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class="vector-toc-list"> <li id="toc-Inverted_versus_non-inverted_retina" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Inverted_versus_non-inverted_retina"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Inverted versus non-inverted retina</span> </div> </a> <ul id="toc-Inverted_versus_non-inverted_retina-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Retinal_layers" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Retinal_layers"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Retinal layers</span> </div> </a> <ul id="toc-Retinal_layers-sublist" class="vector-toc-list"> <li id="toc-Layers_imagable_with_optical_coherence_tomography" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Layers_imagable_with_optical_coherence_tomography"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2.1</span> <span>Layers imagable with optical coherence tomography</span> </div> </a> <ul id="toc-Layers_imagable_with_optical_coherence_tomography-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Development"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Development</span> </div> </a> <ul id="toc-Development-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Blood_supply" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Blood_supply"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Blood supply</span> </div> </a> <ul id="toc-Blood_supply-sublist" class="vector-toc-list"> <li id="toc-Energy_requirements" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Energy_requirements"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.1</span> <span>Energy requirements</span> </div> </a> <ul id="toc-Energy_requirements-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_birds" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#In_birds"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4.2</span> <span>In birds</span> </div> </a> <ul id="toc-In_birds-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Biometric_identification_and_diagnosis_of_disease" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Biometric_identification_and_diagnosis_of_disease"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Biometric identification and diagnosis of disease</span> </div> </a> <ul id="toc-Biometric_identification_and_diagnosis_of_disease-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Function</span> </div> </a> <button aria-controls="toc-Function-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 Function subsection</span> </button> <ul id="toc-Function-sublist" class="vector-toc-list"> <li id="toc-Spatial_encoding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Spatial_encoding"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Spatial encoding</span> </div> </a> <ul id="toc-Spatial_encoding-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Clinical_significance" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinical_significance"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Clinical significance</span> </div> </a> <button aria-controls="toc-Clinical_significance-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 Clinical significance subsection</span> </button> <ul id="toc-Clinical_significance-sublist" class="vector-toc-list"> <li id="toc-Diagnosis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Diagnosis"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Diagnosis</span> </div> </a> <ul id="toc-Diagnosis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Treatment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Treatment"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Treatment</span> </div> </a> <ul id="toc-Treatment-sublist" class="vector-toc-list"> <li id="toc-Common_treatment_modalities" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Common_treatment_modalities"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Common treatment modalities</span> </div> </a> <ul id="toc-Common_treatment_modalities-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Uncommon_treatment_modalities" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Uncommon_treatment_modalities"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.2</span> <span>Uncommon treatment modalities</span> </div> </a> <ul id="toc-Uncommon_treatment_modalities-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-History" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#History"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Additional_images" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Additional_images"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Additional images</span> </div> </a> <ul id="toc-Additional_images-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</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" title="Table of Contents" > <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">Retina</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 75 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-75" 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">75 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Netvlies" title="Netvlies – Afrikaans" lang="af" hreflang="af" data-title="Netvlies" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B4%D8%A8%D9%83%D9%8A%D8%A9" title="شبكية – Arabic" lang="ar" hreflang="ar" data-title="شبكية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-arc mw-list-item"><a href="https://arc.wikipedia.org/wiki/%DC%A2%DC%AB%DC%92%DC%90_%DC%95%DC%A5%DC%9D%DC%A2%DC%90" title="ܢܫܒܐ ܕܥܝܢܐ – Aramaic" lang="arc" hreflang="arc" data-title="ܢܫܒܐ ܕܥܝܢܐ" data-language-autonym="ܐܪܡܝܐ" data-language-local-name="Aramaic" class="interlanguage-link-target"><span>ܐܪܡܝܐ</span></a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Retina" title="Retina – Asturian" lang="ast" hreflang="ast" data-title="Retina" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target"><span>Asturianu</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Retina" title="Retina – Azerbaijani" lang="az" hreflang="az" data-title="Retina" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%85%E0%A6%95%E0%A7%8D%E0%A6%B7%E0%A6%BF%E0%A6%AA%E0%A6%9F" title="অক্ষিপট – Bangla" lang="bn" hreflang="bn" data-title="অক্ষিপট" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%A1%D0%B5%D1%82%D0%BA%D0%B0%D0%B2%D1%96%D1%86%D0%B0" title="Сеткавіца – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Сеткавіца" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A0%D0%B5%D1%82%D0%B8%D0%BD%D0%B0" title="Ретина – Bulgarian" lang="bg" hreflang="bg" data-title="Ретина" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Mre%C5%BEnja%C4%8Da" title="Mrežnjača – Bosnian" lang="bs" hreflang="bs" data-title="Mrežnjača" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Retina" title="Retina – Catalan" lang="ca" hreflang="ca" data-title="Retina" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/S%C3%ADtnice" title="Sítnice – Czech" lang="cs" hreflang="cs" data-title="Sítnice" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Retina" title="Retina – Welsh" lang="cy" hreflang="cy" data-title="Retina" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target"><span>Cymraeg</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Nethinde" title="Nethinde – Danish" lang="da" hreflang="da" data-title="Nethinde" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-se mw-list-item"><a href="https://se.wikipedia.org/wiki/Fierbmecuozza" title="Fierbmecuozza – Northern Sami" lang="se" hreflang="se" data-title="Fierbmecuozza" data-language-autonym="Davvisámegiella" data-language-local-name="Northern Sami" class="interlanguage-link-target"><span>Davvisámegiella</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Netzhaut" title="Netzhaut – German" lang="de" hreflang="de" data-title="Netzhaut" 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/V%C3%B5rkkest" title="Võrkkest – Estonian" lang="et" hreflang="et" data-title="Võrkkest" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%91%CE%BC%CF%86%CE%B9%CE%B2%CE%BB%CE%B7%CF%83%CF%84%CF%81%CE%BF%CE%B5%CE%B9%CE%B4%CE%AE%CF%82_%CF%87%CE%B9%CF%84%CF%8E%CE%BD%CE%B1%CF%82" title="Αμφιβληστροειδής χιτώνας – Greek" lang="el" hreflang="el" data-title="Αμφιβληστροειδής χιτώνας" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Retina" title="Retina – Spanish" lang="es" hreflang="es" data-title="Retina" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Retino" title="Retino – Esperanto" lang="eo" hreflang="eo" data-title="Retino" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Erretina" title="Erretina – Basque" lang="eu" hreflang="eu" data-title="Erretina" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B4%D8%A8%DA%A9%DB%8C%D9%87" title="شبکیه – Persian" lang="fa" hreflang="fa" data-title="شبکیه" 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/R%C3%A9tine" title="Rétine – French" lang="fr" hreflang="fr" data-title="Rétine" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Reitine" title="Reitine – Irish" lang="ga" hreflang="ga" data-title="Reitine" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Retina" title="Retina – Galician" lang="gl" hreflang="gl" data-title="Retina" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%A7%9D%EB%A7%89" title="망막 – Korean" lang="ko" hreflang="ko" data-title="망막" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%A6%E0%A5%83%E0%A4%B7%E0%A5%8D%E0%A4%9F%E0%A4%BF_%E0%A4%AA%E0%A4%9F%E0%A4%B2" title="दृष्टि पटल – Hindi" lang="hi" hreflang="hi" data-title="दृष्टि पटल" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Mre%C5%BEnica_(oko)" title="Mrežnica (oko) – Croatian" lang="hr" hreflang="hr" data-title="Mrežnica (oko)" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Retino" title="Retino – Ido" lang="io" hreflang="io" data-title="Retino" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target"><span>Ido</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Retina" title="Retina – Indonesian" lang="id" hreflang="id" data-title="Retina" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Sj%C3%B3nhimna" title="Sjónhimna – Icelandic" lang="is" hreflang="is" data-title="Sjónhimna" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Retina" title="Retina – Italian" lang="it" hreflang="it" data-title="Retina" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A8%D7%A9%D7%AA%D7%99%D7%AA" title="רשתית – Hebrew" lang="he" hreflang="he" data-title="רשתית" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Retina" title="Retina – Javanese" lang="jv" hreflang="jv" data-title="Retina" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%B0%E0%B3%86%E0%B2%9F%E0%B2%BF%E0%B2%A8%E0%B2%BE" title="ರೆಟಿನಾ – Kannada" lang="kn" hreflang="kn" data-title="ರೆಟಿನಾ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target"><span>ಕನ್ನಡ</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%A2%D0%BE%D1%80%D0%BB%D1%8B_%D2%9B%D0%B0%D0%B1%D1%8B%D2%9B" title="Торлы қабық – Kazakh" lang="kk" hreflang="kk" data-title="Торлы қабық" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Retin" title="Retin – Haitian Creole" lang="ht" hreflang="ht" data-title="Retin" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/Tore" title="Tore – Kurdish" lang="ku" hreflang="ku" data-title="Tore" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target"><span>Kurdî</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%A2%D0%B0%D0%BC%D1%8B%D1%80%D0%B4%D1%83%D1%83_%D1%87%D0%B5%D0%BB" title="Тамырдуу чел – Kyrgyz" lang="ky" hreflang="ky" data-title="Тамырдуу чел" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Retina_(anatomia)" title="Retina (anatomia) – Latin" lang="la" hreflang="la" data-title="Retina (anatomia)" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/T%C4%ABklene" title="Tīklene – Latvian" lang="lv" hreflang="lv" data-title="Tīklene" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Tinklain%C4%97" title="Tinklainė – Lithuanian" lang="lt" hreflang="lt" data-title="Tinklainė" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Receh%C3%A1rtya" title="Recehártya – Hungarian" lang="hu" hreflang="hu" data-title="Recehártya" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%9C%D1%80%D0%B5%D0%B6%D0%BD%D0%B8%D1%86%D0%B0" title="Мрежница – Macedonian" lang="mk" hreflang="mk" data-title="Мрежница" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B1%E0%B5%86%E0%B4%B1%E0%B5%8D%E0%B4%B1%E0%B4%BF%E0%B4%A8" title="റെറ്റിന – Malayalam" lang="ml" hreflang="ml" data-title="റെറ്റിന" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%A6%E0%A5%83%E0%A4%B7%E0%A5%8D%E0%A4%9F%E0%A4%BF%E0%A4%AA%E0%A4%9F%E0%A4%B2" title="दृष्टिपटल – Marathi" lang="mr" hreflang="mr" data-title="दृष्टिपटल" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target"><span>मराठी</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Retina" title="Retina – Malay" lang="ms" hreflang="ms" data-title="Retina" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%99%E1%80%BB%E1%80%80%E1%80%BA%E1%80%80%E1%80%BC%E1%80%B1%E1%80%AC%E1%80%9C%E1%80%BD%E1%80%BE%E1%80%AC" title="မျက်ကြောလွှာ – Burmese" lang="my" hreflang="my" data-title="မျက်ကြောလွှာ" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target"><span>မြန်မာဘာသာ</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Netvlies" title="Netvlies – Dutch" lang="nl" hreflang="nl" data-title="Netvlies" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%B6%B2%E8%86%9C" title="網膜 – Japanese" lang="ja" hreflang="ja" data-title="網膜" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Netthinne" title="Netthinne – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Netthinne" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Siatk%C3%B3wka_(anatomia)" title="Siatkówka (anatomia) – Polish" lang="pl" hreflang="pl" data-title="Siatkówka (anatomia)" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Retina" title="Retina – Portuguese" lang="pt" hreflang="pt" data-title="Retina" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Retin%C4%83" title="Retină – Romanian" lang="ro" hreflang="ro" data-title="Retină" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A1%D0%B5%D1%82%D1%87%D0%B0%D1%82%D0%BA%D0%B0" title="Сетчатка – Russian" lang="ru" hreflang="ru" data-title="Сетчатка" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Retina" title="Retina – Albanian" lang="sq" hreflang="sq" data-title="Retina" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Retina" title="Retina – Simple English" lang="en-simple" hreflang="en-simple" data-title="Retina" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Sietnica" title="Sietnica – Slovak" lang="sk" hreflang="sk" data-title="Sietnica" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Mre%C5%BEnica" title="Mrežnica – Slovenian" lang="sl" hreflang="sl" data-title="Mrežnica" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AA%DB%86%DA%95%DB%8C%D9%86%DB%95" title="تۆڕینە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="تۆڕینە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9C%D1%80%D0%B5%D0%B6%D1%9A%D0%B0%D1%87%D0%B0" title="Мрежњача – Serbian" lang="sr" hreflang="sr" data-title="Мрежњача" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Mre%C5%BEnja%C4%8Da" title="Mrežnjača – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Mrežnjača" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/R%C3%A9tina" title="Rétina – Sundanese" lang="su" hreflang="su" data-title="Rétina" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target"><span>Sunda</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Verkkokalvo" title="Verkkokalvo – Finnish" lang="fi" hreflang="fi" data-title="Verkkokalvo" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/N%C3%A4thinna" title="Näthinna – Swedish" lang="sv" hreflang="sv" data-title="Näthinna" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%B5%E0%AE%BF%E0%AE%B4%E0%AE%BF%E0%AE%A4%E0%AF%8D%E0%AE%A4%E0%AE%BF%E0%AE%B0%E0%AF%88" title="விழித்திரை – Tamil" lang="ta" hreflang="ta" data-title="விழித்திரை" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target"><span>தமிழ்</span></a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%A7%D0%B5%D0%BB%D1%82%D3%99%D1%80%D0%BA%D0%B0%D1%82%D0%BB%D0%B0%D1%83" title="Челтәркатлау – Tatar" lang="tt" hreflang="tt" data-title="Челтәркатлау" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target"><span>Татарча / tatarça</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%88%E0%B8%AD%E0%B8%95%E0%B8%B2" title="จอตา – Thai" lang="th" hreflang="th" data-title="จอตา" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Retina" title="Retina – Turkish" lang="tr" hreflang="tr" data-title="Retina" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D1%96%D1%82%D0%BA%D1%96%D0%B2%D0%BA%D0%B0" title="Сітківка – Ukrainian" lang="uk" hreflang="uk" data-title="Сітківка" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B4%D8%A8%DA%A9%DB%8C%DB%81" title="شبکیہ – Urdu" lang="ur" hreflang="ur" data-title="شبکیہ" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/V%C3%B5ng_m%E1%BA%A1c" title="Võng mạc – Vietnamese" lang="vi" hreflang="vi" data-title="Võng mạc" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E8%A7%86%E7%BD%91%E8%86%9C" title="视网膜 – Wu" lang="wuu" hreflang="wuu" data-title="视网膜" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target"><span>吴语</span></a></li><li class="interlanguage-link interwiki-yi mw-list-item"><a href="https://yi.wikipedia.org/wiki/%D7%A8%D7%A2%D7%98%D7%99%D7%A0%D7%A2" title="רעטינע – Yiddish" lang="yi" hreflang="yi" data-title="רעטינע" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target"><span>ייִדיש</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E8%A6%96%E7%B6%B2%E8%86%9C" title="視網膜 – Cantonese" lang="yue" hreflang="yue" data-title="視網膜" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E8%A7%86%E7%BD%91%E8%86%9C" title="视网膜 – Chinese" lang="zh" hreflang="zh" data-title="视网膜" 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 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class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Part of the eye</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">For the electronics screen trademark, see <a href="/wiki/Retina_display" title="Retina display">Retina display</a>. For other uses, see <a href="/wiki/Retina_(disambiguation)" class="mw-disambig" title="Retina (disambiguation)">Retina (disambiguation)</a>.</div> <p class="mw-empty-elt"> </p> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:dimgray; color: white">Retina</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Human_eye_cross-sectional_view_grayscale.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ed/Human_eye_cross-sectional_view_grayscale.png/250px-Human_eye_cross-sectional_view_grayscale.png" decoding="async" width="250" height="179" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/ed/Human_eye_cross-sectional_view_grayscale.png 1.5x" data-file-width="350" data-file-height="250" /></a></span><div class="infobox-caption">Right <a href="/wiki/Human_eye" title="Human eye">human eye</a> cross-sectional view; eyes vary significantly among animals.</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Pronunciation</th><td class="infobox-data"><span class="rt-commentedText nowrap"><style data-mw-deduplicate="TemplateStyles:r1177148991">.mw-parser-output .IPA-label-small{font-size:85%}.mw-parser-output .references .IPA-label-small,.mw-parser-output .infobox .IPA-label-small,.mw-parser-output .navbox .IPA-label-small{font-size:100%}</style><span class="IPA-label IPA-label-small"><a href="/wiki/British_English" title="British English">UK</a>: </span><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="/ˈ/: primary stress follows">ˈ</span><span title="'r' in 'rye'">r</span><span title="/ɛ/: 'e' in 'dress'">ɛ</span><span title="'t' in 'tie'">t</span><span title="/ɪ/: 'i' in 'kit'">ɪ</span><span title="'n' in 'nigh'">n</span><span title="/ə/: 'a' in 'about'">ə</span></span>/</a></span></span>,<br /><span class="rt-commentedText nowrap"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1177148991" /><span class="IPA-label IPA-label-small"><a href="/wiki/American_English" title="American English">US</a>: </span><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/<span style="border-bottom:1px dotted"><span title="/ˈ/: primary stress follows">ˈ</span><span title="'r' in 'rye'">r</span><span title="/ɛ/: 'e' in 'dress'">ɛ</span><span title="'t' in 'tie'">t</span><span title="/ən/: 'on' in 'button'">ən</span><span title="/ə/: 'a' in 'about'">ə</span></span>/</a></span></span>,<br /><a href="/wiki/Plural" title="Plural">pl.</a> retinae <span class="rt-commentedText nowrap"><span class="IPA nopopups noexcerpt" lang="en-fonipa"><a href="/wiki/Help:IPA/English" title="Help:IPA/English">/-<span style="border-bottom:1px dotted"><span title="'n' in 'nigh'">n</span><span title="/i/: 'y' in 'happy'">i</span></span>/</a></span></span></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Part of</th><td class="infobox-data"><a href="/wiki/Eye" title="Eye">Eye</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Organ_system" title="Organ system">System</a></th><td class="infobox-data"><a href="/wiki/Visual_system" title="Visual system">Visual system</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Artery" title="Artery">Artery</a></th><td class="infobox-data"><a href="/wiki/Central_retinal_artery" title="Central retinal artery">Central retinal artery</a></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>rēte, tunica interna bulbi</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Medical_Subject_Headings" title="Medical Subject Headings">MeSH</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?ui=D012160">D012160</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Anatomica" title="Terminologia Anatomica">TA98</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ifaa.unifr.ch/Public/EntryPage/TA98%20Tree/Entity%20TA98%20EN/15.2.04.002%20Entity%20TA98%20EN.htm">A15.2.04.002</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Terminologia_Anatomica" title="Terminologia Anatomica">TA2</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://ta2viewer.openanatomy.org/?id=6776">6776</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma58301">58301</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terminology" title="Anatomical terminology">Anatomical terminology</a><div style="text-align: right;"><small class="noprint">[<a href="https://www.wikidata.org/wiki/Q169342" class="extiw" title="d:Q169342">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>The <b>retina</b> (from <a href="/wiki/Latin_language" class="mw-redirect" title="Latin language">Latin</a> <i> <i lang="la"><a href="https://en.wiktionary.org/wiki/rete#Latin" class="extiw" title="wikt:rete">rete</a></i></i> <span class="gloss-quot">'</span><span class="gloss-text">net</span><span class="gloss-quot">'</span>; <abbr title="plural">pl.</abbr><span style="white-space:nowrap;"> <b>retinae</b></span> or <b>retinas</b>) is the innermost, <a href="/wiki/Photosensitivity" title="Photosensitivity">light-sensitive</a> layer of <a href="/wiki/Tissue_(biology)" title="Tissue (biology)">tissue</a> of the <a href="/wiki/Eye" title="Eye">eye</a> of most <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> and some <a href="/wiki/Mollusca" title="Mollusca">molluscs</a>. The optics of the eye create a <a href="/wiki/Focus_(optics)" title="Focus (optics)">focused</a> two-dimensional image of the visual world on the retina, which then processes that image within the retina and sends nerve impulses along the <a href="/wiki/Optic_nerve" title="Optic nerve">optic nerve</a> to the <a href="/wiki/Visual_cortex" title="Visual cortex">visual cortex</a> to create <a href="/wiki/Visual_perception" title="Visual perception">visual perception</a>. The retina serves a function which is in many ways analogous to that of the <a href="/wiki/Photographic_film" title="Photographic film">film</a> or <a href="/wiki/Image_sensor" title="Image sensor">image sensor</a> in a <a href="/wiki/Camera" title="Camera">camera</a>. </p><p>The neural retina consists of several layers of <a href="/wiki/Neuron" title="Neuron">neurons</a> interconnected by <a href="/wiki/Chemical_synapse" title="Chemical synapse">synapses</a> and is supported by an outer layer of pigmented epithelial cells. The primary light-sensing cells in the retina are the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a>, which are of two types: <a href="/wiki/Rod_cell" title="Rod cell">rods</a> and <a href="/wiki/Cone_cell" title="Cone cell">cones</a>. Rods function mainly in dim light and provide monochromatic vision. Cones function in well-lit conditions and are responsible for the perception of colour through the use of a range of <a href="/wiki/Opsin" title="Opsin">opsins</a>, as well as high-acuity vision used for tasks such as reading. A third type of light-sensing cell, the <a href="/wiki/Intrinsically_photosensitive_retinal_ganglion_cells" class="mw-redirect" title="Intrinsically photosensitive retinal ganglion cells">photosensitive ganglion cell</a>, is important for <a href="/wiki/Entrainment_(chronobiology)" title="Entrainment (chronobiology)">entrainment</a> of <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">circadian rhythms</a> and reflexive responses such as the <a href="/wiki/Pupillary_light_reflex" title="Pupillary light reflex">pupillary light reflex</a>. </p><p>Light striking the retina initiates a cascade of chemical and electrical events that ultimately trigger nerve impulses that are sent to various visual centres of the <a href="/wiki/Brain" title="Brain">brain</a> through the fibres of the <a href="/wiki/Optic_nerve" title="Optic nerve">optic nerve</a>. Neural signals from the rods and cones undergo processing by other neurons, whose output takes the form of <a href="/wiki/Action_potential" title="Action potential">action potentials</a> in <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">retinal ganglion cells</a> whose <a href="/wiki/Axon" title="Axon">axons</a> form the optic nerve.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>In vertebrate <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a>, the retina and the optic nerve originate as outgrowths of the developing brain, specifically the embryonic <a href="/wiki/Diencephalon" title="Diencephalon">diencephalon</a>; thus, the retina is considered part of the <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> (CNS) and is actually brain tissue.<sup id="cite_ref-eb_2-0" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><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> It is the only part of the CNS that can be visualized <a href="/wiki/Invasiveness_of_surgical_procedures#Non-invasive_procedure" class="mw-redirect" title="Invasiveness of surgical procedures">noninvasively</a>. Like most of the brain, the retina is isolated from the vascular system by the <a href="/wiki/Blood%E2%80%93brain_barrier" title="Blood–brain barrier">blood–brain barrier</a>. The retina is the part of the body with the greatest continuous energy demand.<sup id="cite_ref-Viegas_Neuhauss_4-0" class="reference"><a href="#cite_note-Viegas_Neuhauss-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Structure">Structure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Inverted_versus_non-inverted_retina">Inverted versus non-inverted retina</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=2" title="Edit section: Inverted versus non-inverted retina"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The vertebrate retina is <i>inverted</i> in the sense that the light-sensing cells are in the back of the retina, so that light has to pass through layers of neurons and capillaries before it reaches the photosensitive sections of the rods and cones.<sup id="cite_ref-Kolb_5-0" class="reference"><a href="#cite_note-Kolb-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The ganglion cells, whose axons form the optic nerve, are at the front of the retina; therefore, the optic nerve must cross through the retina en route to the brain. No photoreceptors are in this region, giving rise to the <a href="/wiki/Blind_spot_(vision)" title="Blind spot (vision)">blind spot</a>.<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> In contrast, in the <a href="/wiki/Cephalopod" title="Cephalopod">cephalopod</a> retina, the photoreceptors are in front, with processing neurons and capillaries behind them. Because of this, cephalopods do not have a blind spot. </p><p>Although the overlying neural tissue is partly transparent, and the accompanying <a href="/wiki/Glial_cell" class="mw-redirect" title="Glial cell">glial cells</a> have been shown to act as <a href="/wiki/Optical_fiber" title="Optical fiber">fibre-optic</a> channels to transport photons directly to the photoreceptors,<sup id="cite_ref-pmid17485670_7-0" class="reference"><a href="#cite_note-pmid17485670-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><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> <a href="/wiki/Light_scattering" class="mw-redirect" title="Light scattering">light scattering</a> does occur.<sup id="cite_ref-Bringmann2018_9-0" class="reference"><a href="#cite_note-Bringmann2018-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Some vertebrates, including humans, have an area of the central retina adapted for high-acuity vision. This area, termed the <a href="/wiki/Fovea_centralis" title="Fovea centralis">fovea centralis</a>, is avascular (does not have blood vessels), and has minimal neural tissue in front of the photoreceptors, thereby minimizing light scattering.<sup id="cite_ref-Bringmann2018_9-1" class="reference"><a href="#cite_note-Bringmann2018-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>The cephalopods have a non-inverted retina, which is comparable in <a href="/wiki/Angular_resolution" title="Angular resolution">resolving power</a> to the eyes of many vertebrates. Squid eyes do not have an analog of the vertebrate <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a> (RPE). Although their photoreceptors contain a protein, retinochrome, that recycles retinal and replicates one of the functions of the vertebrate RPE, cephalopod photoreceptors are likely not maintained as well as in vertebrates, and that as a result, the useful lifetime of photoreceptors in invertebrates is much shorter than in vertebrates.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Having easily replaced stalk eyes (some lobsters) or retinae (some spiders, such as<i> Deinopis</i><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>) rarely occurs. </p><p>The cephalopod retina does not originate as an outgrowth of the brain, as the vertebrate one does. This difference suggests that vertebrate and cephalopod eyes are not <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous</a>, but have evolved separately. From an evolutionary perspective, a more complex structure such as the inverted retina can generally come about as a consequence of two alternate processes - an advantageous "good" compromise between competing functional limitations, or as a historical maladaptive relic of the convoluted path of organ evolution and transformation. Vision is an important adaptation in higher vertebrates. </p><p>A third view of the "inverted" vertebrate eye is that it combines two benefits - the maintenance of the photoreceptors mentioned above, and the reduction in light intensity necessary to avoid blinding the photoreceptors, which are based on the extremely sensitive eyes of the ancestors of modern hagfish (fish that live in very deep, dark water).<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>A recent study on the evolutionary purpose for the inverted retina structure from the APS (American Physical Society)<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> says that "The directional of glial cells helps increase the clarity of human vision. But we also noticed something rather curious: the colours that best passed through the glial cells were green to red, which the eye needs most for daytime vision. The eye usually receives too much blue—and thus has fewer blue-sensitive cones. </p><p>Further computer simulations showed that green and red are concentrated five to ten times more by the glial cells, and into their respective cones, than blue light. Instead, excess blue light gets scattered to the surrounding rods. This optimization is such that color vision during the day is enhanced, while night-time vision suffers very little". </p> <div class="mw-heading mw-heading3"><h3 id="Retinal_layers">Retinal layers</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=3" title="Edit section: Retinal layers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Gray881.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/Gray881.png/300px-Gray881.png" decoding="async" width="300" height="201" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/Gray881.png/450px-Gray881.png 1.5x, //upload.wikimedia.org/wikipedia/commons/5/55/Gray881.png 2x" data-file-width="500" data-file-height="335" /></a><figcaption>Section of retina</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Retina-diagram.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Retina-diagram.svg/300px-Retina-diagram.svg.png" decoding="async" width="300" height="134" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Retina-diagram.svg/450px-Retina-diagram.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Retina-diagram.svg/600px-Retina-diagram.svg.png 2x" data-file-width="743" data-file-height="331" /></a><figcaption>Rods, cones, and nerve layers in the retina: The front (anterior) of the eye is on the left. Light (from the left) passes through several transparent nerve layers to reach the rods and cones (far right). Chemical changes in the rods and cones send a signal back to the nerves. The signal goes first to the <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar</a> and <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">horizontal cells</a> (yellow layer), then to the <a href="/wiki/Retina_amacrine_cell" class="mw-redirect" title="Retina amacrine cell">amacrine cells</a> and <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion cells</a> (purple layer), then to the optic nerve fibres. The signals are processed in these layers. First, the signals start as raw outputs of points in the rod and cone cells. Then, the nerve layers identify simple shapes, such as bright points surrounded by dark points, edges, and movement. (Based on a drawing by <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Ramón y Cajal</a>, 1911)</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:ConeMosaics.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a6/ConeMosaics.jpg/300px-ConeMosaics.jpg" decoding="async" width="300" height="169" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a6/ConeMosaics.jpg/450px-ConeMosaics.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/a/a6/ConeMosaics.jpg 2x" data-file-width="600" data-file-height="338" /></a><figcaption>Illustration of the distribution of cone cells in the fovea of an individual with normal colour vision (left), and a colourblind (protanopic) retina. The center of the fovea holds very few blue-sensitive cones.</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Human_photoreceptor_distribution.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Human_photoreceptor_distribution.svg/300px-Human_photoreceptor_distribution.svg.png" decoding="async" width="300" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Human_photoreceptor_distribution.svg/450px-Human_photoreceptor_distribution.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Human_photoreceptor_distribution.svg/600px-Human_photoreceptor_distribution.svg.png 2x" data-file-width="512" data-file-height="512" /></a><figcaption>Distribution of rods and cones along a line passing through the fovea and the blind spot of a human eye<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>The vertebrate retina has 10 distinct layers.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> From closest to farthest from the vitreous body: </p> <ol><li><a href="/wiki/Inner_limiting_membrane" class="mw-redirect" title="Inner limiting membrane">Inner limiting membrane</a> – basement membrane elaborated by <a href="/wiki/Muller_glia" class="mw-redirect" title="Muller glia">Müller cells</a></li> <li><a href="/wiki/Nerve_fiber_layer" class="mw-redirect" title="Nerve fiber layer">Nerve fiber layer</a> – axons of the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion cell</a> bodies (a thin layer of Müller cell footplates exists between this layer and the inner limiting membrane)</li> <li><a href="/wiki/Ganglion_cell_layer" title="Ganglion cell layer">Ganglion cell layer</a> – contains nuclei of ganglion cells, the axons of which become the optic nerve fibres, and some displaced <a href="/wiki/Retina_amacrine_cell" class="mw-redirect" title="Retina amacrine cell">amacrine cells</a><sup id="cite_ref-eb_2-1" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Inner_plexiform_layer" title="Inner plexiform layer">Inner plexiform layer</a> – contains the synapse between the <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar cell</a> axons and the dendrites of the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion</a> and amacrine cells<sup id="cite_ref-eb_2-2" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Inner_nuclear_layer" title="Inner nuclear layer">Inner nuclear layer</a> – contains the nuclei and surrounding cell bodies (perikarya) of the <a href="/wiki/Amacrine_cells" class="mw-redirect" title="Amacrine cells">amacrine cells</a>, <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar cells</a>, and <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">horizontal cells</a><sup id="cite_ref-eb_2-3" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Outer_plexiform_layer" title="Outer plexiform layer">Outer plexiform layer</a> – projections of rods and cones ending in the rod spherule and cone pedicle, respectively, these make synapses with dendrites of bipolar cells and horizontal cells.<sup id="cite_ref-eb_2-4" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/Macula" title="Macula">macular</a> region, this is known as the f<i>iber layer of <a href="/wiki/Friedrich_Gustav_Jakob_Henle" title="Friedrich Gustav Jakob Henle">Henle</a></i>.</li> <li><a href="/wiki/Outer_nuclear_layer" title="Outer nuclear layer">Outer nuclear layer</a> – cell bodies of rods and cones</li> <li><a href="/wiki/External_limiting_membrane" title="External limiting membrane">External limiting membrane</a> – layer that separates the inner segment portions of the photoreceptors from their cell nuclei</li> <li><a href="/wiki/Layer_of_rods_and_cones" title="Layer of rods and cones">Inner segment / outer segment layer</a> – inner segments and outer segments of rods and cones, the outer segments contain a highly specialized light-sensing apparatus.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">Retinal pigment epithelium</a> – single layer of cuboidal epithelial cells (with extrusions not shown in diagram). This layer is closest to the choroid, and provides nourishment and supportive functions to the neural retina, The black pigment melanin in the pigment layer prevents light reflection throughout the globe of the eyeball; this is extremely important for clear vision.<sup id="cite_ref-Guyton_and_Hall_Physiology_18-0" class="reference"><a href="#cite_note-Guyton_and_Hall_Physiology-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li></ol> <p>These layers can be grouped into four main processing stages—photoreception; transmission to <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar cells</a>; transmission to <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion cells</a>, which also contain photoreceptors, the <a href="/wiki/Photosensitive_ganglion_cell" class="mw-redirect" title="Photosensitive ganglion cell">photosensitive ganglion cells</a>; and transmission along the optic nerve. At each synaptic stage, <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">horizontal</a> and <a href="/wiki/Retina_amacrine_cell" class="mw-redirect" title="Retina amacrine cell">amacrine cells</a> also are laterally connected. </p><p>The <a href="/wiki/Optic_nerve" title="Optic nerve">optic nerve</a> is a central tract of many axons of ganglion cells connecting primarily to the <a href="/wiki/Lateral_geniculate_body" class="mw-redirect" title="Lateral geniculate body">lateral geniculate body</a>, a visual relay station in the <a href="/wiki/Diencephalon" title="Diencephalon">diencephalon</a> (the rear of the forebrain). It also projects to the <a href="/wiki/Superior_colliculus" title="Superior colliculus">superior colliculus</a>, the <a href="/wiki/Suprachiasmatic_nucleus" title="Suprachiasmatic nucleus">suprachiasmatic nucleus</a>, and the <a href="/w/index.php?title=Nucleus_of_the_optic_tract&action=edit&redlink=1" class="new" title="Nucleus of the optic tract (page does not exist)">nucleus of the optic tract</a>. It passes through the other layers, creating the <a href="/wiki/Optic_disc" title="Optic disc">optic disc</a> in primates.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Additional structures, not directly associated with vision, are found as outgrowths of the retina in some vertebrate groups. In <a href="/wiki/Bird" title="Bird">birds</a>, the <a href="/wiki/Pecten_oculi" title="Pecten oculi">pecten</a> is a vascular structure of complex shape that projects from the retina into the <a href="/wiki/Vitreous_humour" class="mw-redirect" title="Vitreous humour">vitreous humour</a>; it supplies oxygen and nutrients to the eye, and may also aid in vision. <a href="/wiki/Reptile" title="Reptile">Reptiles</a> have a similar, but much simpler, structure.<sup id="cite_ref-VB_22-0" class="reference"><a href="#cite_note-VB-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p><p>In adult humans, the entire retina is about 72% of a sphere about 22 mm in diameter. The entire retina contains about 7 million cones and 75 to 150 million rods. The optic disc, a part of the retina sometimes called "the blind spot" because it lacks photoreceptors, is located at the <a href="/wiki/Optic_disc" title="Optic disc">optic papilla</a>, where the optic-nerve fibres leave the eye. It appears as an oval white area of 3 mm<sup>2</sup>. Temporal (in the direction of the temples) to this disc is the <a href="/wiki/Macula" title="Macula">macula</a>, at whose centre is the <a href="/wiki/Fovea_centralis" title="Fovea centralis">fovea</a>, a pit that is responsible for sharp central vision, but is actually less sensitive to light because of its lack of rods. Human and non-human <a href="/wiki/Primate" title="Primate">primates</a> possess one fovea, as opposed to certain bird species, such as hawks, that are bifoviate, and dogs and cats, that possess no fovea, but a central band known as the visual streak.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="Citations are needed for some of these statements, esp. bifoviate hawks and the lack of fovea in cats and dogs (March 2018)">citation needed</span></a></i>]</sup> Around the fovea extends the central retina for about 6 mm and then the peripheral retina. The farthest edge of the retina is defined by the <a href="/wiki/Ora_serrata" title="Ora serrata">ora serrata</a>. The distance from one ora to the other (or macula), the most sensitive area along the <a href="/wiki/Meridian_(perimetry,_visual_field)" title="Meridian (perimetry, visual field)">horizontal meridian</a>, is about 32 mm.<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="sentence is confusing; is all the area between the ora the macula, and are we talking length of curvature or that of a straight line? (April 2018)">clarification needed</span></a></i>]</sup> </p><p>In section, the retina is no more than 0.5 mm thick. It has three layers of <a href="/wiki/Nerve" title="Nerve">nerve</a> cells and two of <a href="/wiki/Synapse" title="Synapse">synapses</a>, including the unique <a href="/wiki/Ribbon_synapse" title="Ribbon synapse">ribbon synapse</a>. The optic nerve carries the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion-cell</a> <a href="/wiki/Axon" title="Axon">axons</a> to the brain, and the blood vessels that supply the retina. The ganglion cells lie innermost in the eye while the photoreceptive cells lie beyond. Because of this counter-intuitive arrangement, light must first pass through and around the ganglion cells and through the thickness of the retina, (including its capillary vessels, not shown) before reaching the rods and cones. Light is absorbed by the <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a> or the <a href="/wiki/Choroid" title="Choroid">choroid</a> (both of which are opaque). </p><p>The <a href="/wiki/White_blood_cell" title="White blood cell">white blood cells</a> in the <a href="/wiki/Capillaries" class="mw-redirect" title="Capillaries">capillaries</a> in front of the photoreceptors can be perceived as tiny bright moving dots when looking into blue light. This is known as the <a href="/wiki/Blue_field_entoptic_phenomenon" title="Blue field entoptic phenomenon">blue field entoptic phenomenon</a> (or Scheerer's phenomenon). </p><p>Between the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion-cell</a> layer and the rods and cones are two layers of <a href="/wiki/Neuropil" title="Neuropil">neuropils</a>, where synaptic contacts are made. The neuropil layers are the <a href="/wiki/Outer_plexiform_layer" title="Outer plexiform layer">outer plexiform layer</a> and the <a href="/wiki/Inner_plexiform_layer" title="Inner plexiform layer">inner plexiform layer</a>. In the outer neuropil layer, the rods and cones connect to the vertically running <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar cells</a>, and the horizontally oriented <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">horizontal cells</a> connect to ganglion cells. </p><p>The central retina predominantly contains cones, while the peripheral retina predominantly contains rods. In total, the retina has about seven million cones and a hundred million rods. At the centre of the macula is the foveal pit where the cones are narrow and long, and arranged in a hexagonal <a href="/wiki/Retinal_mosaic" title="Retinal mosaic">mosaic</a>, the most dense, in contradistinction to the much fatter cones located more peripherally in the retina.<sup id="cite_ref-ReferenceA_23-0" class="reference"><a href="#cite_note-ReferenceA-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> At the foveal pit, the other retinal layers are displaced, before building up along the foveal slope until the rim of the fovea, or <a href="/wiki/Parafovea" title="Parafovea">parafovea</a>, is reached, which is the thickest portion of the retina. The macula has a yellow pigmentation, from screening pigments, and is known as the macula lutea. The area directly surrounding the fovea has the highest density of rods converging on single bipolar cells. Since its cones have a much lesser convergence of signals, the fovea allows for the sharpest vision the eye can attain.<sup id="cite_ref-eb_2-5" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>Though the rod and cones are a <a href="/wiki/Retinal_mosaic" title="Retinal mosaic">mosaic</a> of sorts, transmission from receptors, to bipolars, to <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">ganglion cells</a> is not direct. Since about 150 million receptors and only 1 million optic nerve fibres exist, convergence and thus mixing of signals must occur. Moreover, the horizontal action of the <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">horizontal</a> and <a href="/wiki/Retina_amacrine_cell" class="mw-redirect" title="Retina amacrine cell">amacrine cells</a> can allow one area of the retina to control another (e.g. one stimulus inhibiting another). This inhibition is key to lessening the sum of messages sent to the higher regions of the brain. In some lower vertebrates (e.g. the <a href="/wiki/Pigeon" class="mw-redirect" title="Pigeon">pigeon</a>), control of messages is "centrifugal" – that is, one layer can control another, or higher regions of the brain can drive the retinal nerve cells, but in primates, this does not occur.<sup id="cite_ref-eb_2-6" class="reference"><a href="#cite_note-eb-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Layers_imagable_with_optical_coherence_tomography">Layers imagable with optical coherence tomography</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=4" title="Edit section: Layers imagable with optical coherence tomography"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Using <a href="/wiki/Optical_coherence_tomography" title="Optical coherence tomography">optical coherence tomography</a> (OCT), at least 13 layers can be identified in the retina. The layers and anatomical correlation are:<sup id="cite_ref-sciencedirect.com_24-0" class="reference"><a href="#cite_note-sciencedirect.com-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_25-0" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Retina-OCT800.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Retina-OCT800.png/220px-Retina-OCT800.png" decoding="async" width="220" height="69" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Retina-OCT800.png/330px-Retina-OCT800.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Retina-OCT800.png/440px-Retina-OCT800.png 2x" data-file-width="800" data-file-height="250" /></a><figcaption>Time-Domain OCT of the macular area of a retina at 800 nm, axial resolution 3 μm</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:SD-OCT_Macula_Cross-Section.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9f/SD-OCT_Macula_Cross-Section.png/220px-SD-OCT_Macula_Cross-Section.png" decoding="async" width="220" height="65" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9f/SD-OCT_Macula_Cross-Section.png/330px-SD-OCT_Macula_Cross-Section.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9f/SD-OCT_Macula_Cross-Section.png/440px-SD-OCT_Macula_Cross-Section.png 2x" data-file-width="1528" data-file-height="453" /></a><figcaption>Spectral-Domain OCT macula cross-section scan</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Macula_Histology_OCT.jpg" class="mw-file-description"><img alt="macula histology (OCT)" src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Macula_Histology_OCT.jpg/220px-Macula_Histology_OCT.jpg" decoding="async" width="220" height="253" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Macula_Histology_OCT.jpg/330px-Macula_Histology_OCT.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Macula_Histology_OCT.jpg/440px-Macula_Histology_OCT.jpg 2x" data-file-width="2000" data-file-height="2299" /></a><figcaption>Macula histology (OCT)</figcaption></figure> <p>From innermost to outermost, the layers identifiable by OCT are as follows: </p> <table class="wikitable"> <tbody><tr> <th>#</th> <th>OCT Layer / Conventional Label</th> <th>Anatomical Correlate </th> <th><a href="/wiki/Reflectivity" class="mw-redirect" title="Reflectivity">Reflectivity</a> <p><a href="/wiki/Optical_coherence_tomography" title="Optical coherence tomography">on OCT</a> </p> </th> <th>Specific <p>anatomical </p><p>boundaries? </p> </th> <th>Additional <p>references </p> </th></tr> <tr> <td>1 </td> <td><a href="/wiki/Vitreous_body" title="Vitreous body">Posterior cortical vitreous</a> </td> <td>Posterior cortical vitreous </td> <td>Hyper-reflective </td> <td>Yes </td> <td><sup id="cite_ref-:0_25-1" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><i>2</i> </td> <td><i><a href="/w/index.php?title=Preretinal_space&action=edit&redlink=1" class="new" title="Preretinal space (page does not exist)">Preretinal space</a></i> </td> <td><i>In eyes where the <a href="/wiki/Vitreous_body" title="Vitreous body">vitreous</a> has fully or partially <a href="/wiki/Posterior_vitreous_detachment" title="Posterior vitreous detachment">detached</a> from the retina, this is the space created between the posterior cortical vitreous face and the internal limiting membrane of the retina.</i> </td> <td><i>Hypo-reflective</i> </td> <td> </td> <td><i><sup id="cite_ref-:0_25-2" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></i> </td></tr> <tr> <td rowspan="2">3</td> <td><a href="/wiki/Internal_limiting_membrane" title="Internal limiting membrane">Internal limiting membrane</a> (ILM)</td> <td>Formed by <a href="/wiki/M%C3%BCller_cell" class="mw-redirect" title="Müller cell">Müller cell</a> endfeet <p><i>(unclear if it can be observed on OCT)</i> </p> </td> <td rowspan="2">Hyper-reflective </td> <td rowspan="2">No </td> <td rowspan="2"><sup id="cite_ref-:0_25-3" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><a href="/wiki/Nerve_fiber_layer" class="mw-redirect" title="Nerve fiber layer">Nerve fiber layer</a> (NFL) </td> <td><a href="/wiki/Ganglion_cell" title="Ganglion cell">Ganglion cell</a> <a href="/wiki/Axons" class="mw-redirect" title="Axons">axons</a> travelling towards the <a href="/wiki/Optic_nerve" title="Optic nerve">optic nerve</a> </td></tr> <tr> <td>4</td> <td><a href="/wiki/Ganglion_cell_layer" title="Ganglion cell layer">Ganglion cell layer</a> (GCL)</td> <td><a href="/wiki/Ganglion_cell" title="Ganglion cell">Ganglion cell</a> bodies (and some displaced <a href="/wiki/Amacrine_cells" class="mw-redirect" title="Amacrine cells">amacrine cells</a>) </td> <td>Hypo-reflective </td> <td> </td> <td><sup id="cite_ref-:0_25-4" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>5</td> <td><a href="/wiki/Inner_plexiform_layer" title="Inner plexiform layer">Inner plexiform layer</a> (IPL)</td> <td>Synapses between <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar</a>, <a href="/wiki/Amacrine" class="mw-redirect" title="Amacrine">amacrine</a> and <a href="/wiki/Ganglion_cells" class="mw-redirect" title="Ganglion cells">ganglion cells</a> </td> <td>Hyper-reflective </td> <td> </td> <td><sup id="cite_ref-:0_25-5" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>6</td> <td><a href="/wiki/Inner_nuclear_layer" title="Inner nuclear layer">Inner nuclear layer</a> (INL)</td> <td>a) <a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">Horizontal</a>, <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar</a> and <a href="/wiki/Amacrine" class="mw-redirect" title="Amacrine">amacrine</a> cell bodies <p>b) <a href="/wiki/M%C3%BCller_cell" class="mw-redirect" title="Müller cell">Müller cell</a> nuclei </p> </td> <td>Hypo-reflective </td> <td> </td> <td><sup id="cite_ref-:0_25-6" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>7</td> <td><a href="/wiki/Outer_plexiform_layer" title="Outer plexiform layer">Outer plexiform layer</a> (OPL)</td> <td><a href="/wiki/Synapses" class="mw-redirect" title="Synapses">Synapses</a> between <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor</a>, <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">bipolar</a> and <a href="/wiki/Horizontal_cells" class="mw-redirect" title="Horizontal cells">horizontal cells</a> </td> <td>Hyper-reflective </td> <td> </td> <td><sup id="cite_ref-:0_25-7" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td rowspan="2">8</td> <td>(Inner half) <a href="/w/index.php?title=Henle%27s_fiber_layer&action=edit&redlink=1" class="new" title="Henle's fiber layer (page does not exist)">Henle's nerve fiber layer</a> (HL) </td> <td><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor</a> axons <p>(obliquely orientated fibres; not present in mid-peripheral or peripheral retina) </p> </td> <td rowspan="2">Hypo-reflective </td> <td rowspan="2">No </td> <td rowspan="2"><sup id="cite_ref-:0_25-8" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>(Outer half) <a href="/wiki/Outer_nuclear_layer" title="Outer nuclear layer">Outer nuclear layer</a> (ONL) </td> <td>The <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor</a> cell bodies </td></tr> <tr> <td>9</td> <td><a href="/wiki/External_limiting_membrane" title="External limiting membrane">External limiting membrane</a> (ELM) </td> <td>Made of <a href="/w/index.php?title=Zonulae_adherens&action=edit&redlink=1" class="new" title="Zonulae adherens (page does not exist)">zonulae adherens</a> between <a href="/wiki/M%C3%BCller_cells" class="mw-redirect" title="Müller cells">Müller cells</a> and <a href="/w/index.php?title=Photoreceptor_inner_segments&action=edit&redlink=1" class="new" title="Photoreceptor inner segments (page does not exist)">photoreceptor inner segments</a> </td> <td>Hyper-reflective </td> <td> </td> <td><sup id="cite_ref-:0_25-9" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>10</td> <td><a href="/w/index.php?title=Myoid_zone&action=edit&redlink=1" class="new" title="Myoid zone (page does not exist)">Myoid zone (MZ)</a></td> <td>The innermost portion of the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor inner segment (IS)</a> containing: <ul><li>Smooth and rough <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a></li> <li><a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">Ribosomes</a></li> <li><a href="/wiki/Golgi_bodies" class="mw-redirect" title="Golgi bodies">Golgi bodies</a></li> <li><a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">Microtubules</a></li> <li><i>(rarely: mitochondria)</i></li></ul> </td> <td>Hypo-reflective </td> <td>No </td> <td><sup id="cite_ref-Hildebrand_2011_39–65_27-0" class="reference"><a href="#cite_note-Hildebrand_2011_39–65-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td rowspan="2">11</td> <td><a href="/w/index.php?title=Ellipsoid_zone&action=edit&redlink=1" class="new" title="Ellipsoid zone (page does not exist)">Ellipsoid zone (EZ)</a></td> <td>The outermost portion of the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor inner segment (IS)</a> packed with <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> </td> <td rowspan="2"><b>Very Hyper-reflective</b> </td> <td rowspan="2">No </td> <td rowspan="2"><sup id="cite_ref-sciencedirect.com_24-1" class="reference"><a href="#cite_note-sciencedirect.com-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hildebrand_2011_39–65_27-1" class="reference"><a href="#cite_note-Hildebrand_2011_39–65-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_25-10" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><a href="/w/index.php?title=IS/OS_junction&action=edit&redlink=1" class="new" title="IS/OS junction (page does not exist)">IS/OS junction</a> or <a href="/w/index.php?title=Photoreceptor_integrity_line&action=edit&redlink=1" class="new" title="Photoreceptor integrity line (page does not exist)">Photoreceptor integrity line (PIL)</a> </td> <td>The photoreceptor <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">connecting cilia</a> which bridge the inner and outer segments of the photoreceptor cells. </td></tr> <tr> <td>12</td> <td><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor outer segments (OS)</a></td> <td>The <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor outer segments (OS)</a> which contain disks filled with <a href="/wiki/Opsin" title="Opsin">opsin</a>, the molecule that absorbs photons. </td> <td>Hypo-reflective </td> <td> </td> <td rowspan="2"><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_25-11" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>13</td> <td><a href="/w/index.php?title=Interdigitation_zone&action=edit&redlink=1" class="new" title="Interdigitation zone (page does not exist)">Interdigitation zone (IZ)</a></td> <td>Apices of the <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">RPE</a> cells which encase part of the cone OSs. <p><i>Poorly distinguishable from RPE.</i> <i>Previously: "cone outer segment tips line"</i> <i>(COST)</i> </p> </td> <td>Hyper-reflective </td> <td>No </td></tr> <tr> <td rowspan="3">14 </td> <td rowspan="3"><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">RPE</a>/<a href="/w/index.php?title=Bruch%27s&action=edit&redlink=1" class="new" title="Bruch's (page does not exist)">Bruch's</a> complex </td> <td><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">RPE</a> phagosome zone </td> <td><b>Very Hyper-reflective</b> </td> <td rowspan="3">No </td> <td rowspan="3"><sup id="cite_ref-sciencedirect.com_24-2" class="reference"><a href="#cite_note-sciencedirect.com-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_25-12" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">RPE</a> melanosome zone </td> <td>Hypo-reflective </td></tr> <tr> <td><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">RPE</a> mitochondria zone + Junction between the RPE & <a href="/wiki/Bruch%27s_membrane" title="Bruch's membrane">Bruch's membrane</a> </td> <td><b>Very Hyper-reflective</b> </td></tr> <tr> <td>15 </td> <td><a href="/wiki/Choriocapillaris" class="mw-redirect" title="Choriocapillaris">Choriocapillaris</a> </td> <td colspan="2">Thin layer of moderate reflectivity in inner choroid </td> <td>No </td> <td><sup id="cite_ref-:0_25-13" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>16 </td> <td><a href="/wiki/Sattler%27s_layer" title="Sattler's layer">Sattler's layer</a> </td> <td colspan="2">Thick layer of round or ovalshaped hyperreflective profiles, with hyporeflective cores in mid-choroid </td> <td> </td> <td><sup id="cite_ref-:0_25-14" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>17 </td> <td><a href="/w/index.php?title=Haller%27s_layer&action=edit&redlink=1" class="new" title="Haller's layer (page does not exist)">Haller's layer</a> </td> <td colspan="2">Thick layer of oval-shaped hyperreflective profiles, with hyporeflective cores in outer choroid </td> <td> </td> <td><sup id="cite_ref-:0_25-15" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr> <tr> <td>18 </td> <td><a href="/w/index.php?title=Choroidal-scleral_juncture&action=edit&redlink=1" class="new" title="Choroidal-scleral juncture (page does not exist)">Choroidal-scleral juncture</a> </td> <td colspan="2">Zone at the outer choroid with a marked change in texture, in which large circular or ovoid profiles abut a <p>homogenous region of variable reflectivity </p> </td> <td> </td> <td><sup id="cite_ref-:0_25-16" class="reference"><a href="#cite_note-:0-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Development">Development</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=5" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Retinal development begins with the establishment of the eye fields mediated by the <a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">SHH</a> and <a href="/wiki/SIX3" title="SIX3">SIX3</a> proteins, with subsequent development of the optic vesicles regulated by the <a href="/wiki/PAX6" title="PAX6">PAX6</a> and <a href="/wiki/LHX2" title="LHX2">LHX2</a> proteins.<sup id="cite_ref-Heavner_33-0" class="reference"><a href="#cite_note-Heavner-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The role of Pax6 in eye development was elegantly demonstrated by Walter Gehring and colleagues, who showed that ectopic expression of Pax6 can lead to eye formation on <a href="/wiki/Drosophila" title="Drosophila">Drosophila</a> antennae, wings, and legs.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> The optic vesicle gives rise to three structures: the neural retina, the retinal pigmented epithelium, and the optic stalk. The neural retina contains the retinal progenitor cells (RPCs) that give rise to the seven cell types of the retina. Differentiation begins with the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">retinal ganglion cells</a> and concludes with production of the Muller glia.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Although each cell type differentiates from the RPCs in a sequential order, there is considerable overlap in the timing of when individual cell types differentiate.<sup id="cite_ref-Heavner_33-1" class="reference"><a href="#cite_note-Heavner-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The cues that determine a RPC daughter cell fate are coded by multiple transcription factor families including the <a href="/wiki/Basic_helix-loop-helix" class="mw-redirect" title="Basic helix-loop-helix">bHLH</a> and <a href="/wiki/Homeobox" title="Homeobox">homeodomain</a> factors.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_37-0" class="reference"><a href="#cite_note-:1-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p><p>In addition to guiding cell fate determination, cues exist in the retina to determine the dorsal-ventral (D-V) and nasal-temporal (N-T) axes. The D-V axis is established by a ventral to dorsal gradient of <a href="/wiki/VAX1" title="VAX1">VAX2</a>, whereas the N-T axis is coordinated by expression of the forkhead transcription factors <a href="/wiki/FOXD1" class="mw-redirect" title="FOXD1">FOXD1</a> and <a href="/wiki/FOXG1" title="FOXG1">FOXG1</a>. Additional gradients are formed within the retina.<sup id="cite_ref-:1_37-1" class="reference"><a href="#cite_note-:1-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> This spatial distribution may aid in proper targeting of RGC axons that function to establish the retinotopic map.<sup id="cite_ref-Heavner_33-2" class="reference"><a href="#cite_note-Heavner-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Blood_supply">Blood supply</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=6" title="Edit section: Blood supply"><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 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You can help by <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Retina&action=edit&section=">adding to it</a>. <span class="date-container"><i>(<span class="date">October 2016</span>)</i></span></div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444" /><table class="box-Self-published plainlinks metadata ambox ambox-content ambox-self-published" 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/b/b4/Ambox_important.svg/40px-Ambox_important.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/60px-Ambox_important.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/b/b4/Ambox_important.svg/80px-Ambox_important.svg.png 2x" data-file-width="40" data-file-height="40" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>may contain excessive or inappropriate references to <a href="/wiki/Wikipedia:Verifiability#Self-published_sources" title="Wikipedia:Verifiability">self-published sources</a></b>.<span class="hide-when-compact"> Please help <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Retina&action=edit">improve it</a> by removing references to unreliable <a href="/wiki/Wikipedia:Reliable_sources" title="Wikipedia:Reliable sources">sources</a> where they are used inappropriately.</span> <span class="date-container"><i>(<span class="date">October 2016</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> </div> </div><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Fundus_photograph_of_normal_right_eye.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Fundus_photograph_of_normal_right_eye.jpg/220px-Fundus_photograph_of_normal_right_eye.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Fundus_photograph_of_normal_right_eye.jpg/330px-Fundus_photograph_of_normal_right_eye.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/Fundus_photograph_of_normal_right_eye.jpg/440px-Fundus_photograph_of_normal_right_eye.jpg 2x" data-file-width="1411" data-file-height="1411" /></a><figcaption><a href="/wiki/Fundus_photograph" class="mw-redirect" title="Fundus photograph">Fundus photograph</a> showing the blood vessels in a normal human retina. Veins are darker and slightly wider than corresponding arteries. The <a href="/wiki/Optic_disc" title="Optic disc">optic disc</a> is at right, and the <a href="/wiki/Macula_lutea" class="mw-redirect" title="Macula lutea">macula lutea</a> is near the centre.</figcaption></figure> <p>The retina is stratified into distinct layers, each containing specific cell types or cellular compartments<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> that have metabolisms with different nutritional requirements.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> To satisfy these requirements, the <a href="/wiki/Ophthalmic_artery" title="Ophthalmic artery">ophthalmic artery</a> bifurcates and supplies the retina via two distinct vascular networks: the choroidal network, which supplies the choroid and the outer retina, and the retinal network, which supplies the retina's inner layer.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p><p>Although the inverted retina of vertebrates appears counter-intuitive, it is necessary for the proper functioning of the retina. The photoreceptor layer must be embedded in the retinal pigment epithelium (RPE), which performs at least seven vital functions,<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> one of the most obvious being to supply oxygen and other necessary nutrients needed for the photoreceptors to function. </p> <div class="mw-heading mw-heading4"><h4 id="Energy_requirements">Energy requirements</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=7" title="Edit section: Energy requirements"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The energy requirements of the retina are even greater than that of the brain.<sup id="cite_ref-Viegas_Neuhauss_4-1" class="reference"><a href="#cite_note-Viegas_Neuhauss-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This is due to the additional energy needed to continuously renew the photoreceptor outer segments, of which 10% are shed daily.<sup id="cite_ref-Viegas_Neuhauss_4-2" class="reference"><a href="#cite_note-Viegas_Neuhauss-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Energy demands are greatest during dark adaptation when its sensitivity is most enhanced.<sup id="cite_ref-Kaynezhad_Tachtsidis_Sivaprasad_Jeffery_42-0" class="reference"><a href="#cite_note-Kaynezhad_Tachtsidis_Sivaprasad_Jeffery-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> The choroid supplies about 75% of these nutrients to the retina and the retinal vasculature only 25%.<sup id="cite_ref-Kolb_5-1" class="reference"><a href="#cite_note-Kolb-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>When light strikes 11-cis-retinal (in the disks in the rods and cones), 11-cis-retinal changes to all-trans-retinal which then triggers changes in the opsins. Now, the outer segments do not regenerate the retinal back into the cis- form once it has been changed by light. Instead the retinal is pumped out to the surrounding RPE where it is regenerated and transported back into the outer segments of the photoreceptors. This recycling function of the RPE protects the photoreceptors against photo-oxidative damage<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> and allows the photoreceptor cells to have decades-long useful lives. </p> <div class="mw-heading mw-heading4"><h4 id="In_birds">In birds</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=8" title="Edit section: In birds"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The bird retina is devoid of blood vessels, perhaps to give unobscured passage of light for forming images, thus giving better resolution. It is, therefore, a considered view that the bird retina depends for nutrition and oxygen supply on a specialized organ, called the "pecten" or <a href="/wiki/Pecten_oculi" title="Pecten oculi">pecten oculi</a>, located on the blind spot or optic disk. This organ is extremely rich in blood vessels and is thought to supply nutrition and oxygen to the bird retina by diffusion through the vitreous body. The pecten is highly rich in alkaline phosphatase activity and polarized cells in its bridge portion – both befitting its secretory role.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Pecten cells are packed with dark melanin granules, which have been theorized to keep this organ warm with the absorption of stray light falling on the pecten. This is considered to enhance metabolic rate of the pecten, thereby exporting more nutritive molecules to meet the stringent energy requirements of the retina during long periods of exposure to light.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Biometric_identification_and_diagnosis_of_disease">Biometric identification and diagnosis of disease</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=9" title="Edit section: Biometric identification and diagnosis of disease"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Retinal_scan" title="Retinal scan">Retinal scan</a> and <a href="/wiki/Biometrics" title="Biometrics">Biometrics</a></div> <p>The bifurcations and other physical characteristics of the inner retinal vascular network are known to vary among individuals,<sup id="cite_ref-Sherman81_47-0" class="reference"><a href="#cite_note-Sherman81-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> and these individual variances have been used for <a href="/wiki/Retinal_scan" title="Retinal scan">biometric identification</a> and for early detection of the onset of disease. The mapping of vascular bifurcations is one of the basic steps in biometric identification.<sup id="cite_ref-Azzopardi2011_48-0" class="reference"><a href="#cite_note-Azzopardi2011-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Results of such analyses of retinal blood vessel structure can be evaluated against the ground truth data<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> of vascular bifurcations of retinal fundus images that are obtained from the DRIVE dataset.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> In addition, the classes of vessels of the DRIVE dataset have also been identified,<sup id="cite_ref-Qureshi2013_51-0" class="reference"><a href="#cite_note-Qureshi2013-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> and an automated method for accurate extraction of these bifurcations is also available.<sup id="cite_ref-Qureshi2014_52-0" class="reference"><a href="#cite_note-Qureshi2014-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Changes in retinal blood circulation are seen with aging<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> and exposure to air pollution,<sup id="cite_ref-Louwies_54-0" class="reference"><a href="#cite_note-Louwies-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> and may indicate cardiovascular diseases such as hypertension and atherosclerosis.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Chapman2002_56-0" class="reference"><a href="#cite_note-Chapman2002-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Patton06_57-0" class="reference"><a href="#cite_note-Patton06-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Determining the equivalent width of arterioles and venules near the optic disc is also a widely used technique to identify cardiovascular risks.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Function">Function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=10" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Adaptation_(eye)" title="Adaptation (eye)">Adaptation (eye)</a> and <a href="/wiki/Visual_acuity" title="Visual acuity">Visual acuity</a></div> <p>The retina translates an optical image into neural impulses starting with the patterned excitation of the colour-sensitive pigments of its rods and cones, the retina's <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a>. The excitation is processed by the neural system and various parts of the brain working in parallel to form a representation of the external environment in the brain.<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. (July 2024)">citation needed</span></a></i>]</sup> </p><p>The cones respond to bright light and mediate high-resolution colour vision during daylight illumination (also called <a href="/wiki/Photopic_vision" title="Photopic vision">photopic vision</a>). The rod responses are saturated at daylight levels and do not contribute to pattern vision. However, rods do respond to dim light and mediate lower-resolution, monochromatic vision under very low levels of illumination (called <a href="/wiki/Scotopic_vision" title="Scotopic vision">scotopic vision</a>). The illumination in most office settings falls between these two levels and is called <a href="/wiki/Mesopic_vision" title="Mesopic vision">mesopic vision</a>. At mesopic light levels, both the rods and cones are actively contributing pattern information. What contribution the rod information makes to pattern vision under these circumstances is unclear. </p><p>The response of cones to various wavelengths of light is called their spectral sensitivity. In normal human vision, the spectral sensitivity of a cone falls into one of three subtypes, often called blue, green, and red, but more accurately known as short, medium, and long wavelength-sensitive cone subtypes. It is a lack of one or more of the cone subtypes that causes individuals to have deficiencies in colour vision or various kinds of <a href="/wiki/Colour_blindness" class="mw-redirect" title="Colour blindness">colour blindness</a>. These individuals are not blind to objects of a particular colour, but are unable to distinguish between colours that can be distinguished by people with normal vision. Humans have this <a href="/wiki/Trichromatic_vision" class="mw-redirect" title="Trichromatic vision">trichromatic vision</a>, while most other mammals lack cones with red sensitive pigment and therefore have poorer dichromatic colour vision. However, some animals have four spectral subtypes, e.g. the trout adds an ultraviolet subgroup to short, medium, and long subtypes that are similar to humans. Some fish are sensitive to the polarization of light as well. </p><p>In the photoreceptors, exposure to light hyperpolarizes the membrane in a series of graded shifts. The outer cell segment contains a <a href="/wiki/Photopigment" title="Photopigment">photopigment</a>. Inside the cell the normal levels of <a href="/wiki/Cyclic_guanosine_monophosphate" title="Cyclic guanosine monophosphate">cyclic guanosine monophosphate</a> (cGMP) keep the Na+ channel open, and thus in the resting state the cell is depolarised. The <a href="/wiki/Photon" title="Photon">photon</a> causes the <a href="/wiki/Retinal" title="Retinal">retinal</a> bound to the receptor protein to <a href="/wiki/Isomerism" class="mw-redirect" title="Isomerism">isomerise</a> to <a href="/wiki/Retinal" title="Retinal">trans-retinal</a>. This causes the receptor to activate multiple <a href="/wiki/G-protein" class="mw-redirect" title="G-protein">G-proteins</a>. This in turn causes the Ga-subunit of the protein to activate a phosphodiesterase (PDE6), which degrades cGMP, resulting in the closing of Na+ <a href="/wiki/Cyclic_nucleotide-gated_ion_channel" class="mw-redirect" title="Cyclic nucleotide-gated ion channel">cyclic nucleotide-gated ion channels</a> (CNGs). Thus the cell is hyperpolarised. The amount of neurotransmitter released is reduced in bright light and increases as light levels fall. The actual photopigment is bleached away in bright light and only replaced as a chemical process, so in a transition from bright light to darkness the eye can take up to thirty minutes to reach full sensitivity. </p><p>When thus excited by light, the photoceptor sends a proportional response <a href="/wiki/Synapse" title="Synapse">synaptically</a> to <a href="/wiki/Bipolar_cell" class="mw-redirect" title="Bipolar cell">bipolar cells</a> which in turn signal the <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">retinal ganglion cells</a>. The photoreceptors are also cross-linked by <a href="/wiki/Horizontal_cell" class="mw-redirect" title="Horizontal cell">horizontal cells</a> and <a href="/wiki/Amacrine_cell" title="Amacrine cell">amacrine cells</a>, which modify the synaptic signal before it reaches the ganglion cells, the neural signals being intermixed and combined. Of the retina's nerve cells, only the retinal ganglion cells and few amacrine cells create <a href="/wiki/Action_potential" title="Action potential">action potentials</a>. </p><p>In the retinal ganglion cells there are two types of response, depending on the <a href="/wiki/Receptive_field" title="Receptive field">receptive field</a> of the cell. The receptive fields of retinal ganglion cells comprise a central, approximately circular area, where light has one effect on the firing of the cell, and an annular surround, where light has the opposite effect. In ON cells, an increment in light intensity in the centre of the receptive field causes the firing rate to increase. In OFF cells, it makes it decrease. In a linear model, this response profile is well described by a <a href="/wiki/Difference_of_Gaussians" title="Difference of Gaussians">difference of Gaussians</a> and is the basis for <a href="/wiki/Edge_detection" title="Edge detection">edge detection</a> algorithms. Beyond this simple difference, ganglion cells are also differentiated by chromatic sensitivity and the type of spatial summation. Cells showing linear spatial summation are termed X cells (also called parvocellular, P, or midget ganglion cells), and those showing non-linear summation are Y cells (also called magnocellular, M, or parasol retinal ganglion cells), although the correspondence between X and Y cells (in the cat retina) and P and M cells (in the primate retina) is not as simple as it once seemed. </p><p>In the transfer of visual signals to the brain, the <a href="/wiki/Visual_pathway" class="mw-redirect" title="Visual pathway">visual pathway</a>, the retina is vertically divided in two, a temporal (nearer to the temple) half and a nasal (nearer to the nose) half. The axons from the nasal half cross the brain at the <a href="/wiki/Optic_chiasma" class="mw-redirect" title="Optic chiasma">optic chiasma</a> to join with axons from the temporal half of the other eye before passing into the <a href="/wiki/Lateral_geniculate_body" class="mw-redirect" title="Lateral geniculate body">lateral geniculate body</a>. </p><p>Although there are more than 130 million retinal receptors, there are only approximately 1.2 million fibres (axons) in the optic nerve. So, a large amount of pre-processing is performed within the retina. The <a href="/wiki/Fovea_centralis" title="Fovea centralis">fovea</a> produces the most accurate information. Despite occupying about 0.01% of the visual field (less than 2° of <a href="/wiki/Visual_angle" title="Visual angle">visual angle</a>), about 10% of axons in the optic nerve are devoted to the fovea. The resolution limit of the fovea has been determined to be around 10,000 points. The information capacity is estimated at 500,000 bits per second (for more information on bits, see <a href="/wiki/Information_theory" title="Information theory">information theory</a>) without colour or around 600,000 bits per second including colour.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Spatial_encoding">Spatial encoding</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=11" title="Edit section: Spatial encoding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Receptive_field" title="Receptive field">Receptive field, for figures and more information on centre–surround structures</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Receptive_field.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Receptive_field.png/300px-Receptive_field.png" decoding="async" width="300" height="549" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/1/16/Receptive_field.png 1.5x" data-file-width="403" data-file-height="737" /></a><figcaption>On-centres and off-centres of the retina</figcaption></figure> <p>When the retina sends neural impulses representing an image to the brain, it spatially encodes (compresses) those impulses to fit the limited capacity of the optic nerve. Compression is necessary because there are 100 times more <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cells</a> than <a href="/wiki/Ganglion_cell" title="Ganglion cell">ganglion cells</a>. This is done by "<a href="/wiki/Decorrelation" title="Decorrelation">decorrelation</a>", which is carried out by the "centre–surround structures", which are implemented by the bipolar and ganglion cells. </p><p>There are two types of centre–surround structures in the retina – on-centres and off-centres. On-centres have a positively weighted centre and a negatively weighted surround. Off-centres are just the opposite. Positive weighting is more commonly known as <a href="/wiki/Chemical_synapse#Receptor_binding" title="Chemical synapse">excitatory</a>, and negative weighting as <a href="/wiki/Chemical_synapse#Receptor_binding" title="Chemical synapse">inhibitory</a>. </p><p>These centre–surround structures are not physical apparent, in the sense that one cannot see them by staining samples of tissue and examining the retina's anatomy. The centre–surround structures are logical (i.e., mathematically abstract) in the sense that they depend on the connection strengths between bipolar and ganglion cells. It is believed that the connection strength between cells is caused by the number and types of <a href="/wiki/Ion_channel" title="Ion channel">ion channels</a> embedded in the <a href="/wiki/Synapse" title="Synapse">synapses</a> between the bipolar and ganglion cells. </p><p>The centre–surround structures are mathematically equivalent to the <a href="/wiki/Edge_detection" title="Edge detection">edge detection</a> algorithms used by computer programmers to extract or enhance the edges in a digital photograph. Thus, the retina performs operations on the image-representing impulses to enhance the edges of objects within its visual field. For example, in a picture of a dog, a cat and a car, it is the edges of these objects that contain the most information. In order for higher functions in the brain (or in a computer for that matter) to extract and classify objects such as a dog and a cat, the retina is the first step to separating out the various objects within the scene. </p><p>As an example, the following <a href="/wiki/Matrix_(mathematics)" title="Matrix (mathematics)">matrix</a> is at the heart of a computer <a href="/wiki/Algorithm" title="Algorithm">algorithm</a> that implements edge detection. This matrix is the computer equivalent to the centre–surround structure. In this example, each box (element) within this matrix would be connected to one photoreceptor. The photoreceptor in the centre is the current receptor being processed. The centre photoreceptor is multiplied by the +1 weight factor. The surrounding photoreceptors are the "nearest neighbors" to the centre and are multiplied by the −1/8 value. The sum of all nine of these elements is finally calculated. This summation is repeated for every photoreceptor in the image by shifting left to the end of a row and then down to the next line. </p> <table class="wikitable"> <tbody><tr> <td style="background:lightyellow;">-1/8</td> <td style="background:lightyellow;">-1/8</td> <td style="background:lightyellow;">-1/8 </td></tr> <tr> <td style="background:lightyellow;">-1/8</td> <td style="background:lightblue;">+1</td> <td style="background:lightyellow;">-1/8 </td></tr> <tr> <td style="background:lightyellow;">-1/8</td> <td style="background:lightyellow;">-1/8</td> <td style="background:lightyellow;">-1/8 </td></tr></tbody></table> <p>The total sum of this matrix is zero, if all the inputs from the nine photoreceptors are of the same value. The zero result indicates the image was uniform (non-changing) within this small patch. Negative or positive sums mean the image was varying (changing) within this small patch of nine photoreceptors. </p><p>The above matrix is only an approximation to what really happens inside the retina. The differences are: </p> <ul><li>The above example is called "balanced". The term balanced means that the sum of the negative weights is equal to the sum of the positive weights so that they cancel out perfectly. Retinal ganglion cells are almost never perfectly balanced.</li> <li>The table is square while the centre–surround structures in the retina are circular.</li> <li>Neurons operate on <a href="/wiki/Spike_train" class="mw-redirect" title="Spike train">spike trains</a> traveling down nerve cell <a href="/wiki/Axons" class="mw-redirect" title="Axons">axons</a>. Computers operate on a single <a href="/wiki/Floating-point_arithmetic" title="Floating-point arithmetic">floating-point number</a> that is essentially constant from each input <a href="/wiki/Pixel" title="Pixel">pixel</a>. (The computer pixel is basically the equivalent of a biological photoreceptor.)</li> <li>The retina performs all these calculations in parallel while the computer operates on each pixel one at a time. The retina performs no repeated summations and shifting as would a computer.</li> <li>Finally, the <a href="/wiki/Horizontal_cell" class="mw-redirect" title="Horizontal cell">horizontal</a> and <a href="/wiki/Amacrine_cell" title="Amacrine cell">amacrine cells</a> play a significant role in this process, but that is not represented here.</li></ul> <p>Here is an example of an input image and how edge detection would modify it. </p><p><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Edge-detection-2.jpg" class="mw-file-description"><img alt="input image" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/62/Edge-detection-2.jpg/550px-Edge-detection-2.jpg" decoding="async" width="550" height="105" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/6/62/Edge-detection-2.jpg 1.5x" data-file-width="672" data-file-height="128" /></a></span> </p><p>Once the image is spatially encoded by the centre–surround structures, the signal is sent out along the optic nerve (via the axons of the ganglion cells) through the <a href="/wiki/Optic_chiasm" title="Optic chiasm">optic chiasm</a> to the LGN (<a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">lateral geniculate nucleus</a>). The exact function of the LGN is unknown at this time. The output of the LGN is then sent to the back of the brain. Specifically, the output of the LGN "radiates" out to the V1 <a href="/wiki/Primary_visual_cortex" class="mw-redirect" title="Primary visual cortex">primary visual cortex</a>. </p><p>Simplified signal flow: Photoreceptors → Bipolar → Ganglion → Chiasm → LGN → V1 cortex </p><p><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:ERP_-_optic_cabling.jpg" class="mw-file-description"><img alt="ERP optic cabling" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/ERP_-_optic_cabling.jpg/500px-ERP_-_optic_cabling.jpg" decoding="async" width="500" height="375" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/ERP_-_optic_cabling.jpg/750px-ERP_-_optic_cabling.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/d/d0/ERP_-_optic_cabling.jpg 2x" data-file-width="800" data-file-height="600" /></a></span> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_significance">Clinical significance<span class="anchor" id="Diseases_and_disorders"></span></h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=12" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are many inherited and acquired diseases or disorders that may affect the retina. Some of them include: </p> <ul><li><a href="/wiki/Retinitis_pigmentosa" title="Retinitis pigmentosa">Retinitis pigmentosa</a> is a group of genetic diseases that affect the retina and cause the loss of night vision and peripheral vision.</li> <li><a href="/wiki/Macular_degeneration" title="Macular degeneration">Macular degeneration</a> describes a group of diseases characterized by loss of central vision because of death or impairment of the cells in the <a href="/wiki/Macula" title="Macula">macula</a>.</li> <li><a href="/wiki/Cone-rod_dystrophy" class="mw-redirect" title="Cone-rod dystrophy">Cone-rod dystrophy</a> (CORD) describes a number of diseases where vision loss is caused by deterioration of the <a href="/wiki/Cone_cell" title="Cone cell">cones</a> and/or <a href="/wiki/Rod_cell" title="Rod cell">rods</a> in the retina.</li> <li>In <a href="/wiki/Retinal_separation" class="mw-redirect" title="Retinal separation">retinal separation</a>, the retina detaches from the back of the eyeball. <a href="/wiki/Ignipuncture" title="Ignipuncture">Ignipuncture</a> is an outdated treatment method. The term <a href="/wiki/Retinal_detachment" title="Retinal detachment">retinal detachment</a> is used to describe a separation of the neurosensory retina from the <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a>.<sup id="cite_ref-Retina_60-0" class="reference"><a href="#cite_note-Retina-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> There are several modern treatment methods for fixing a retinal detachment: <a href="/wiki/Pneumatic_retinopexy" class="mw-redirect" title="Pneumatic retinopexy">pneumatic retinopexy</a>, <a href="/wiki/Scleral_buckle" title="Scleral buckle">scleral buckle</a>, <a href="/wiki/Cryotherapy" title="Cryotherapy">cryotherapy</a>, <a href="/wiki/Laser_photocoagulation" class="mw-redirect" title="Laser photocoagulation">laser photocoagulation</a> and pars plana <a href="/wiki/Vitrectomy" title="Vitrectomy">vitrectomy</a>.</li> <li>Both <a href="/wiki/Hypertension" title="Hypertension">hypertension</a> and <a href="/wiki/Diabetes_mellitus" class="mw-redirect" title="Diabetes mellitus">diabetes mellitus</a> can cause damage to the tiny blood vessels that supply the retina, leading to <a href="/wiki/Hypertensive_retinopathy" title="Hypertensive retinopathy">hypertensive retinopathy</a> and <a href="/wiki/Diabetic_retinopathy" title="Diabetic retinopathy">diabetic retinopathy</a>.</li> <li><a href="/wiki/Retinoblastoma" title="Retinoblastoma">Retinoblastoma</a> is a <a href="/wiki/Cancer" title="Cancer">cancer</a> of the retina.</li> <li>Retinal diseases in <a href="/wiki/Dog" title="Dog">dogs</a> include <a href="/wiki/Retinal_dysplasia" title="Retinal dysplasia">retinal dysplasia</a>, <a href="/wiki/Progressive_retinal_atrophy" title="Progressive retinal atrophy">progressive retinal atrophy</a>, and <a href="/wiki/Sudden_acquired_retinal_degeneration" class="mw-redirect" title="Sudden acquired retinal degeneration">sudden acquired retinal degeneration</a>.</li> <li><i><a href="/wiki/Lipaemia_retinalis" title="Lipaemia retinalis">Lipaemia retinalis</a></i> is a white appearance of the retina, and can occur by lipid deposition in <a href="/wiki/Lipoprotein_lipase_deficiency" title="Lipoprotein lipase deficiency">lipoprotein lipase deficiency</a>.</li> <li><i><a href="/wiki/Retinal_Detachment" class="mw-redirect" title="Retinal Detachment">Retinal Detachment</a></i>. The neural retina occasionally detaches from the pigment epithelium. In some instances, the cause of such detachment is injury to the eyeball that allows fluid or blood to collect between the neural retina and the pigment epithelium. Detachment is occasionally caused by contracture of fine collagenous fibrils in the vitreous humor, which pull areas of the retina toward the interior of the globe.<sup id="cite_ref-ReferenceA_23-1" class="reference"><a href="#cite_note-ReferenceA-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup></li> <li><i><a href="/wiki/Night_blindness" class="mw-redirect" title="Night blindness">Night blindness</a></i>: Night blindness occurs in any person with severe vitamin A deficiency. The reason for this is that without vitamin A, the amounts of retinal and rhodopsin that can be formed are severely depressed. This condition is called night blindness because the amount of light available at night is too little to permit adequate vision in vitamin A–deficient persons.<sup id="cite_ref-Guyton_and_Hall_Physiology_18-1" class="reference"><a href="#cite_note-Guyton_and_Hall_Physiology-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup></li></ul> <p>In addition, the retina has been described as a "window" into the brain and body, given that abnormalities detected through an examination of the retina can discover both neurological and systemic diseases.<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Diagnosis">Diagnosis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=13" title="Edit section: Diagnosis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A number of different instruments are available for the diagnosis of diseases and disorders affecting the retina. <a href="/wiki/Ophthalmoscopy" title="Ophthalmoscopy">Ophthalmoscopy</a> and <a href="/wiki/Fundus_photography" title="Fundus photography">fundus photography</a> have long been used to examine the retina. Recently, <a href="/wiki/Adaptive_optics" title="Adaptive optics">adaptive optics</a> has been used to image individual rods and cones in the living human retina, and a company based in Scotland has engineered technology that allows physicians to observe the complete retina without any discomfort to patients.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>The <a href="/wiki/Electroretinography" title="Electroretinography">electroretinogram</a> is used to <a href="/wiki/Non-invasive_(medical)" class="mw-redirect" title="Non-invasive (medical)">non-invasively</a> measure the retina's electrical activity, which is affected by certain diseases. A relatively new technology, now becoming widely available, is <a href="/wiki/Optical_coherence_tomography" title="Optical coherence tomography">optical coherence tomography</a> (OCT). This non-invasive technique allows one to obtain a <a href="/wiki/Dimension" title="Dimension">3D</a> volumetric or high resolution cross-sectional <a href="/wiki/Tomogram" class="mw-redirect" title="Tomogram">tomogram</a> of the fine structures of the retina, with <a href="/wiki/Histologic" class="mw-redirect" title="Histologic">histologic</a> quality. <a href="/wiki/Retinal_vessel_analysis" title="Retinal vessel analysis">Retinal vessel analysis</a> is a non-invasive method to examine the small arteries and veins in the retina which allows to draw conclusions about the morphology and the function of small vessels elsewhere in the human body. It has been established as a predictor of <a href="/wiki/Cardiovascular_disease" title="Cardiovascular disease">cardiovascular disease</a><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> and seems to have, according to a study published in 2019, potential in the early detection of Alzheimer's disease.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Treatment">Treatment</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=14" title="Edit section: Treatment"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Treatment depends upon the nature of the disease or disorder. </p> <div class="mw-heading mw-heading4"><h4 id="Common_treatment_modalities">Common treatment modalities</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=15" title="Edit section: Common treatment modalities"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The following are commonly modalities of management for retinal disease: </p> <ul><li><a href="/wiki/Intravitreal" class="mw-redirect" title="Intravitreal">Intravitreal</a> medication, such as <a href="/wiki/Anti%E2%80%93vascular_endothelial_growth_factor_therapy" class="mw-redirect" title="Anti–vascular endothelial growth factor therapy">anti-VEGF</a> or <a href="/wiki/Corticosteroid" title="Corticosteroid">corticosteroid</a> agents</li> <li><a href="/wiki/Vitreoretinal_surgery" class="mw-redirect" title="Vitreoretinal surgery">Vitreoretinal surgery</a></li> <li>Use of nutritional supplements</li> <li>Modification of systemic risk factors for retinal disease</li></ul> <div class="mw-heading mw-heading4"><h4 id="Uncommon_treatment_modalities">Uncommon treatment modalities</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=16" title="Edit section: Uncommon treatment modalities"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><b>Retinal gene therapy</b> </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Adeno_associated_virus_and_gene_therapy_of_the_human_retina" class="mw-redirect" title="Adeno associated virus and gene therapy of the human retina">Adeno associated virus and gene therapy of the human retina</a></div> <p><a href="/wiki/Gene_therapy" title="Gene therapy">Gene therapy</a> holds promise as a potential avenue to cure a wide range of retinal diseases. This involves using a non-infectious virus to shuttle a gene into a part of the retina. Recombinant <a href="/wiki/Adeno-associated_virus" title="Adeno-associated virus">adeno-associated virus</a> (rAAV) vectors possess a number of features that render them ideally suited for retinal gene therapy, including a lack of pathogenicity, minimal immunogenicity, and the ability to transduce postmitotic cells in a stable and efficient manner.<sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> rAAV vectors are increasingly utilized for their ability to mediate efficient transduction of <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a> (RPE), <a href="/wiki/Photoreceptor_cells" class="mw-redirect" title="Photoreceptor cells">photoreceptor cells</a> and <a href="/wiki/Retinal_ganglion_cells" class="mw-redirect" title="Retinal ganglion cells">retinal ganglion cells</a>. Each cell type can be specifically targeted by choosing the appropriate combination of AAV <a href="/wiki/Serotype" title="Serotype">serotype</a>, promoter, and intraocular injection site. </p><p>Several clinical trials have already reported positive results using rAAV to treat <a href="/wiki/Leber%27s_congenital_amaurosis" class="mw-redirect" title="Leber's congenital amaurosis">Leber's congenital amaurosis</a>, showing that the therapy was both safe and effective.<sup id="cite_ref-Cideciyan_2009_66-0" class="reference"><a href="#cite_note-Cideciyan_2009-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Simonelli_2010_67-0" class="reference"><a href="#cite_note-Simonelli_2010-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> There were no serious adverse events, and patients in all three studies showed improvement in their visual function as measured by a number of methods. The methods used varied among the three trials, but included both functional methods such as visual acuity<sup id="cite_ref-Simonelli_2010_67-1" class="reference"><a href="#cite_note-Simonelli_2010-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Maguire_2008_68-0" class="reference"><a href="#cite_note-Maguire_2008-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Maguire_2009_69-0" class="reference"><a href="#cite_note-Maguire_2009-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> and functional mobility<sup id="cite_ref-Maguire_2008_68-1" class="reference"><a href="#cite_note-Maguire_2008-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Maguire_2009_69-1" class="reference"><a href="#cite_note-Maguire_2009-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bainbridge_2008_70-0" class="reference"><a href="#cite_note-Bainbridge_2008-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> as well as objective measures that are less susceptible to bias, such as the pupil's ability to respond to light<sup id="cite_ref-Cideciyan_2009_66-1" class="reference"><a href="#cite_note-Cideciyan_2009-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hauswirth_2008_71-0" class="reference"><a href="#cite_note-Hauswirth_2008-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> and improvements on functional MRI.<sup id="cite_ref-Ashtari_2011_72-0" class="reference"><a href="#cite_note-Ashtari_2011-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> Improvements were sustained over the long-term, with patients continuing to do well after more than 1.5 years.<sup id="cite_ref-Cideciyan_2009_66-2" class="reference"><a href="#cite_note-Cideciyan_2009-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Simonelli_2010_67-2" class="reference"><a href="#cite_note-Simonelli_2010-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p><p>The unique architecture of the retina and its relatively immune-privileged environment help this process.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Tight_junction" title="Tight junction">Tight junctions</a> that form the <a href="/wiki/Blood_retinal_barrier" class="mw-redirect" title="Blood retinal barrier">blood retinal barrier</a> separate the subretinal space from the blood supply, thus protecting it from microbes and most immune-mediated damage, and enhancing its potential to respond to vector-mediated therapies. The highly compartmentalized anatomy of the eye facilitates accurate delivery of therapeutic vector suspensions to specific tissues under direct visualization using microsurgical techniques.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> In the sheltered environment of the retina, AAV vectors are able to maintain high levels of <a href="/wiki/Transgene" title="Transgene">transgene</a> expression in the retinal pigmented epithelium (RPE), photoreceptors, or ganglion cells for long periods of time after a single treatment. In addition, the eye and the visual system can be routinely and easily monitored for visual function and retinal structural changes after injections with noninvasive advanced technology, such as visual acuities, <a href="/wiki/Contrast_sensitivity" class="mw-redirect" title="Contrast sensitivity">contrast sensitivity</a>, <a href="/wiki/Fundus_(eye)" title="Fundus (eye)">fundus</a> auto-fluorescence (FAF), dark-adapted visual thresholds, vascular diameters, pupillometry, <a href="/wiki/Electroretinography" title="Electroretinography">electroretinography</a> (ERG), multifocal ERG and <a href="/wiki/Optical_coherence_tomography" title="Optical coherence tomography">optical coherence tomography</a> (OCT).<sup id="cite_ref-Hollander2008_75-0" class="reference"><a href="#cite_note-Hollander2008-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> </p><p>This strategy is effective against a number of retinal diseases that have been studied, including neovascular diseases that are features of <a href="/wiki/Age-related_macular_degeneration" class="mw-redirect" title="Age-related macular degeneration">age-related macular degeneration</a>, <a href="/wiki/Diabetic_retinopathy" title="Diabetic retinopathy">diabetic retinopathy</a> and <a href="/wiki/Retinopathy_of_prematurity" title="Retinopathy of prematurity">retinopathy of prematurity</a>. Since the regulation of vascularization in the mature retina involves a balance between endogenous positive <a href="/wiki/Growth_factors" class="mw-redirect" title="Growth factors">growth factors</a>, such as <a href="/wiki/Vascular_endothelial_growth_factor" title="Vascular endothelial growth factor">vascular endothelial growth factor</a> (VEGF) and inhibitors of <a href="/wiki/Angiogenesis" title="Angiogenesis">angiogenesis</a>, such as pigment epithelium-derived factor (<a href="/wiki/PEDF" title="PEDF">PEDF</a>), rAAV-mediated expression of PEDF, angiostatin, and the soluble VEGF receptor sFlt-1, which are all antiangiogenic proteins, have been shown to reduce aberrant vessel formation in animal models.<sup id="cite_ref-Rolling2004_76-0" class="reference"><a href="#cite_note-Rolling2004-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> Since specific gene therapies cannot readily be used to treat a significant fraction of patients with retinal dystrophy, there is a major interest in developing a more generally applicable survival factor therapy. <a href="/wiki/Neurotrophic_factors" title="Neurotrophic factors">Neurotrophic factors</a> have the ability to modulate neuronal growth during development to maintain existing cells and to allow recovery of injured neuronal populations in the eye. AAV encoding neurotrophic factors such as fibroblast growth factor (FGF) family members and GDNF either protected photoreceptors from apoptosis or slowed down cell death.<sup id="cite_ref-Rolling2004_76-1" class="reference"><a href="#cite_note-Rolling2004-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> </p><p><b>Organ transplantation</b> <a href="/wiki/Organ_transplant" class="mw-redirect" title="Organ transplant">Transplantation</a> of retinas has been attempted, but without much success. At <a href="/wiki/Massachusetts_Institute_of_Technology" title="Massachusetts Institute of Technology">MIT</a>, The University of Southern California, RWTH Aachen University, and the <a href="/wiki/University_of_New_South_Wales" title="University of New South Wales">University of New South Wales</a>, an "artificial retina" is under development: an implant which will bypass the photoreceptors of the retina and stimulate the attached nerve cells directly, with signals from a digital camera. </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=17" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Around 300 <a href="/wiki/BCE" class="mw-redirect" title="BCE">BCE</a>, <a href="/wiki/Herophilos" title="Herophilos">Herophilos</a> identified the retina from dissections of cadaver eyes. He called it the <i>arachnoid</i> layer, from its resemblance to a spider web, and <i>retiform</i>, from its resemblance to a casting net. The term <a href="/wiki/Arachnoid_mater" title="Arachnoid mater">arachnoid</a> came to refer to a layer around the brain; the term <i>retiform</i> came to refer to the <i>retina</i>.<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> </p><p>Between 1011 and 1021 CE, <a href="/wiki/Ibn_Al-Haytham" class="mw-redirect" title="Ibn Al-Haytham">Ibn Al-Haytham</a> published numerous experiments demonstrating that sight occurs from light reflecting from objects into the eye. This is consistent with <a href="/wiki/Intromission_theory" class="mw-redirect" title="Intromission theory">intromission theory</a> and against <a href="/wiki/Emission_theory_(vision)" title="Emission theory (vision)">emission theory</a>, the theory that sight occurs from rays emitted by the eyes. However, Ibn Al-Haytham decided that the retina could not be responsible for the beginnings of vision because the image formed on it was inverted. Instead he decided it must begin at the surface of the lens.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p><p>In 1604, <a href="/wiki/Johannes_Kepler" title="Johannes Kepler">Johannes Kepler</a> worked out the optics of the eye and decided that the retina must be where sight begins. He left it up to other scientists to reconcile the inverted retinal image with our perception of the world as upright.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p><p>In 1894, <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a> published the first major characterization of retinal neurons in <i>Retina der Wirbelthiere</i> (<i>The Retina of Vertebrates</i>).<sup id="cite_ref-nobelprize.org_80-0" class="reference"><a href="#cite_note-nobelprize.org-80"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/George_Wald" title="George Wald">George Wald</a>, <a href="/wiki/Haldan_Keffer_Hartline" title="Haldan Keffer Hartline">Haldan Keffer Hartline</a>, and <a href="/wiki/Ragnar_Granit" title="Ragnar Granit">Ragnar Granit</a> won the 1967 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> for their scientific research on the retina.<sup id="cite_ref-81" class="reference"><a href="#cite_note-81"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p><p>A recent <a href="/wiki/University_of_Pennsylvania" title="University of Pennsylvania">University of Pennsylvania</a> study calculated that the approximate <a href="/wiki/Bandwidth_(computing)" title="Bandwidth (computing)">bandwidth</a> of human retinas is 8.75 megabits per second, whereas a <a href="/wiki/Guinea_pig" title="Guinea pig">guinea pig</a>'s retinal transfer rate is 875 kilobits per second.<sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Robert_MacLaren" title="Robert MacLaren">MacLaren</a> & Pearson and colleagues at <a href="/wiki/University_College_London" title="University College London">University College London</a> and <a href="/wiki/Moorfields_Eye_Hospital" title="Moorfields Eye Hospital">Moorfields Eye Hospital</a> in London, in 2006, showed that <a href="/wiki/Photoreceptor_cells" class="mw-redirect" title="Photoreceptor cells">photoreceptor cells</a> could be transplanted successfully in the mouse retina if donor cells were at a critical developmental stage.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> Recently Ader and colleagues in Dublin showed, using the electron microscope, that transplanted photoreceptors formed synaptic connections.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> </p><p>In 2012, <a href="/wiki/Sebastian_Seung" title="Sebastian Seung">Sebastian Seung</a> and his laboratory at <a href="/wiki/Massachusetts_Institute_of_Technology" title="Massachusetts Institute of Technology">MIT</a> launched <a href="/wiki/EyeWire" class="mw-redirect" title="EyeWire">EyeWire</a>, an online <a href="/wiki/Citizen_science" title="Citizen science">Citizen science</a> game where players trace neurons in the retina.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> The goals of the EyeWire project are to identify specific cell types within the known broad classes of retinal cells, and to <a href="/wiki/Brain_mapping" title="Brain mapping">map the connections</a> between neurons in the retina, which will help to determine how vision works.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Additional_images">Additional images</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=18" title="Edit section: Additional images"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-traditional center"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Three_Main_Layers_of_the_Eye.png" class="mw-file-description" title="The structures of the eye labeled"><img alt="The structures of the eye labeled" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Three_Main_Layers_of_the_Eye.png/120px-Three_Main_Layers_of_the_Eye.png" decoding="async" width="120" height="94" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Three_Main_Layers_of_the_Eye.png/180px-Three_Main_Layers_of_the_Eye.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d0/Three_Main_Layers_of_the_Eye.png/240px-Three_Main_Layers_of_the_Eye.png 2x" data-file-width="765" data-file-height="599" /></a></span></div> <div class="gallerytext">The structures of the eye labeled</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Three_Internal_chambers_of_the_Eye.svg" class="mw-file-description" title="Another view of the eye and the structures of the eye labeled"><img alt="Another view of the eye and the structures of the eye labeled" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Three_Internal_chambers_of_the_Eye.svg/117px-Three_Internal_chambers_of_the_Eye.svg.png" decoding="async" width="117" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Three_Internal_chambers_of_the_Eye.svg/176px-Three_Internal_chambers_of_the_Eye.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e9/Three_Internal_chambers_of_the_Eye.svg/235px-Three_Internal_chambers_of_the_Eye.svg.png 2x" data-file-width="704" data-file-height="720" /></a></span></div> <div class="gallerytext">Another view of the eye and the structures of the eye labeled</div> </li> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Retinal_Image.png" class="mw-file-description" title="Illustration of image as 'seen' by the retina independent of optic nerve and striate cortex processing"><img alt="Illustration of image as 'seen' by the retina independent of optic nerve and striate cortex processing" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Retinal_Image.png/120px-Retinal_Image.png" decoding="async" width="120" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Retinal_Image.png/180px-Retinal_Image.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Retinal_Image.png/240px-Retinal_Image.png 2x" data-file-width="2048" data-file-height="2048" /></a></span></div> <div class="gallerytext">Illustration of image as 'seen' by the retina independent of optic nerve and striate cortex processing</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=19" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1266661725">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><span class="portalbox-image"><span class="noviewer" typeof="mw:File"><span><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/12px-WHO_Rod.svg.png" decoding="async" width="12" height="28" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/18px-WHO_Rod.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/WHO_Rod.svg/24px-WHO_Rod.svg.png 2x" data-file-width="107" data-file-height="250" /></span></span></span><span class="portalbox-link"><a href="/wiki/Portal:Medicine" title="Portal:Medicine">Medicine portal</a></span></li></ul> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">This article uses <a href="/wiki/Anatomical_terminology" title="Anatomical terminology">anatomical terminology</a>.</div> <ul><li><a href="/wiki/Adeno_associated_virus_and_gene_therapy_of_the_human_retina" class="mw-redirect" title="Adeno associated virus and gene therapy of the human retina">Adeno associated virus and gene therapy of the human retina</a></li> <li><a href="/wiki/Charles_Schepens" title="Charles Schepens">Charles Schepens</a> – "the father of modern retinal surgery"</li> <li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Evolution of the eye</a></li> <li><a href="/wiki/Duplex_retina" title="Duplex retina">Duplex retina</a></li> <li><a href="/wiki/Retinal_scan" title="Retinal scan">Retinal scan</a></li> <li><a href="/wiki/Retinal_vein_occlusion" class="mw-redirect" title="Retinal vein occlusion">Retinal vein occlusion</a></li> <li><a href="/wiki/List_of_xanthoma_variants_associated_with_hyperlipoproteinemia_subtypes" title="List of xanthoma variants associated with hyperlipoproteinemia subtypes">List of xanthoma variants associated with hyperlipoproteinemia subtypes</a></li> <li><a href="/wiki/Rhodopsin" title="Rhodopsin">Rhodopsin</a></li> <li><a href="/wiki/Persistence_of_vision" title="Persistence of vision">Persistence of vision</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Retina&action=edit&section=20" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></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="CITEREFJ2005" class="citation book cs1">J, Krause William (2005). <i>Krause's Essential Human Histology for Medical Students</i>. 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Ramón y Cajal</a>, <i>Histologie du Système Nerveux de l'Homme et des Vertébrés</i>, Maloine, Paris, 1911.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFRodieck_RW1965" class="citation journal cs1">Rodieck RW (1965). 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Rev</i>. <b>71</b> (2): <span class="nowrap">447–</span>480. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1152%2Fphysrev.1991.71.2.447">10.1152/physrev.1991.71.2.447</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/2006220">2006220</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Physiol.+Rev.&rft.atitle=Functional+architecture+of+the+mammalian+retina&rft.volume=71&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E447-%3C%2Fspan%3E480&rft.date=1991&rft_id=info%3Adoi%2F10.1152%2Fphysrev.1991.71.2.447&rft_id=info%3Apmid%2F2006220&rft.aulast=W%C3%A4ssle&rft.aufirst=H&rft.au=Boycott%2C+BB&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARetina" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFSchulzGoetzKaschkoetoeWeber2004" class="citation journal cs1">Schulz HL, Goetz T, Kaschkoetoe J, Weber BH (2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC512282">"The Retinome – Defining a reference transcriptome of the adult mammalian retina/retinal pigment epithelium"</a>. <i>BMC Genomics</i> (about a <a href="/wiki/Transcriptome" title="Transcriptome">transcriptome</a> for eye colour). <b>5</b> (1): 50. <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/10.1186%2F1471-2164-5-50">10.1186/1471-2164-5-50</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/PMC512282">512282</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/15283859">15283859</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BMC+Genomics&rft.atitle=The+Retinome+%E2%80%93+Defining+a+reference+transcriptome+of+the+adult+mammalian+retina%2Fretinal+pigment+epithelium&rft.volume=5&rft.issue=1&rft.pages=50&rft.date=2004&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC512282%23id-name%3DPMC&rft_id=info%3Apmid%2F15283859&rft_id=info%3Adoi%2F10.1186%2F1471-2164-5-50&rft.aulast=Schulz&rft.aufirst=HL&rft.au=Goetz%2C+T&rft.au=Kaschkoetoe%2C+J&rft.au=Weber%2C+BH&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC512282&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARetina" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFDowling2007" class="citation journal cs1">Dowling, John (2007). <a rel="nofollow" class="external text" href="https://doi.org/10.4249%2Fscholarpedia.3487">"Retina"</a>. <i><a href="/wiki/Scholarpedia" title="Scholarpedia">Scholarpedia</a></i>. <b>2</b> (12): 3487. <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/2007SchpJ...2.3487D">2007SchpJ...2.3487D</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/10.4249%2Fscholarpedia.3487">10.4249/scholarpedia.3487</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scholarpedia&rft.atitle=Retina&rft.volume=2&rft.issue=12&rft.pages=3487&rft.date=2007&rft_id=info%3Adoi%2F10.4249%2Fscholarpedia.3487&rft_id=info%3Abibcode%2F2007SchpJ...2.3487D&rft.aulast=Dowling&rft.aufirst=John&rft_id=https%3A%2F%2Fdoi.org%2F10.4249%252Fscholarpedia.3487&rfr_id=info%3Asid%2Fen.wikipedia.org%3ARetina" class="Z3988"></span></li></ul> <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=Retina&action=edit&section=22" 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="https://www.youtube.com/watch?v=nELWup_Gghw">Histology of the Eye</a>, edited by William Krause, Dept. Pathology and Anatomical science, University of Missouri School of Medicine</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20070314135325/http://neuro.med.harvard.edu/site/dh/index.html">Eye, Brain, and Vision – online book – by David Hubel</a></li> <li>Kolb, H., Fernandez, E., & Nelson, R. (2003). <a rel="nofollow" class="external text" href="http://webvision.med.utah.edu/">Webvision: The neural organization of the vertebrate retina</a>. Salt Lake City, Utah: John Moran Eye Center, <a href="/wiki/University_of_Utah" title="University of Utah">University of Utah</a>. Retrieved 22 July 2014.</li> <li><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=neurosci.figgrp.740">Retinal layers image.</a> NeuroScience 2nd Ed at <a href="/wiki/United_States_National_Library_of_Medicine" title="United States National Library of Medicine">United States National Library of Medicine</a></li> <li><a rel="nofollow" class="external text" href="https://www.ibiology.org/neuroscience/color-vision/">Jeremy Nathans's Seminars: "The Vertebrate Retina: Structure, Function, and Evolution"</a></li> <li><a rel="nofollow" class="external text" href="http://ccdb.ucsd.edu/sand/main?mpid=54&event=displaySum">Retina – Cell Centered Database</a></li> <li><a rel="nofollow" class="external text" href="https://www.bu.edu/phpbin/medlib/histology/p/07901loa.htm">Histology image: 07901loa</a> – Histology Learning System at Boston University</li> <li><i><a href="/wiki/MedlinePlus" title="MedlinePlus">MedlinePlus Encyclopedia</a></i>: <a 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eye)">globe</a> of the <a href="/wiki/Human_eye" title="Human eye">human eye</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Fibrous_tunic_of_eyeball" title="Fibrous tunic of eyeball">Fibrous tunic</a> <br />(outer)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sclera" title="Sclera">Sclera</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/Episcleral_layer" title="Episcleral layer">Episcleral layer</a></li> <li><a href="/wiki/Schlemm%27s_canal" title="Schlemm's canal">Schlemm's canal</a></li> <li><a href="/wiki/Trabecular_meshwork" title="Trabecular meshwork">Trabecular meshwork</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cornea" title="Cornea">Cornea</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/Corneal_limbus" title="Corneal limbus">Limbus</a></li> <li><i>layers</i> <ul><li><a href="/wiki/Corneal_epithelium" title="Corneal epithelium">Epithelium</a></li> <li><a href="/wiki/Bowman%27s_layer" title="Bowman's layer">Bowman's</a></li> <li><a href="/wiki/Stroma_of_cornea" title="Stroma of cornea">Stroma</a></li> <li><a href="/wiki/Dua%27s_layer" title="Dua's layer">Dua's layer</a></li> <li><a href="/wiki/Descemet%27s_membrane" title="Descemet's membrane">Descemet's</a></li> <li><a href="/wiki/Corneal_endothelium" title="Corneal endothelium">Endothelium</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><figure class="noresize mw-ext-imagemap-desc-bottom-right" typeof="mw:File"><span title="1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina"><img alt="1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Eye-diagram_no_circles_border.svg/200px-Eye-diagram_no_circles_border.svg.png" decoding="async" width="200" height="195" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Eye-diagram_no_circles_border.svg/300px-Eye-diagram_no_circles_border.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Eye-diagram_no_circles_border.svg/400px-Eye-diagram_no_circles_border.svg.png 2x" data-file-width="1237" data-file-height="1208" usemap="#ImageMap_0a6142ee86469b4f" resource="/wiki/File:Eye-diagram_no_circles_border.svg" /></span><map name="ImageMap_0a6142ee86469b4f"><area href="/wiki/Posterior_segment_of_eyeball" shape="circle" coords="50,21,7" alt="1: posterior segment" title="1: posterior segment" /><area href="/wiki/Ora_serrata" shape="circle" coords="38,30,7" alt="2: ora serrata" title="2: ora serrata" /><area href="/wiki/Ciliary_muscle" shape="circle" coords="28,40,7" alt="3: ciliary muscle" title="3: ciliary muscle" /><area href="/wiki/Zonule_of_Zinn" shape="circle" coords="20,53,7" alt="4: ciliary zonules" title="4: ciliary zonules" /><area href="/wiki/Schlemm%27s_canal" shape="circle" coords="12,66,7" alt="5: Schlemm's canal" title="5: Schlemm's canal" /><area href="/wiki/Pupil" shape="circle" coords="9,80,7" alt="6: pupil" title="6: pupil" /><area href="/wiki/Anterior_chamber_of_eyeball" shape="circle" coords="7,96,7" alt="7: anterior chamber" title="7: anterior chamber" /><area href="/wiki/Cornea" shape="circle" coords="7,111,7" alt="8: cornea" title="8: cornea" /><area href="/wiki/Iris_(anatomy)" shape="circle" coords="9,125,7" alt="9: iris" title="9: iris" /><area href="/wiki/Lens_cortex" shape="circle" coords="15,139,7" alt="10: lens cortex" title="10: lens cortex" /><area href="/wiki/Lens_nucleus" shape="circle" coords="22,152,7" alt="11: lens nucleus" title="11: lens nucleus" /><area href="/wiki/Ciliary_processes" shape="circle" coords="30,163,7" alt="12: ciliary process" title="12: ciliary process" /><area href="/wiki/Conjunctiva" shape="circle" coords="41,173,7" alt="13: conjunctiva" title="13: conjunctiva" /><area href="/wiki/Inferior_oblique_muscle" shape="circle" coords="133,189,7" alt="14: inferior oblique muscule" title="14: inferior oblique muscule" /><area href="/wiki/Inferior_rectus_muscle" shape="circle" coords="147,182,7" alt="15: inferior rectus muscule" title="15: inferior rectus muscule" /><area href="/wiki/Medial_rectus_muscle" shape="circle" coords="160,174,7" alt="16: medial rectus muscle" title="16: medial rectus muscle" /><area href="/wiki/Retinal_arteries_and_veins" shape="circle" coords="171,163,7" alt="17: retinal arteries and veins" title="17: retinal arteries and veins" /><area href="/wiki/Optic_disc" shape="circle" coords="180,152,7" alt="18: optic disc" title="18: optic disc" /><area href="/wiki/Dura_mater" shape="circle" coords="187,139,7" alt="19: dura mater" title="19: dura mater" /><area href="/wiki/Central_retinal_artery" shape="circle" coords="191,125,7" alt="20: central retinal artery" title="20: central retinal artery" /><area href="/wiki/Central_retinal_vein" shape="circle" coords="188,66,7" alt="21: central retinal vein" title="21: central retinal vein" /><area href="/wiki/Optic_nerve" shape="circle" coords="182,53,7" alt="22: optic nerve" title="22: optic nerve" /><area href="/wiki/Vorticose_veins" shape="circle" coords="174,41,7" alt="23: vorticose vein" title="23: vorticose vein" /><area href="/wiki/Tenon%27s_capsule" shape="circle" coords="162,30,7" alt="24: bulbar sheath" title="24: bulbar sheath" /><area href="/wiki/Macula_of_retina" shape="circle" coords="151,22,7" alt="25: macula" title="25: macula" /><area href="/wiki/Fovea_centralis" shape="circle" coords="136,13,7" alt="26: fovea" title="26: fovea" /><area href="/wiki/Sclera" shape="circle" coords="121,8,7" alt="27: sclera" title="27: sclera" /><area href="/wiki/Choroid" shape="circle" coords="107,7,7" alt="28: choroid" title="28: choroid" /><area href="/wiki/Superior_rectus_muscle" shape="circle" coords="93,7,7" alt="29: superior rectus muscle" title="29: superior rectus muscle" /><area href="/wiki/Retina" shape="circle" coords="79,8,7" alt="30: retina" title="30: retina" /></map><figcaption>1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina</figcaption></figure></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Uvea" title="Uvea">Uvea / vascular <br />tunic</a> (middle)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Choroid" title="Choroid">Choroid</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/Capillary_lamina_of_choroid" title="Capillary lamina of choroid">Capillary lamina of choroid</a></li> <li><a href="/wiki/Bruch%27s_membrane" title="Bruch's membrane">Bruch's membrane</a></li> <li><a href="/wiki/Sattler%27s_layer" title="Sattler's layer">Sattler's layer</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ciliary_body" title="Ciliary body">Ciliary body</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/Ciliary_processes" title="Ciliary processes">Ciliary processes</a></li> <li><a href="/wiki/Ciliary_muscle" title="Ciliary muscle">Ciliary muscle</a></li> <li><a href="/wiki/Pars_plicata" title="Pars plicata">Pars plicata</a></li> <li><a href="/wiki/Pars_plana" title="Pars plana">Pars plana</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Iris_(anatomy)" title="Iris (anatomy)">Iris</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/Stroma_of_iris" title="Stroma of iris">Stroma</a></li> <li><a href="/wiki/Pupil" title="Pupil">Pupil</a></li> <li><a href="/wiki/Iris_dilator_muscle" title="Iris dilator muscle">Iris dilator muscle</a></li> <li><a href="/wiki/Iris_sphincter_muscle" title="Iris sphincter muscle">Iris sphincter muscle</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a class="mw-selflink selflink">Retina</a> (inner)</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Layers</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/Inner_limiting_membrane" class="mw-redirect" title="Inner limiting membrane">Inner limiting membrane</a></li> <li><a href="/wiki/Nerve_fiber_layer" class="mw-redirect" title="Nerve fiber layer">Nerve fiber layer</a></li> <li><a href="/wiki/Ganglion_cell_layer" title="Ganglion cell layer">Ganglion cell layer</a></li> <li><a href="/wiki/Inner_plexiform_layer" title="Inner plexiform layer">Inner plexiform layer</a></li> <li><a href="/wiki/Inner_nuclear_layer" title="Inner nuclear layer">Inner nuclear layer</a></li></ul> <ul><li><a href="/wiki/Outer_plexiform_layer" title="Outer plexiform layer">Outer plexiform layer</a></li> <li><a href="/wiki/Outer_nuclear_layer" title="Outer nuclear layer">Outer nuclear layer</a></li></ul> <ul><li><a href="/wiki/External_limiting_membrane" title="External limiting membrane">External limiting membrane</a></li> <li><a href="/wiki/Layer_of_rods_and_cones" title="Layer of rods and cones">Layer of rods and cones</a></li> <li><a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">Retinal pigment epithelium</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cells</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor cells</a> (<a href="/wiki/Cone_cell" title="Cone cell">Cone cell</a>, <a href="/wiki/Rod_cell" title="Rod cell">Rod cell</a>) → (<a href="/wiki/Retina_horizontal_cell" title="Retina horizontal cell">Horizontal cell</a>) → <a href="/wiki/Retina_bipolar_cell" title="Retina bipolar cell">Bipolar cell</a> → (<a href="/wiki/Amacrine_cell" title="Amacrine cell">Amacrine cell</a>) → <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">Retina ganglion cell</a> (<a href="/wiki/Midget_cell" title="Midget cell">Midget cell</a>, <a href="/wiki/Parasol_cell" title="Parasol cell">Parasol cell</a>, <a href="/wiki/Bistratified_cell" title="Bistratified cell">Bistratified cell</a>, <a href="/wiki/Giant_retinal_ganglion_cells" title="Giant retinal ganglion cells">Giant retina ganglion cells</a>, <a href="/wiki/Intrinsically_photosensitive_retinal_ganglion_cells" class="mw-redirect" title="Intrinsically photosensitive retinal ganglion cells">Photosensitive ganglion cell</a>) → <i>Diencephalon</i>: <a href="/wiki/Parvocellular_cell" title="Parvocellular cell">P cell</a>, <a href="/wiki/Magnocellular_cell" title="Magnocellular cell">M cell</a>, <a href="/wiki/Koniocellular_cell" title="Koniocellular cell">K cell</a>, <a href="/wiki/Muller_glia" class="mw-redirect" title="Muller glia">Muller glia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Macula_of_retina" class="mw-redirect" title="Macula of retina">Macula</a> <ul><li><a href="/wiki/Perifovea" title="Perifovea">Perifoveal area</a></li> <li><a href="/wiki/Parafovea" title="Parafovea">Parafoveal area</a></li> <li><a href="/wiki/Fovea_centralis" title="Fovea centralis">Fovea</a> <ul><li><a href="/wiki/Foveal_avascular_zone" title="Foveal avascular zone">Foveal avascular zone</a></li> <li><a href="/wiki/Foveola" title="Foveola">Foveola</a></li></ul></li></ul></li> <li><a href="/wiki/Optic_disc" title="Optic disc">Optic disc</a> <ul><li><a href="/wiki/Optic_cup_(anatomical)" title="Optic cup (anatomical)">Optic cup</a></li></ul></li> <li><a href="/wiki/Ora_serrata" title="Ora serrata">Ora serrata</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Anatomical regions <br />of the eye</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Anterior_segment_of_eyeball" title="Anterior segment of eyeball">Anterior segment</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/Accessory_visual_structures" title="Accessory visual structures">Adnexa</a> (<a href="/wiki/Eyebrow" title="Eyebrow">Eyebrow</a>, <a href="/wiki/Eyelid" title="Eyelid">Eyelid</a>, <a href="/wiki/Conjunctiva" title="Conjunctiva">Conjunctiva</a>, <a href="/wiki/Lacrimal_system" class="mw-redirect" title="Lacrimal system">Lacrimal system</a>, <a href="/wiki/Orbit_(anatomy)" title="Orbit (anatomy)">Orbit</a>)</li> <li><a href="/wiki/Fibrous_tunic" class="mw-redirect" title="Fibrous tunic">Fibrous tunic</a></li> <li><a href="/wiki/Anterior_chamber_of_eyeball" title="Anterior chamber of eyeball">Anterior chamber</a></li> <li><a href="/wiki/Aqueous_humour" title="Aqueous humour">Aqueous humour</a></li> <li><a href="/wiki/Iris_(anatomy)" title="Iris (anatomy)">Iris</a></li> <li><a href="/wiki/Posterior_chamber_of_eyeball" title="Posterior chamber of eyeball">Posterior chamber</a></li> <li><a href="/wiki/Ciliary_body" title="Ciliary body">Ciliary body</a></li> <li><a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">Lens</a> <ul><li><a href="/wiki/Capsule_of_lens" title="Capsule of lens">Capsule of lens</a></li> <li><a href="/wiki/Zonule_of_Zinn" title="Zonule of Zinn">Zonule of Zinn</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Posterior_segment_of_eyeball" title="Posterior segment of eyeball">Posterior segment</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/Vitreous_chamber" title="Vitreous chamber">Vitreous chamber</a> <ul><li><a href="/wiki/Vitreous_body" title="Vitreous body">Vitreous body</a></li></ul></li> <li><a class="mw-selflink selflink">Retina</a></li> <li><a href="/wiki/Choroid" title="Choroid">Choroid</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Corneal_keratocyte" title="Corneal keratocyte">Keratocytes</a></li> <li><a href="/wiki/Ocular_immune_system" title="Ocular immune system">Ocular immune system</a></li> <li><a href="/wiki/Optical_coherence_tomography#Ophthalmology" title="Optical coherence tomography">Optical coherence tomography</a></li> <li><a href="/wiki/Eye_care_professional" title="Eye care professional">Eye care professional</a></li> <li><a href="/wiki/Eye_disease" title="Eye disease">Eye disease</a></li> <li><a href="/wiki/Refractive_error" title="Refractive error">Refractive error</a></li> <li><a href="/wiki/Accommodation_(eye)" class="mw-redirect" title="Accommodation (eye)">Accommodation</a></li> <li><i>Physiological Optics</i></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235" /></div><div role="navigation" class="navbox" aria-labelledby="Optical_illusions_(list)89" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231" /><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Optical_illusions" title="Template:Optical illusions"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Optical_illusions" title="Template talk:Optical illusions"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Optical_illusions" title="Special:EditPage/Template:Optical illusions"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Optical_illusions_(list)89" style="font-size:114%;margin:0 4em"><a href="/wiki/Optical_illusion" title="Optical illusion">Optical illusions</a> (<a href="/wiki/List_of_optical_illusions" title="List of optical illusions">list</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Illusions</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/Afterimage" title="Afterimage">Afterimage</a></li> <li><a href="/wiki/Ambigram" title="Ambigram">Ambigram</a></li> <li><a href="/wiki/Ambiguous_image" title="Ambiguous image">Ambiguous image</a></li> <li><a href="/wiki/Ames_room" title="Ames room">Ames room</a></li> <li><a href="/wiki/Autostereogram" title="Autostereogram">Autostereogram</a></li> <li><a href="/wiki/Barberpole_illusion" title="Barberpole illusion">Barberpole</a></li> <li><a href="/wiki/Bezold_effect" title="Bezold effect">Bezold</a></li> <li><a href="/wiki/Caf%C3%A9_wall_illusion" title="Café wall illusion">Café wall</a></li> <li><a href="/wiki/Checker_shadow_illusion" title="Checker shadow illusion">Checker shadow</a></li> <li><a href="/wiki/Chubb_illusion" title="Chubb illusion">Chubb</a></li> <li><a href="/wiki/Cornsweet_illusion" title="Cornsweet illusion">Cornsweet</a></li> <li><a href="/wiki/Delboeuf_illusion" title="Delboeuf illusion">Delboeuf</a></li> <li><a href="/wiki/Ebbinghaus_illusion" title="Ebbinghaus illusion">Ebbinghaus</a></li> <li><a href="/wiki/Ehrenstein_illusion" title="Ehrenstein illusion">Ehrenstein</a></li> <li><a href="/wiki/Flash_lag_illusion" title="Flash lag illusion">Flash lag</a></li> <li><a href="/wiki/Fraser_spiral_illusion" title="Fraser spiral illusion">Fraser spiral</a></li> <li><a href="/wiki/Gravity_hill" title="Gravity hill">Gravity hill</a></li> <li><a href="/wiki/Grid_illusion" title="Grid illusion">Grid</a></li> <li><a href="/wiki/Hering_illusion" title="Hering illusion">Hering</a></li> <li><a href="/wiki/Impossible_trident" title="Impossible trident">Impossible trident</a></li> <li><a href="/wiki/Irradiation_illusion" title="Irradiation illusion">Irradiation</a></li> <li><a href="/wiki/Jastrow_illusion" title="Jastrow illusion">Jastrow</a></li> <li><a href="/wiki/Lilac_chaser" title="Lilac chaser">Lilac chaser</a></li> <li><a href="/wiki/Mach_bands" title="Mach bands">Mach bands</a></li> <li><a href="/wiki/McCollough_effect" title="McCollough effect">McCollough</a></li> <li><a href="/wiki/M%C3%BCller-Lyer_illusion" title="Müller-Lyer illusion">Müller-Lyer</a></li> <li><a href="/wiki/Necker_cube" title="Necker cube">Necker cube</a></li> <li><a href="/wiki/Oppel-Kundt_illusion" class="mw-redirect" title="Oppel-Kundt illusion">Oppel-Kundt</a></li> <li><a href="/wiki/Orbison_illusion" title="Orbison illusion">Orbison</a></li> <li><a href="/wiki/Penrose_stairs" title="Penrose stairs">Penrose stairs</a></li> <li><a href="/wiki/Penrose_triangle" title="Penrose triangle">Penrose triangle</a></li> <li><a href="/wiki/Peripheral_drift_illusion" title="Peripheral drift illusion">Peripheral drift</a></li> <li><a href="/wiki/Poggendorff_illusion" title="Poggendorff illusion">Poggendorff</a></li> <li><a href="/wiki/Ponzo_illusion" title="Ponzo illusion">Ponzo</a></li> <li><a href="/wiki/Rubin_vase" title="Rubin vase">Rubin vase</a></li> <li><a href="/wiki/Sander_illusion" title="Sander illusion">Sander</a></li> <li><a href="/wiki/Schroeder_stairs" title="Schroeder stairs">Schroeder stairs</a></li> <li><a href="/wiki/Shepard_tables" title="Shepard tables">Shepard tables</a></li> <li><a href="/wiki/Spinning_dancer" title="Spinning dancer">Spinning dancer</a></li> <li><a href="/wiki/Ternus_illusion" title="Ternus illusion">Ternus</a></li> <li><a href="/wiki/Vertical%E2%80%93horizontal_illusion" title="Vertical–horizontal illusion">Vertical–horizontal</a></li> <li><a href="/wiki/White%27s_illusion" title="White's illusion">White's</a></li> <li><a href="/wiki/Wundt_illusion" title="Wundt illusion">Wundt</a></li> <li><a href="/wiki/Z%C3%B6llner_illusion" title="Zöllner illusion">Zöllner</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="3" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Optical_Illustion-Ambiguous_Patterns.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Optical_Illustion-Ambiguous_Patterns.svg/30px-Optical_Illustion-Ambiguous_Patterns.svg.png" decoding="async" width="30" height="112" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Optical_Illustion-Ambiguous_Patterns.svg/45px-Optical_Illustion-Ambiguous_Patterns.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Optical_Illustion-Ambiguous_Patterns.svg/60px-Optical_Illustion-Ambiguous_Patterns.svg.png 2x" data-file-width="117" data-file-height="438" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Popular culture</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/Op_art" title="Op art">Op art</a></li> <li><i><a href="/wiki/Trompe-l%27%C5%93il" title="Trompe-l'œil">Trompe-l'œil</a></i></li> <li><i><a href="/wiki/Spectropia" title="Spectropia">Spectropia</a></i> (1864 book)</li> <li><i><a href="/wiki/Ascending_and_Descending" title="Ascending and Descending">Ascending and Descending</a></i> (1960 drawing)</li> <li><i><a href="/wiki/Waterfall_(M._C._Escher)" title="Waterfall (M. C. Escher)">Waterfall</a></i> (1961 drawing)</li> <li><i><a href="/wiki/The_dress" title="The dress">The dress</a></i> (2015 photograph)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</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/Accidental_viewpoint" title="Accidental viewpoint">Accidental viewpoint</a></li> <li><a href="/wiki/Auditory_illusion" title="Auditory illusion">Auditory illusions</a></li> <li><a href="/wiki/Illusion" title="Illusion">Illusions</a></li> <li><a href="/wiki/Tactile_illusion" title="Tactile illusion">Tactile illusions</a></li> <li><a href="/wiki/Time_perception" title="Time perception">Temporal illusion</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374" /><link rel="mw-deduplicated-inline-style" 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