export </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="coll-download-as-rl" class="mw-list-item"><a href="/w/index.php?title=Special:DownloadAsPdf&page=Visual_phototransduction&action=show-download-screen" title="Download this page as a PDF file"><span>Download as PDF</span></a></li><li id="t-print" class="mw-list-item"><a href="/w/index.php?title=Visual_phototransduction&printable=yes" title="Printable version of this page [p]" accesskey="p"><span>Printable version</span></a></li> </ul> </div> </div> <div id="p-wikibase-otherprojects" class="vector-menu mw-portlet mw-portlet-wikibase-otherprojects" > <div class="vector-menu-heading"> In other projects </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="t-wikibase" class="wb-otherproject-link wb-otherproject-wikibase-dataitem mw-list-item"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q445081" title="Structured data on this page hosted by Wikidata [g]" accesskey="g"><span>Wikidata item</span></a></li> </ul> </div> </div> </div> </div> </div> </div> </nav> </div> </div> </div> <div class="vector-column-end"> <div class="vector-sticky-pinned-container"> <nav class="vector-page-tools-landmark" aria-label="Page tools"> <div id="vector-page-tools-pinned-container" class="vector-pinned-container"> </div> </nav> <nav class="vector-appearance-landmark" aria-label="Appearance"> <div id="vector-appearance-pinned-container" class="vector-pinned-container"> <div id="vector-appearance" class="vector-appearance vector-pinnable-element"> <div class="vector-pinnable-header vector-appearance-pinnable-header vector-pinnable-header-pinned" data-feature-name="appearance-pinned" data-pinnable-element-id="vector-appearance" data-pinned-container-id="vector-appearance-pinned-container" data-unpinned-container-id="vector-appearance-unpinned-container" > <div class="vector-pinnable-header-label">Appearance</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Process by which light activates retinal cells</div> <p><b>Visual phototransduction</b> is the <a href="/wiki/Transduction_(physiology)" title="Transduction (physiology)">sensory transduction</a> process of the <a href="/wiki/Visual_system" title="Visual system">visual system</a> by which <a href="/wiki/Light" title="Light">light</a> is detected by <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a> (<a href="/wiki/Rod_cell" title="Rod cell">rods</a> and <a href="/wiki/Cone_cell" title="Cone cell">cones</a>) in the vertebrate <a href="/wiki/Retina" title="Retina">retina</a>. A <a href="/wiki/Photon" title="Photon">photon</a> is absorbed by a <a href="/wiki/Retinal" title="Retinal">retinal</a> <a href="/wiki/Chromophore" title="Chromophore">chromophore</a> (each bound to an <a href="/wiki/Opsin" title="Opsin">opsin</a>), which initiates a signal cascade through several intermediate cells, then through the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">retinal ganglion cells</a> (RGCs) comprising the optic nerve. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=1" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Light enters the eye, passes through the optical media, then the inner neural layers of the retina before finally reaching the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a> in the outer layer of the retina. The light may be absorbed by a <a href="/wiki/Chromophore" title="Chromophore">chromophore</a> bound to an <a href="/wiki/Opsin" title="Opsin">opsin</a>, which <a href="/wiki/Photoisomerization" title="Photoisomerization">photoisomerizes</a> the chromophore, initiating both the <a href="/wiki/Visual_cycle" title="Visual cycle">visual cycle</a>, which "resets" the chromophore, and the phototransduction cascade, which transmits the visual signal to the brain. The cascade begins with graded polarisation (an <a href="/wiki/Analog_signal" title="Analog signal">analog signal</a>) of the excited photoreceptor cell, as its <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a> increases from a <a href="/wiki/Resting_potential" title="Resting potential">resting potential</a> of -70 mV, proportional to the light intensity. At rest, the photoreceptor cells are continually releasing glutamate at the <a href="/wiki/Synaptic_terminal" class="mw-redirect" title="Synaptic terminal">synaptic terminal</a> to maintain the potential.<sup id="cite_ref-Bert20_1-0" class="reference"><a href="#cite_note-Bert20-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The transmitter release rate is lowered (<a href="/wiki/Hyperpolarization_(biology)" title="Hyperpolarization (biology)">hyperpolarization</a>) as light intensity increases. Each synaptic terminal makes up to 500 contacts with <a href="/wiki/Horizontal_cell" class="mw-redirect" title="Horizontal cell">horizontal cells</a> and <a href="/wiki/Bipolar_cell" class="mw-redirect" title="Bipolar cell">bipolar cells</a>.<sup id="cite_ref-Bert20_1-1" class="reference"><a href="#cite_note-Bert20-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> These intermediate cells (along with <a href="/wiki/Amacrine_cell" title="Amacrine cell">amacrine cells</a>) perform comparisons of photoreceptor signals within a <a href="/wiki/Receptive_field#Visual_system" title="Receptive field">receptive field</a>, but their precise functionalities are not well understood. The signal remains as a graded polarization in all cells until it reaches the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">RGCs</a>, where it is converted to an <a href="/wiki/Action_potential" title="Action potential">action potential</a> and transmitted to the brain.<sup id="cite_ref-Bert20_1-2" class="reference"><a href="#cite_note-Bert20-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Photoreceptors">Photoreceptors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=2" title="Edit section: Photoreceptors"><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/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cell</a></div> <p>The <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a> involved in vertebrate vision are the <a href="/wiki/Rod_cell" title="Rod cell">rods</a>, the <a href="/wiki/Cone_cell" title="Cone cell">cones</a>, and the <a href="/wiki/Photosensitive_ganglion_cell" class="mw-redirect" title="Photosensitive ganglion cell">photosensitive ganglion cells</a> (ipRGCs). These cells contain a <a href="/wiki/Chromophore" title="Chromophore">chromophore</a> (<a href="/wiki/11-cis_retinal" class="mw-redirect" title="11-cis retinal">11-<i>cis</i>-retinal</a>, the <a href="/wiki/Aldehyde" title="Aldehyde">aldehyde</a> of <a href="/wiki/Vitamin_a" class="mw-redirect" title="Vitamin a">vitamin A1</a> and light-absorbing portion) that is bound to a cell membrane protein, <a href="/wiki/Opsin" title="Opsin">opsin</a>. Rods are responsible for vision under low light intensity and contrast detections. Because they all have the same response across frequencies, no color information can be deduced from the rods only, as in low light conditions for example. Cones, on the other hand, are of different kinds with different frequency response, such that color can be perceived through comparison of the outputs of different kinds of cones. Each cone type responds best to certain <a href="/wiki/Wavelength" title="Wavelength">wavelengths</a>, or colors, of light because each type has a slightly different opsin. The three types of cones are L-cones, M-cones and S-cones that respond optimally to long wavelengths (reddish color), medium wavelengths (greenish color), and short wavelengths (bluish color) respectively. Humans have <a href="/wiki/Trichromatic" class="mw-redirect" title="Trichromatic">trichromatic</a> <a href="/wiki/Photopic_vision" title="Photopic vision">photopic vision</a> consisting of three <a href="/wiki/Opponent_process" title="Opponent process">opponent process</a> channels that enable <a href="/wiki/Color_vision" title="Color vision">color vision</a>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Rod photoreceptors are the most common cell type in the retina and develop quite late. Most cells become postmitotic before birth, but differentiation occurs after birth. In the first week after birth, cells mature and the eye becomes fully functional at the time of opening. The visual pigment rhodopsin (rho) is the first known sign of differentiation in rods. <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> </p> <div class="mw-heading mw-heading2"><h2 id="Transduction_process">Transduction process</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=3" title="Edit section: Transduction process"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>To understand the photoreceptor's behavior to light intensities, it is necessary to understand the roles of different currents. </p><p>There is an ongoing outward <a href="/wiki/Potassium" title="Potassium">potassium</a> current through nongated K<sup>+</sup>-selective channels. This outward current tends to hyperpolarize the photoreceptor at around −70 mV (the equilibrium potential for K<sup>+</sup>). </p><p>There is also an inward sodium current carried by <a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cGMP</a>-gated <a href="/wiki/Sodium_channel" title="Sodium channel">sodium channels</a>. This "<a href="/wiki/Dark_current_(biochemistry)" class="mw-redirect" title="Dark current (biochemistry)">dark current</a>" depolarizes the cell to around −40 mV. This is significantly more depolarized than most other neurons. </p><p>A high density of <a href="/wiki/Sodium%E2%80%93potassium_pump" title="Sodium–potassium pump">Na<sup>+</sup>-K<sup>+</sup> pumps</a> enables the photoreceptor to maintain a steady intracellular concentration of Na<sup>+</sup> and K<sup>+</sup>. </p><p>When light intensity increases, the potential of the membrane decreases (hyperpolarization). Because as the intensity increases, the release of the stimulating neurotransmitter glutamate of the photoreceptors is reduced. When light intensity decreases, that is, in the dark environment, glutamate release by photoreceptors increases. This increases the membrane potential and produces membrane depolarization.<sup id="cite_ref-Bert20_1-3" class="reference"><a href="#cite_note-Bert20-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_the_dark">In the dark</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=4" title="Edit section: In the dark"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Photoreceptor_cells" class="mw-redirect" title="Photoreceptor cells">Photoreceptor cells</a> are unusual cells in that they depolarize in response to absence of stimuli or scotopic conditions (darkness). In photopic conditions (light), photoreceptors hyperpolarize to a potential of −60 mV. </p><p>In the dark, cGMP levels are high and keep cGMP-gated sodium channels open allowing a steady inward current, called the dark current. This dark current keeps the cell depolarized at about −40 mV, leading to <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> release which inhibits excitation of neurons. </p><p>The depolarization of the cell membrane in <a href="/wiki/Scotopic" class="mw-redirect" title="Scotopic">scotopic</a> conditions opens voltage-gated calcium channels. An increased intracellular concentration of <a href="/wiki/Ca2%2B" class="mw-redirect" title="Ca2+">Ca<sup>2+</sup></a> causes <a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">vesicles</a> containing glutamate, a <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a>, to merge with the cell membrane, therefore releasing glutamate into the <a href="/wiki/Synaptic_cleft" class="mw-redirect" title="Synaptic cleft">synaptic cleft</a>, an area between the end of one cell and the beginning of another <a href="/wiki/Neuron" title="Neuron">neuron</a>. Glutamate, though usually excitatory, functions here as an inhibitory neurotransmitter. </p><p>In the cone pathway, glutamate: </p> <ul><li>Hyperpolarizes on-center <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar cells</a>. Glutamate that is released from the photoreceptors in the dark binds to metabotropic glutamate receptors (<a href="/wiki/Metabotropic_glutamate_receptor_6" title="Metabotropic glutamate receptor 6">mGluR6</a>), which, through a G-protein coupling mechanism, causes non-specific cation channels in the cells to close, thus hyperpolarizing the bipolar cell.</li> <li>Depolarizes off-center bipolar cells. Binding of glutamate to ionotropic glutamate receptors results in an inward cation current that depolarizes the bipolar cell.</li></ul> <div class="mw-heading mw-heading3"><h3 id="In_the_light">In the light</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=5" title="Edit section: In the light"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In summary: <i>Light</i> closes cGMP-gated sodium channels, reducing the influx of both Na<sup>+</sup> and Ca<sup>2+</sup> ions. Stopping the influx of Na<sup>+</sup> ions effectively switches <i>off</i> the dark current. Reducing this dark current causes the photoreceptor to <i>hyperpolarise</i>, which reduces glutamate release which thus reduces the <i>inhibition</i> of retinal nerves, leading to <i>excitation</i> of these nerves. This reduced Ca<sup>2+</sup> influx during phototransduction enables deactivation and recovery from phototransduction, as discussed below in <a href="#Deactivation_of_the_phototransduction_cascade">§ Deactivation of the phototransduction cascade</a>. </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Phototransduction.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Phototransduction.png/500px-Phototransduction.png" decoding="async" width="500" height="211" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/de/Phototransduction.png/750px-Phototransduction.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/de/Phototransduction.png/1000px-Phototransduction.png 2x" data-file-width="1327" data-file-height="561" /></a><figcaption>Representation of molecular steps in photoactivation (modified from Leskov et al., 2000<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>). Depicted is an outer membrane disk in a rod. <b>Step 1</b>: Incident photon (hν) is absorbed and activates a <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a> (likewise <a href="/wiki/Photopsin" class="mw-redirect" title="Photopsin">photopsin</a>) by conformational change in the disk membrane to R*. <b>Step 2</b>: Next, R* makes repeated contacts with transducin molecules, catalyzing its activation to G* by the release of bound GDP in exchange for cytoplasmic GTP, which expels its β and γ subunits. <b>Step 3</b>: G* binds inhibitory γ subunits of the phosphodiesterase (PDE) activating its α and β subunits. <b>Step 4</b>: Activated PDE hydrolyzes cGMP. <b>Step 5</b>: Guanylyl cyclase (GC) synthesizes cGMP, the second messenger in the phototransduction cascade. Reduced levels of cytosolic cGMP cause cyclic nucleotide gated channels to close preventing further influx of Na<sup>+</sup> and Ca<sup>2+</sup>.</figcaption></figure> <ol><li>A photon interacts with a <a href="/wiki/Retinal" title="Retinal">retinal</a> molecule in an <a href="/wiki/Opsin" title="Opsin">opsin</a> complex in a <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cell</a>. The retinal undergoes <a href="/wiki/Isomerisation" class="mw-redirect" title="Isomerisation">isomerisation</a>, changing from the 11-<i>cis</i>-retinal to the all-<i>trans</i>-retinal configuration.</li> <li>Opsin therefore undergoes a conformational change to metarhodopsin II.</li> <li>Metarhodopsin II activates a <a href="/wiki/G_protein" title="G protein">G protein</a> known as <a href="/wiki/Transducin" title="Transducin">transducin</a>. This causes transducin to dissociate from its bound <a href="/wiki/Guanosine_diphosphate" title="Guanosine diphosphate">GDP</a>, and bind <a href="/wiki/Guanosine_triphosphate" title="Guanosine triphosphate">GTP</a>; then the alpha subunit of transducin dissociates from the beta and gamma subunits, with the GTP still bound to the alpha subunit.</li> <li>The alpha subunit-GTP complex activates <a href="/wiki/Phosphodiesterase" title="Phosphodiesterase">phosphodiesterase</a>, also known as PDE6. It binds to one of two regulatory subunits of PDE (which itself is a tetramer) and stimulates its activity.</li> <li>PDE hydrolyzes <a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cGMP</a>, forming <a href="/wiki/Guanosine_monophosphate" title="Guanosine monophosphate">GMP</a>. This lowers the intracellular concentration of cGMP and therefore the sodium channels close.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup></li> <li>Closure of the sodium channels causes hyperpolarization of the cell due to the ongoing efflux of potassium ions.</li> <li>Hyperpolarization of the cell causes voltage-gated calcium channels to close.</li> <li>As the calcium level in the photoreceptor cell drops, the amount of the neurotransmitter glutamate that is released by the cell also drops. This is because calcium is required for the glutamate-containing vesicles to fuse with cell membrane and release their contents (see <a href="/wiki/SNARE_(protein)" class="mw-redirect" title="SNARE (protein)">SNARE proteins</a>).</li> <li>A decrease in the amount of glutamate released by the photoreceptors causes depolarization of on-center bipolar cells (rod and cone On bipolar cells) and hyperpolarization of cone off-center bipolar cells.</li></ol> <div class="mw-heading mw-heading3"><h3 id="Deactivation_of_the_phototransduction_cascade">Deactivation of the phototransduction cascade</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=6" title="Edit section: Deactivation of the phototransduction cascade"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In light, low cGMP levels close Na<sup>+</sup> and Ca<sup>2+</sup> channels, reducing intracellular Na<sup>+</sup> and Ca<sup>2+</sup>. During recovery (<a href="/wiki/Dark_adaptation" class="mw-redirect" title="Dark adaptation">dark adaptation</a>), the low Ca<sup>2+</sup> levels induce recovery (termination of the phototransduction cascade), as follows: </p> <ol><li>Low intracellular Ca<sup>2+</sup> causes Ca<sup>2+</sup> to dissociate from <a href="/wiki/Guanylate_cyclase_activator" title="Guanylate cyclase activator">guanylate cyclase activating protein</a> (GCAP). The liberated GCAP ultimately restores depleted cGMP levels, which re-opens the cGMP-gated cation channels (restoring dark current).</li> <li>Low intracellular Ca<sup>2+</sup> causes Ca<sup>2+</sup> to dissociate from <a href="/wiki/GTPase-activating_protein" title="GTPase-activating protein">GTPase-activating protein</a> (GAP), also known as <a href="/wiki/Regulator_of_G_protein_signaling" title="Regulator of G protein signaling">regulator of G protein signaling</a>. The liberated GAP deactivates transducin, terminating the phototransduction cascade (restoring dark current).</li> <li>Low intracellular Ca<sup>2+</sup> makes intracellular Ca-recoverin-RK dissociate into Ca<sup>2+</sup> and <a href="/wiki/Recoverin" title="Recoverin">recoverin</a> and <a href="/wiki/Rhodopsin_kinase" title="Rhodopsin kinase">rhodopsin kinase</a> (RK). The liberated RK then phosphorylates the Metarhodopsin II, reducing its binding affinity for transducin. <a href="/wiki/Arrestin" title="Arrestin">Arrestin</a> then completely deactivates the phosphorylated-metarhodopsin II, terminating the phototransduction cascade (restoring dark current).</li> <li>Low intracellular Ca<sup>2+</sup> make the Ca<sup>2+</sup>/<a href="/wiki/Calmodulin" title="Calmodulin">calmodulin</a> complex within the cGMP-gated cation channels more sensitive to low cGMP levels (thereby, keeping the cGMP-gated cation channel open even at low cGMP levels, restoring dark current)<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li></ol> <p>In more detail: </p><p>GTPase Accelerating Protein (GAP) of RGS (regulators of G protein signaling) interacts with the alpha subunit of transducin, and causes it to hydrolyse its bound GTP to GDP, and thus halts the action of phosphodiesterase, stopping the transformation of cGMP to GMP. This deactivation step of the phototransduction cascade (the deactivation of the G protein transducer) was found to be the rate limiting step in the deactivation of the phototransduction cascade.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><p>In other words: Guanylate Cyclase Activating Protein (GCAP) is a calcium binding protein, and as the calcium levels in the cell have decreased, GCAP dissociates from its bound calcium ions, and interacts with Guanylate Cyclase, activating it. Guanylate Cyclase then proceeds to transform GTP to cGMP, replenishing the cell's cGMP levels and thus reopening the sodium channels that were closed during phototransduction. </p><p>Finally, Metarhodopsin II is deactivated. Recoverin, another calcium binding protein, is normally bound to Rhodopsin Kinase when calcium is present. When the calcium levels fall during phototransduction, the calcium dissociates from recoverin, and rhodopsin kinase is released and phosphorylates <a href="/w/index.php?title=Metarhodopsin_II&action=edit&redlink=1" class="new" title="Metarhodopsin II (page does not exist)">metarhodopsin II</a>, which decreases its affinity for transducin. Finally, arrestin, another protein, binds the phosphorylated metarhodopsin II, completely deactivating it. Thus, finally, phototransduction is deactivated, and the dark current and glutamate release is restored. It is this pathway, where Metarhodopsin II is phosphorylated and bound to arrestin and thus deactivated, which is thought to be responsible for the S2 component of dark adaptation. The S2 component represents a linear section of the dark adaptation function present at the beginning of dark adaptation for all bleaching intensities. </p> <div class="mw-heading mw-heading2"><h2 id="Visual_cycle">Visual cycle</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=7" title="Edit section: Visual cycle"><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:1415_Retinal_Isomers.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/1415_Retinal_Isomers.jpg/220px-1415_Retinal_Isomers.jpg" decoding="async" width="220" height="208" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/1415_Retinal_Isomers.jpg/330px-1415_Retinal_Isomers.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f8/1415_Retinal_Isomers.jpg/440px-1415_Retinal_Isomers.jpg 2x" data-file-width="2237" data-file-height="2112" /></a><figcaption>The absorption of light leads to an isomeric change in the retinal molecule.</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/Visual_cycle" title="Visual cycle">Visual cycle</a></div> <p>The visual cycle occurs via <a href="/wiki/G-protein_coupled_receptor" class="mw-redirect" title="G-protein coupled receptor">G-protein coupled receptors</a> called <a href="/wiki/Retinylidene_protein" title="Retinylidene protein">retinylidene proteins</a> which consists of a visual <a href="/wiki/Opsin" title="Opsin">opsin</a> and a <a href="/wiki/Chromophore" title="Chromophore">chromophore</a> <a href="/wiki/11-cis_retinal" class="mw-redirect" title="11-cis retinal">11-<i>cis</i>-retinal</a>. The 11-<i>cis</i>-retinal is covalently linked to the <a href="/wiki/Opsin" title="Opsin">opsin</a> receptor via <a href="/wiki/Schiff_base" title="Schiff base">Schiff base</a>. When it absorbs a <a href="/wiki/Photon" title="Photon">photon</a>, 11-<i>cis</i>-retinal undergoes <a href="/wiki/Photoisomerization" title="Photoisomerization">photoisomerization</a> to <a href="/wiki/All-trans-retinal" class="mw-redirect" title="All-trans-retinal">all-<i>trans</i>-retinal</a>, which changes the conformation of the opsin <a href="/wiki/GPCR" class="mw-redirect" title="GPCR">GPCR</a> leading to <a href="/wiki/Signal_transduction" title="Signal transduction">signal transduction</a> cascades which causes closure of cyclic GMP-gated cation channel, and hyperpolarization of the photoreceptor cell. Following photoisomerization, all-<i>trans</i>-retinal is released from the opsin protein and reduced to all-<i>trans</i>-<a href="/wiki/Retinol" title="Retinol">retinol</a>, which travels to the <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a> to be "recharged". It is first <a href="/wiki/Ester" title="Ester">esterified</a> by <a href="/wiki/Lecithin_retinol_acyltransferase" title="Lecithin retinol acyltransferase">lecithin retinol acyltransferase</a> (LRAT) and then converted to 11-<i>cis</i>-retinol by the isomerohydrolase <a href="/wiki/RPE65" title="RPE65">RPE65</a>. The isomerase activity of RPE65 has been shown; it is uncertain whether it also acts as the hydrolase.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Finally, it is oxidized to 11-<i>cis</i>-retinal before traveling back to the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cell</a> outer segment where it is again conjugated to an <a href="/wiki/Opsin" title="Opsin">opsin</a> to form new, functional visual pigment (<a href="/wiki/Retinylidene_protein" title="Retinylidene protein">retinylidene protein</a>), namely <a href="/wiki/Photopsin" class="mw-redirect" title="Photopsin">photopsin</a> or <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a>. </p> <div class="mw-heading mw-heading2"><h2 id="In_invertebrates">In invertebrates</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=8" title="Edit section: In invertebrates"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <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-Technical plainlinks metadata ambox ambox-style ambox-technical" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>may be too technical for most readers to understand</b>.<span class="hide-when-compact"> Please <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Visual_phototransduction&action=edit">help improve it</a> to <a href="/wiki/Wikipedia:Make_technical_articles_understandable" title="Wikipedia:Make technical articles understandable">make it understandable to non-experts</a>, without removing the technical details.</span> <span class="date-container"><i>(<span class="date">January 2024</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> <p>Visual phototransduction in <a href="/wiki/Invertebrate" title="Invertebrate">invertebrates</a> like the <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">fruit fly</a> differs from that of vertebrates, described up to now. The primary basis of invertebrate phototransduction is the <a href="/wiki/PI(4,5)P2_Cycle" title="PI(4,5)P2 Cycle">PI(4,5)P<sub>2</sub> cycle</a>. Here, light induces the conformational change into <a href="/wiki/Rhodopsin" title="Rhodopsin">rhodopsin</a> and converts it into meta-rhodopsin. This helps in dissociation of G-protein complex. Alpha sub-unit of this complex activates the <a href="/wiki/Phospholipase_C" title="Phospholipase C">PLC</a> enzyme (PLC-beta) which hydrolyze the <a href="/wiki/Phosphatidylinositol_4,5-bisphosphate" title="Phosphatidylinositol 4,5-bisphosphate">PIP2</a> into <a href="/wiki/Diglyceride" title="Diglyceride">DAG</a>. This hydrolysis leads to opening of <a href="/wiki/Transient_receptor_potential_channel" title="Transient receptor potential channel">TRP</a> channels and influx of calcium.<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. (January 2024)">citation needed</span></a></i>]</sup> </p><p>Invertebrate <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a> differ morphologically and physiologically from their vertebrate counterparts. Visual stimulation in vertebrates causes a hyperpolarization (weakening) of the photoreceptor membrane potential, whereas invertebrates experience a depolarization with light intensity. Single-photon events produced under identical conditions in invertebrates differ from vertebrates in time course and size. Likewise, multi-photon events are longer than single-photon responses in invertebrates. However, in vertebrates, the multi-photon response is similar to the single-photon response. Both phyla have light adaptation and single-photon events are smaller and faster. Calcium plays an important role in this adaptation. Light adaptation in vertebrates is primarily attributable to calcium feedback, but in invertebrates cyclic AMP is another control on dark adaptation.<sup id="cite_ref-Rayer90_9-0" class="reference"><a href="#cite_note-Rayer90-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability"><span title="The material near this tag needs to be fact-checked with the cited source(s). (January 2024)">verification needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=9" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-Bert20-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Bert20_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Bert20_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Bert20_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Bert20_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBertalmío2020" class="citation journal cs1">Bertalmío, Marcelo (2020). "The biological basis of vision: the retina". <i>Vision Models for High Dynamic Range and Wide Colour Gamut Imaging</i>: 11–46. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FB978-0-12-813894-6.00007-7">10.1016/B978-0-12-813894-6.00007-7</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-813894-6" title="Special:BookSources/978-0-12-813894-6"><bdi>978-0-12-813894-6</bdi></a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:209571302">209571302</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Vision+Models+for+High+Dynamic+Range+and+Wide+Colour+Gamut+Imaging&rft.atitle=The+biological+basis+of+vision%3A+the+retina&rft.pages=11-46&rft.date=2020&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A209571302%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1016%2FB978-0-12-813894-6.00007-7&rft.isbn=978-0-12-813894-6&rft.aulast=Bertalm%C3%ADo&rft.aufirst=Marcelo&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEbreyKoutalos,_Yiannis2001" class="citation journal cs1">Ebrey, Thomas; Koutalos, Yiannis (January 2001). "Vertebrate Photoreceptors". <i>Progress in Retinal and Eye Research</i>. <b>20</b> (1): 49–94. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS1350-9462%2800%2900014-8">10.1016/S1350-9462(00)00014-8</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11070368">11070368</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:2789591">2789591</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Progress+in+Retinal+and+Eye+Research&rft.atitle=Vertebrate+Photoreceptors&rft.volume=20&rft.issue=1&rft.pages=49-94&rft.date=2001-01&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2789591%23id-name%3DS2CID&rft_id=info%3Apmid%2F11070368&rft_id=info%3Adoi%2F10.1016%2FS1350-9462%2800%2900014-8&rft.aulast=Ebrey&rft.aufirst=Thomas&rft.au=Koutalos%2C+Yiannis&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"> E.Y. Popova; C.J. Barnstable;(2019). "Chapter 15 - Insights Into the Epigenetics of Retinal Development and Diseases". <a rel="nofollow" class="external free" href="https://doi.org/10.1016/B978-0-12-814879-2.00016-9">https://doi.org/10.1016/B978-0-12-814879-2.00016-9</a> </span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeskovKlenchin,_Handy,_Whitlock,_Govardovskii,_Bownds,_Lamb,_Pugh,_Arshavsky2000" class="citation journal cs1">Leskov, Ilya; Klenchin, Handy, Whitlock, Govardovskii, Bownds, Lamb, Pugh, Arshavsky (September 2000). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0896-6273%2800%2900063-5">"The Gain of Rod Phototransduction: Reconciliation of Biochemical and Electrophysiological Measurements"</a>. <i>Neuron</i>. <b>27</b> (3): 525–537. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0896-6273%2800%2900063-5">10.1016/S0896-6273(00)00063-5</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11055435">11055435</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:15573966">15573966</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuron&rft.atitle=The+Gain+of+Rod+Phototransduction%3A+Reconciliation+of+Biochemical+and+Electrophysiological+Measurements&rft.volume=27&rft.issue=3&rft.pages=525-537&rft.date=2000-09&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A15573966%23id-name%3DS2CID&rft_id=info%3Apmid%2F11055435&rft_id=info%3Adoi%2F10.1016%2FS0896-6273%2800%2900063-5&rft.aulast=Leskov&rft.aufirst=Ilya&rft.au=Klenchin%2C+Handy%2C+Whitlock%2C+Govardovskii%2C+Bownds%2C+Lamb%2C+Pugh%2C+Arshavsky&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252FS0896-6273%252800%252900063-5&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span><span class="cs1-maint citation-comment"><code class="cs1-code">{{<a href="/wiki/Template:Cite_journal" title="Template:Cite journal">cite journal</a>}}</code>: CS1 maint: multiple names: authors list (<a href="/wiki/Category:CS1_maint:_multiple_names:_authors_list" title="Category:CS1 maint: multiple names: authors list">link</a>)</span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArshavskyLambPugh2002" class="citation journal cs1">Arshavsky, Vadim Y.; Lamb, Trevor D.; Pugh, Edward N. (2002). "G Proteins and Phototransduction". <i>Annual Review of Physiology</i>. <b>64</b> (1): 153–187. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.physiol.64.082701.102229">10.1146/annurev.physiol.64.082701.102229</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11826267">11826267</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Physiology&rft.atitle=G+Proteins+and+Phototransduction&rft.volume=64&rft.issue=1&rft.pages=153-187&rft.date=2002&rft_id=info%3Adoi%2F10.1146%2Fannurev.physiol.64.082701.102229&rft_id=info%3Apmid%2F11826267&rft.aulast=Arshavsky&rft.aufirst=Vadim+Y.&rft.au=Lamb%2C+Trevor+D.&rft.au=Pugh%2C+Edward+N.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHsuMolday1993" class="citation journal cs1">Hsu, Yi-Te; Molday, Robert S. (1993). "Modulation of the CGMP-gated channel of rod photoreceptor cells by calmodulin". <i>Nature</i>. <b>361</b> (6407): 76–79. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1993Natur.361...76H">1993Natur.361...76H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F361076a0">10.1038/361076a0</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/7678445">7678445</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4362581">4362581</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Modulation+of+the+CGMP-gated+channel+of+rod+photoreceptor+cells+by+calmodulin&rft.volume=361&rft.issue=6407&rft.pages=76-79&rft.date=1993&rft_id=info%3Adoi%2F10.1038%2F361076a0&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4362581%23id-name%3DS2CID&rft_id=info%3Apmid%2F7678445&rft_id=info%3Abibcode%2F1993Natur.361...76H&rft.aulast=Hsu&rft.aufirst=Yi-Te&rft.au=Molday%2C+Robert+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKrispelChenMellingChen2006" class="citation journal cs1">Krispel, CM; Chen, D; Melling, D; Chen, YJ; Martemyanov, KA; Quillinan, N; Arshavsky, VY; Wensel, TG; Chen, CK; Burns, ME (2006). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuron.2006.07.010">"RGS expression rate-limits recovery of rod photoresponses"</a>. <i>Neuron</i>. <b>51</b> (4): 409–416. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2006Neuro.51...409K">2006Neuro.51...409K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuron.2006.07.010">10.1016/j.neuron.2006.07.010</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16908407">16908407</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuron&rft.atitle=RGS+expression+rate-limits+recovery+of+rod+photoresponses.&rft.volume=51&rft.issue=4&rft.pages=409-416&rft.date=2006&rft_id=info%3Apmid%2F16908407&rft_id=info%3Adoi%2F10.1016%2Fj.neuron.2006.07.010&rft_id=info%3Abibcode%2F2006Neuro.51...409K&rft.aulast=Krispel&rft.aufirst=CM&rft.au=Chen%2C+D&rft.au=Melling%2C+D&rft.au=Chen%2C+YJ&rft.au=Martemyanov%2C+KA&rft.au=Quillinan%2C+N&rft.au=Arshavsky%2C+VY&rft.au=Wensel%2C+TG&rft.au=Chen%2C+CK&rft.au=Burns%2C+ME&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.neuron.2006.07.010&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span>.</span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMoiseyevChenTakahashiWu2005" class="citation journal cs1">Moiseyev, Gennadiy; Chen, Ying; Takahashi, Yusuke; Wu, Bill X.; Ma, Jian-xing (30 August 2005). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1194921">"RPE65 is the isomerohydrolase in the retinoid visual cycle"</a>. <i>Proceedings of the National Academy of Sciences</i>. <b>102</b> (35): 12413–12418. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2005PNAS..10212413M">2005PNAS..10212413M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0503460102">10.1073/pnas.0503460102</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1194921">1194921</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16116091">16116091</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences&rft.atitle=RPE65+is+the+isomerohydrolase+in+the+retinoid+visual+cycle&rft.volume=102&rft.issue=35&rft.pages=12413-12418&rft.date=2005-08-30&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1194921%23id-name%3DPMC&rft_id=info%3Apmid%2F16116091&rft_id=info%3Adoi%2F10.1073%2Fpnas.0503460102&rft_id=info%3Abibcode%2F2005PNAS..10212413M&rft.aulast=Moiseyev&rft.aufirst=Gennadiy&rft.au=Chen%2C+Ying&rft.au=Takahashi%2C+Yusuke&rft.au=Wu%2C+Bill+X.&rft.au=Ma%2C+Jian-xing&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1194921&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> <li id="cite_note-Rayer90-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-Rayer90_9-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRayerNaynertStieve1990" class="citation journal cs1">Rayer, B.; Naynert, M.; Stieve, H. (November 1990). "New trends in photobiology". <i>Journal of Photochemistry and Photobiology B: Biology</i>. <b>7</b> (2–4): 107–148. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2F1011-1344%2890%2985151-L">10.1016/1011-1344(90)85151-L</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/2150859">2150859</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Photochemistry+and+Photobiology+B%3A+Biology&rft.atitle=New+trends+in+photobiology&rft.volume=7&rft.issue=2%E2%80%934&rft.pages=107-148&rft.date=1990-11&rft_id=info%3Adoi%2F10.1016%2F1011-1344%2890%2985151-L&rft_id=info%3Apmid%2F2150859&rft.aulast=Rayer&rft.aufirst=B.&rft.au=Naynert%2C+M.&rft.au=Stieve%2C+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AVisual+phototransduction" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Visual_phototransduction&action=edit&section=10" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://webvision.med.utah.edu/">Visual pigments and visual transduction at med.utah.edu</a></li> <li><a rel="nofollow" class="external text" href="https://prezi.com/view/zsG4HF8oit2XgZlla33P/">Transduction of Light Prezi</a></li> <li><a rel="nofollow" class="external text" href="http://serendip.brynmawr.edu/bb/neuro/neuro00/web1/Patel.html">A General Overview on Visual Perception at brynmawr.edu</a></li> <li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Phototransduction">Phototransduction</a> at the U.S. National Library of Medicine <a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · ";font-weight:bold}.mw-parser-output .hlist dd:last-child::after,.mw-parser-output .hlist dt:last-child::after,.mw-parser-output .hlist li:last-child::after{content:none}.mw-parser-output .hlist dd dd:first-child::before,.mw-parser-output .hlist dd dt:first-child::before,.mw-parser-output .hlist dd li:first-child::before,.mw-parser-output .hlist dt dd:first-child::before,.mw-parser-output .hlist dt dt:first-child::before,.mw-parser-output .hlist dt li:first-child::before,.mw-parser-output .hlist li dd:first-child::before,.mw-parser-output .hlist li dt:first-child::before,.mw-parser-output .hlist li li:first-child::before{content:" (";font-weight:normal}.mw-parser-output .hlist dd dd:last-child::after,.mw-parser-output .hlist dd dt:last-child::after,.mw-parser-output .hlist dd li:last-child::after,.mw-parser-output .hlist dt dd:last-child::after,.mw-parser-output .hlist dt dt:last-child::after,.mw-parser-output .hlist dt li:last-child::after,.mw-parser-output .hlist li dd:last-child::after,.mw-parser-output .hlist li dt:last-child::after,.mw-parser-output .hlist li li:last-child::after{content:")";font-weight:normal}.mw-parser-output .hlist ol{counter-reset:listitem}.mw-parser-output .hlist ol>li{counter-increment:listitem}.mw-parser-output .hlist ol>li::before{content:" "counter(listitem)"\a0 "}.mw-parser-output .hlist dd ol>li:first-child::before,.mw-parser-output .hlist dt ol>li:first-child::before,.mw-parser-output .hlist li ol>li:first-child::before{content:" ("counter(listitem)"\a0 "}</style><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 1em;line-height:1.5em;text-align:center}.mw-parser-output .navbox-group{white-space:nowrap;text-align:right}.mw-parser-output .navbox,.mw-parser-output .navbox-subgroup{background-color:#fdfdfd}.mw-parser-output .navbox-list{line-height:1.5em;border-color:#fdfdfd}.mw-parser-output .navbox-list-with-group{text-align:left;border-left-width:2px;border-left-style:solid}.mw-parser-output tr+tr>.navbox-abovebelow,.mw-parser-output tr+tr>.navbox-group,.mw-parser-output tr+tr>.navbox-image,.mw-parser-output tr+tr>.navbox-list{border-top:2px solid #fdfdfd}.mw-parser-output .navbox-title{background-color:#ccf}.mw-parser-output .navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Cell_signaling_/_Signal_transduction" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Signal_transduction" title="Template:Signal transduction"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Signal_transduction" title="Template talk:Signal transduction"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Signal_transduction" title="Special:EditPage/Template:Signal transduction"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_signaling_/_Signal_transduction" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_signaling" title="Cell signaling">Cell signaling</a> / <a href="/wiki/Signal_transduction" title="Signal transduction">Signal transduction</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Signaling pathways</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/G_protein-coupled_receptor" title="G protein-coupled receptor">GPCR</a></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt</a></li> <li><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">RTK</a> <ul><li><a href="/wiki/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta</a></li> <li><a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK</a></li></ul></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch</a></li> <li><a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT</a></li> <li><a href="/wiki/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB</a></li> <li><a href="/wiki/Apoptosis" title="Apoptosis">Fas apoptosis</a></li> <li><a href="/wiki/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo</a></li> <li><a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a></li> <li><a href="/wiki/Integrin#Signal_transduction" title="Integrin">Integrin receptors</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Agents</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Receptor ligands</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hormone" title="Hormone">Hormones</a></li> <li><a href="/wiki/Neurotransmitter" title="Neurotransmitter">Neurotransmitters</a>/<a href="/wiki/Neuropeptide" title="Neuropeptide">Neuropeptides</a>/<a href="/wiki/Neurohormone" title="Neurohormone">Neurohormones</a></li> <li><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a></li> <li><a href="/wiki/Growth_factor" title="Growth factor">Growth factors</a></li> <li><a href="/wiki/Cell_signaling" title="Cell signaling">Signaling molecules</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">Cell surface</a></li> <li><a href="/wiki/Intracellular_receptor" title="Intracellular receptor">Intracellular</a></li> <li><a href="/wiki/Co-receptor" title="Co-receptor">Co-receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Second_messenger_system" title="Second messenger system">Second messenger</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP-dependent pathway</a></li> <li><a href="/wiki/Calcium_signaling" title="Calcium signaling">Ca<sup>2+</sup> signaling</a></li> <li><a href="/wiki/Lipid_signaling" title="Lipid signaling">Lipid signaling</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Assistants:</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/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">Signal transducing adaptor protein</a></li> <li><a href="/wiki/Scaffold_protein" title="Scaffold protein">Scaffold protein</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factors</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/General_transcription_factor" title="General transcription factor">General</a></li> <li><a href="/wiki/Transcription_preinitiation_complex" title="Transcription preinitiation complex">Transcription preinitiation complex</a></li> <li><a href="/wiki/Transcription_factor_II_D" title="Transcription factor II D">TFIID</a></li> <li><a href="/wiki/Transcription_factor_II_H" title="Transcription factor II H">TFIIH</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By distance</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/Juxtacrine_signalling" title="Juxtacrine signalling">Juxtacrine</a></li> <li><a href="/wiki/Autocrine_signalling" class="mw-redirect" title="Autocrine signalling">Autocrine</a> / <a href="/wiki/Paracrine_signalling" class="mw-redirect" title="Paracrine signalling">Paracrine</a></li> <li><a href="/wiki/Endocrine_system" title="Endocrine system">Endocrine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other concepts</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/Intracrine" title="Intracrine">Intracrine action</a></li> <li>Neurocrine signaling <ul><li><a href="/wiki/Synaptic_transmission" class="mw-redirect" title="Synaptic transmission">Synaptic transmission</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a></li></ul></li> <li><a href="/wiki/Neuroendocrine_cell" title="Neuroendocrine cell">Neuroendocrine signaling</a></li> <li><a href="/wiki/Exocrine_gland" title="Exocrine gland">Exocrine signalling</a> <ul><li><a href="/wiki/Pheromone" title="Pheromone">Pheromones</a></li></ul></li> <li><a href="/wiki/Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a></li> <li><a class="mw-selflink selflink">Phototransduction</a></li> <li><a href="/wiki/Ion_channel" title="Ion channel">Ion channel gating</a></li> <li><a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Visual_phototransduction&oldid=1244956549">https://en.wikipedia.org/w/index.php?title=Visual_phototransduction&oldid=1244956549</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Articles_needing_attention" title="Category:Articles needing attention">Articles needing attention</a></li><li><a href="/wiki/Category:Visual_system" title="Category:Visual system">Visual system</a></li><li><a href="/wiki/Category:Nervous_system" title="Category:Nervous system">Nervous system</a></li><li><a href="/wiki/Category:Sensory_receptors" title="Category:Sensory receptors">Sensory receptors</a></li><li><a href="/wiki/Category:Metabolism" title="Category:Metabolism">Metabolism</a></li></ul></div><div id="mw-hidden-catlinks" class="mw-hidden-catlinks mw-hidden-cats-hidden">Hidden categories: <ul><li><a href="/wiki/Category:CS1_maint:_multiple_names:_authors_list" title="Category:CS1 maint: multiple names: authors list">CS1 maint: multiple names: authors list</a></li><li><a href="/wiki/Category:Articles_with_short_description" title="Category:Articles with short description">Articles with short description</a></li><li><a href="/wiki/Category:Short_description_is_different_from_Wikidata" title="Category:Short description is different from Wikidata">Short description is different from Wikidata</a></li><li><a href="/wiki/Category:Wikipedia_articles_that_are_too_technical_from_January_2024" title="Category:Wikipedia articles that are too technical from January 2024">Wikipedia articles that are too technical from January 2024</a></li><li><a href="/wiki/Category:All_articles_that_are_too_technical" title="Category:All articles that are too technical">All articles that are too technical</a></li><li><a href="/wiki/Category:All_articles_with_unsourced_statements" title="Category:All articles with unsourced statements">All articles with unsourced statements</a></li><li><a href="/wiki/Category:Articles_with_unsourced_statements_from_January_2024" title="Category:Articles with unsourced statements from January 2024">Articles with unsourced statements from January 2024</a></li><li><a href="/wiki/Category:All_pages_needing_factual_verification" title="Category:All pages needing factual verification">All pages needing factual verification</a></li><li><a href="/wiki/Category:Wikipedia_articles_needing_factual_verification_from_January_2024" title="Category:Wikipedia articles needing factual verification from January 2024">Wikipedia articles needing factual verification from January 2024</a></li></ul></div></div> </div> </main> </div> <div class="mw-footer-container"> <footer id="footer" class="mw-footer" > <ul id="footer-info"> <li id="footer-info-lastmod"> This page was last edited on 10 September 2024, at 05:14<span class="anonymous-show"> (UTC)</span>.</li> <li id="footer-info-copyright">Text is available under the <a href="/wiki/Wikipedia:Text_of_the_Creative_Commons_Attribution-ShareAlike_4.0_International_License" title="Wikipedia:Text of the Creative Commons Attribution-ShareAlike 4.0 International License">Creative Commons Attribution-ShareAlike 4.0 License</a>; additional terms may apply. By using this site, you agree to the <a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Terms_of_Use" class="extiw" title="foundation:Special:MyLanguage/Policy:Terms of Use">Terms of Use</a> and <a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy" class="extiw" title="foundation:Special:MyLanguage/Policy:Privacy policy">Privacy Policy</a>. Wikipedia® is a registered trademark of the <a rel="nofollow" class="external text" href="https://wikimediafoundation.org/">Wikimedia Foundation, Inc.</a>, a non-profit organization.</li> </ul> <ul id="footer-places"> <li id="footer-places-privacy"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy">Privacy policy</a></li> <li id="footer-places-about"><a href="/wiki/Wikipedia:About">About Wikipedia</a></li> <li id="footer-places-disclaimers"><a href="/wiki/Wikipedia:General_disclaimer">Disclaimers</a></li> <li id="footer-places-contact"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us">Contact Wikipedia</a></li> <li id="footer-places-wm-codeofconduct"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Universal_Code_of_Conduct">Code of Conduct</a></li> <li id="footer-places-developers"><a href="https://developer.wikimedia.org">Developers</a></li> <li id="footer-places-statslink"><a href="https://stats.wikimedia.org/#/en.wikipedia.org">Statistics</a></li> <li id="footer-places-cookiestatement"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Cookie_statement">Cookie statement</a></li> <li id="footer-places-mobileview"><a href="//en.m.wikipedia.org/w/index.php?title=Visual_phototransduction&mobileaction=toggle_view_mobile" class="noprint stopMobileRedirectToggle">Mobile view</a></li> </ul> <ul id="footer-icons" class="noprint"> <li id="footer-copyrightico"><a href="https://wikimediafoundation.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/static/images/footer/wikimedia-button.svg" width="84" height="29" alt="Wikimedia Foundation" loading="lazy"></a></li> <li id="footer-poweredbyico"><a href="https://www.mediawiki.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/w/resources/assets/poweredby_mediawiki.svg" alt="Powered by MediaWiki" width="88" height="31" loading="lazy"></a></li> </ul> </footer> </div> </div> </div> <div class="vector-settings" id="p-dock-bottom"> <ul></ul> </div><script>(RLQ=window.RLQ||[]).push(function(){mw.config.set({"wgHostname":"mw-web.codfw.main-f69cdc8f6-t4zsm","wgBackendResponseTime":175,"wgPageParseReport":{"limitreport":{"cputime":"0.377","walltime":"0.472","ppvisitednodes":{"value":1483,"limit":1000000},"postexpandincludesize":{"value":55537,"limit":2097152},"templateargumentsize":{"value":4931,"limit":2097152},"expansiondepth":{"value":15,"limit":100},"expensivefunctioncount":{"value":6,"limit":500},"unstrip-depth":{"value":1,"limit":20},"unstrip-size":{"value":45119,"limit":5000000},"entityaccesscount":{"value":0,"limit":400},"timingprofile":["100.00% 414.932 1 -total"," 40.58% 168.368 1 Template:Reflist"," 35.43% 147.016 8 Template:Cite_journal"," 20.36% 84.471 3 Template:Navbox"," 19.68% 81.650 1 Template:Signal_transduction"," 12.43% 51.556 1 Template:Short_description"," 10.96% 45.487 1 Template:Needsattention"," 8.67% 35.967 1 Template:Yesno"," 8.32% 34.532 1 Template:Technical"," 7.90% 32.787 1 Template:Ambox"]},"scribunto":{"limitreport-timeusage":{"value":"0.239","limit":"10.000"},"limitreport-memusage":{"value":5917279,"limit":52428800}},"cachereport":{"origin":"mw-web.codfw.main-f69cdc8f6-nbpzk","timestamp":"20241122152050","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Visual phototransduction","url":"https:\/\/en.wikipedia.org\/wiki\/Visual_phototransduction","sameAs":"http:\/\/www.wikidata.org\/entity\/Q445081","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q445081","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2005-05-14T21:01:23Z","dateModified":"2024-09-10T05:14:40Z","headline":"sensory transduction of the visual system"}</script> </body> </html>