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class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Possible role in conscious sight</span> </div> </a> <ul id="toc-Possible_role_in_conscious_sight-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Violet-to-blue_light" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Violet-to-blue_light"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Violet-to-blue light</span> </div> </a> <ul id="toc-Violet-to-blue_light-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Discovery" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Discovery"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Discovery</span> </div> </a> <ul id="toc-Discovery-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Research" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a 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<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-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">8</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 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class="mw-page-title-main">Intrinsically photosensitive retinal ganglion cell</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. 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الجوهرية" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Fotoosjetljive_ganglijske_%C4%87elije" title="Fotoosjetljive ganglijske ćelije – Bosnian" lang="bs" hreflang="bs" data-title="Fotoosjetljive ganglijske ćelije" 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/C%C3%A8l%C2%B7lula_ganglionar_de_la_retina_intr%C3%ADnsecament_fotosensible" title="Cèl·lula ganglionar de la retina intrínsecament fotosensible – Catalan" lang="ca" hreflang="ca" data-title="Cèl·lula ganglionar de la retina intrínsecament fotosensible" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Fotosensitive_Ganglienzelle" title="Fotosensitive Ganglienzelle – German" lang="de" hreflang="de" data-title="Fotosensitive Ganglienzelle" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D9%84%D9%88%D9%84_%D9%87%D8%A7%DB%8C_%DA%AF%D8%A7%D9%86%DA%AF%D9%84%DB%8C%D9%88%D9%86%DB%8C_%D8%B0%D8%A7%D8%AA%D8%A7%D9%8B_%D8%AD%D8%B3%D8%A7%D8%B3_%D8%A8%D9%87_%D9%86%D9%88%D8%B1_%D8%B4%D8%A8%DA%A9%DB%8C%D9%87" title="سلول های گانگلیونی ذاتاً حساس به نور شبکیه – Persian" lang="fa" hreflang="fa" data-title="سلول های گانگلیونی ذاتاً حساس به نور شبکیه" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/C%C3%A9lula_ganglionar_da_retina_fotosensible" title="Célula ganglionar da retina fotosensible – Galician" lang="gl" hreflang="gl" data-title="Célula ganglionar da retina fotosensible" 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%82%B4%EC%9E%AC%EC%A0%81%EC%9D%B8_%EA%B0%90%EA%B4%91%EC%84%B1_%EB%A7%9D%EB%A7%89_%EC%8B%A0%EA%B2%BD%EC%A0%88_%EC%84%B8%ED%8F%AC" title="내재적인 감광성 망막 신경절 세포 – Korean" lang="ko" hreflang="ko" data-title="내재적인 감광성 망막 신경절 세포" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Lichtgevoelige_ganglioncel" title="Lichtgevoelige ganglioncel – Dutch" lang="nl" hreflang="nl" data-title="Lichtgevoelige ganglioncel" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Fotosenzitivne_ganglijske_%C4%87elije" title="Fotosenzitivne ganglijske ćelije – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Fotosenzitivne ganglijske ćelije" 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-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%85%A7%E5%9C%A8%E5%85%89%E6%95%8F%E8%A6%96%E7%B6%B2%E8%86%9C%E7%A5%9E%E7%B6%93%E7%AF%80%E7%B4%B0%E8%83%9E" title="內在光敏視網膜神經節細胞 – Chinese" lang="zh" 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class="mw-redirect" title="Photosensitive ganglion cell">Photosensitive ganglion cell</a>)</span></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Type of neuron in the retina of the mammalian eye</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Overview_of_the_retina_photoreceptors_(a).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Overview_of_the_retina_photoreceptors_%28a%29.png/400px-Overview_of_the_retina_photoreceptors_%28a%29.png" decoding="async" width="400" height="124" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/29/Overview_of_the_retina_photoreceptors_%28a%29.png/600px-Overview_of_the_retina_photoreceptors_%28a%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/29/Overview_of_the_retina_photoreceptors_%28a%29.png/800px-Overview_of_the_retina_photoreceptors_%28a%29.png 2x" data-file-width="1266" data-file-height="391" /></a><figcaption>Overview of the retina photoreceptors. ipRGCs labelled at the top-right.</figcaption></figure> <p><b>Intrinsically photosensitive retinal ganglion cells</b> (<b>ipRGCs</b>), also called <b>photosensitive retinal ganglion cells</b> (<b>pRGC</b>), or <b>melanopsin-containing retinal ganglion cells</b> (<b>mRGCs</b>), are a type of <a href="/wiki/Neuron" title="Neuron">neuron</a> in the <a href="/wiki/Retina" title="Retina">retina</a> of the <a href="/wiki/Mammalian_eye" title="Mammalian eye">mammalian eye</a>. The presence of an additional photoreceptor was first suspected in 1927 when mice lacking rods and cones still responded to changing light levels through <a href="/wiki/Pupil_constriction" class="mw-redirect" title="Pupil constriction">pupil constriction</a>;<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> this suggested that rods and cones are not the only light-sensitive tissue.<sup id="cite_ref-Keeler_2-0" class="reference"><a href="#cite_note-Keeler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> However, it was unclear whether this light sensitivity arose from an additional retinal photoreceptor or elsewhere in the body. Recent research has shown that these <a href="/wiki/Retinal_ganglion_cell" title="Retinal ganglion cell">retinal ganglion cells</a>, unlike other retinal ganglion cells, are intrinsically <a href="/wiki/Photosensitive" class="mw-redirect" title="Photosensitive">photosensitive</a> due to the presence of <a href="/wiki/Melanopsin" title="Melanopsin">melanopsin</a>, a light-sensitive protein. Therefore, they constitute a third class of photoreceptors, in addition to <a href="/wiki/Rod_cells" class="mw-redirect" title="Rod cells">rod</a> and <a href="/wiki/Cone_cells" class="mw-redirect" title="Cone cells">cone cells</a>.<sup id="cite_ref-:1_3-0" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=1" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Melanopsin_stain.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Melanopsin_stain.jpg/340px-Melanopsin_stain.jpg" decoding="async" width="340" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Melanopsin_stain.jpg/510px-Melanopsin_stain.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/58/Melanopsin_stain.jpg/680px-Melanopsin_stain.jpg 2x" data-file-width="1031" data-file-height="668" /></a><figcaption>An ipRGC, shown here as a complied image of the retina from proximal inner nuclear layer to the ganglion cell layer with fluorescent labeling of melanopsin</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Overview_of_the_retina_photoreceptors_(b).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Overview_of_the_retina_photoreceptors_%28b%29.png/340px-Overview_of_the_retina_photoreceptors_%28b%29.png" decoding="async" width="340" height="124" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Overview_of_the_retina_photoreceptors_%28b%29.png/510px-Overview_of_the_retina_photoreceptors_%28b%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7c/Overview_of_the_retina_photoreceptors_%28b%29.png/680px-Overview_of_the_retina_photoreceptors_%28b%29.png 2x" data-file-width="1181" data-file-height="432" /></a><figcaption>Spectral sensitivities of the photoreceptors in the human eye.<sup id="cite_ref-pmid31534436_4-0" class="reference"><a href="#cite_note-pmid31534436-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>Compared to the rods and cones, the ipRGCs respond more sluggishly and signal the presence of light over the long term.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> They represent a very small subset (~1%) of the retinal ganglion cells.<sup id="cite_ref-:3_6-0" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Their functional roles are non-image-forming and fundamentally different from those of pattern vision; they provide a stable representation of ambient light intensity. They have at least three primary functions: </p> <ul><li>They play a major role in synchronizing <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">circadian rhythms</a> to the 24-hour light/dark cycle, providing primarily length-of-day and length-of-night information. They send light information via the <a href="/wiki/Retinohypothalamic_tract" title="Retinohypothalamic tract">retinohypothalamic tract</a> (RHT) directly to the <a href="/wiki/Circadian" class="mw-redirect" title="Circadian">circadian</a> pacemaker of the <a href="/wiki/Brain" title="Brain">brain</a>, the <a href="/wiki/Suprachiasmatic_nucleus" title="Suprachiasmatic nucleus">suprachiasmatic nucleus</a> of the <a href="/wiki/Hypothalamus" title="Hypothalamus">hypothalamus</a>. The physiological properties of these ganglion cells match known properties of the daily light <a href="/wiki/Entrainment_(chronobiology)" title="Entrainment (chronobiology)">entrainment</a> (<a href="/wiki/Synchronization" title="Synchronization">synchronization</a>) mechanism regulating circadian rhythms. In addition, ipRGCs could also influence peripheral tissues such as the hair follicle regeneration through SCN-sympathetic nerve circuit.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup></li> <li>Photosensitive ganglion cells innervate other brain targets, such as the center of <a href="/wiki/Pupillary_reflex" title="Pupillary reflex">pupillary control</a>, the <a href="/wiki/Olivary_pretectal_nucleus" class="mw-redirect" title="Olivary pretectal nucleus">olivary pretectal nucleus</a> of the <a href="/wiki/Midbrain" title="Midbrain">midbrain</a>. They contribute to the regulation of <a href="/wiki/Pupil" title="Pupil">pupil</a> size and other behavioral responses to ambient lighting conditions.<sup id="cite_ref-Ecker_8-0" class="reference"><a href="#cite_note-Ecker-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li> <li>They contribute to photic regulation and acute photic suppression of release of the hormone <a href="/wiki/Melatonin" title="Melatonin">melatonin</a>.<sup id="cite_ref-Ecker_8-1" class="reference"><a href="#cite_note-Ecker-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li> <li>In rats, they play some role in conscious visual perception, including perception of regular gratings, light levels, and spatial information.<sup id="cite_ref-Ecker_8-2" class="reference"><a href="#cite_note-Ecker-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li></ul> <p>Photoreceptive ganglion cells have been isolated in humans, where, in addition to regulating the circadian rhythm, they have been shown to mediate a degree of light recognition in rodless, coneless subjects suffering with disorders of rod and cone photoreceptors.<sup id="cite_ref-pmid18082405_9-0" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Work by <a href="/w/index.php?title=Farhan_H._Zaidi&action=edit&redlink=1" class="new" title="Farhan H. Zaidi (page does not exist)">Farhan H. Zaidi</a> and colleagues showed that photoreceptive ganglion cells may have some visual function in humans. </p><p>The photopigment of photoreceptive ganglion cells, melanopsin, is excited by light mainly in the blue portion of the visible spectrum (absorption peaks at ~480 nanometers<sup id="cite_ref-Berson_10-0" class="reference"><a href="#cite_note-Berson-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>). The <a href="/wiki/Phototransduction" class="mw-redirect" title="Phototransduction">phototransduction</a> mechanism in these cells is not fully understood, but seems likely to resemble that in invertebrate <a href="/wiki/Rhabdom" class="mw-redirect" title="Rhabdom">rhabdomeric</a> photoreceptors. In addition to responding directly to light, these cells may receive excitatory and inhibitory influences from rods and cones by way of synaptic connections in the retina. </p><p>The axons from these ganglia innervate regions of the brain related to object recognition, including the <a href="/wiki/Superior_colliculus" title="Superior colliculus">superior colliculus</a> and dorsal <a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">lateral geniculate nucleus</a>.<sup id="cite_ref-Ecker_8-3" class="reference"><a href="#cite_note-Ecker-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <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=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=2" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="ipRGC_receptor">ipRGC receptor</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=3" title="Edit section: ipRGC receptor"><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:Melanopsin.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Melanopsin.png/220px-Melanopsin.png" decoding="async" width="220" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Melanopsin.png/330px-Melanopsin.png 1.5x, //upload.wikimedia.org/wikipedia/commons/7/79/Melanopsin.png 2x" data-file-width="350" data-file-height="356" /></a><figcaption>Melanopsin structure</figcaption></figure> <p>These photoreceptor cells project both throughout the retina and into the brain. They contain the photopigment melanopsin in varying quantities along the cell membrane, including on the axons up to the optic disc, the soma, and dendrites of the cell.<sup id="cite_ref-:1_3-1" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> ipRGCs contain membrane receptors for the neurotransmitters glutamate, <a href="/wiki/Glycine" title="Glycine">glycine</a>, and <a href="/wiki/Gamma-Aminobutyric_acid" class="mw-redirect" title="Gamma-Aminobutyric acid">GABA</a>.<sup id="cite_ref-:0_11-0" class="reference"><a href="#cite_note-:0-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Photosensitive ganglion cells respond to light by depolarizing, thus increasing the rate at which they fire nerve impulses, which is opposite to that of other photoreceptor cells, which hyperpolarize in response to light.<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>Results of studies in mice suggest that the axons of ipRGCs are <a href="/wiki/Myelin" title="Myelin">unmyelinated</a>.<sup id="cite_ref-:1_3-2" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Melanopsin">Melanopsin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=4" title="Edit section: Melanopsin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Visual_cycle" title="Visual cycle">Visual cycle</a></div> <p>Unlike other photoreceptor pigments, <a href="/wiki/Melanopsin" title="Melanopsin">melanopsin</a> has the ability to act as both the excitable photopigment and as a photoisomerase. Unlike the visual opsins in <a href="/wiki/Rod_cell" title="Rod cell">rod cells</a> and <a href="/wiki/Cone_cell" title="Cone cell">cone cells</a>, which rely on the standard <a href="/wiki/Visual_cycle" title="Visual cycle">visual cycles</a> for recharging <i>all-trans</i>-retinal back into the photosensitive <a href="/wiki/11-cis_retinal" class="mw-redirect" title="11-cis retinal"><i>11-cis</i>-retinal</a>, melanopsin is able to isomerize <i>all-trans-</i>retinal into <i>11-cis-</i>retinal itself when stimulated with another photon.<sup id="cite_ref-:0_11-1" class="reference"><a href="#cite_note-:0-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> An ipRGC therefore does not rely on <a href="/wiki/M%C3%BCller_glia" title="Müller glia">Müller cells</a> and/or <a href="/wiki/Retinal_pigment_epithelium" title="Retinal pigment epithelium">retinal pigment epithelium</a> cells for this conversion. </p><p>The two isoforms of melanopsin differ in their spectral sensitivity, for the <i>11-cis</i>-retinal isoform is more responsive to shorter wavelengths of light, while the <i>all-trans</i> isoform is more responsive to longer wavelengths of light.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Synaptic_inputs_and_outputs">Synaptic inputs and outputs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=5" title="Edit section: Synaptic inputs and outputs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg/900px-Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg" decoding="async" width="900" height="301" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/93/Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg/1350px-Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/9/93/Diagram_of_inputs_and_outputs_of_ipRGC_1.jpg 2x" data-file-width="1566" data-file-height="524" /></a><figcaption>Synaptic inputs and outputs of ipRGCs and their corresponding location in the brain</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Inputs">Inputs</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=6" title="Edit section: Inputs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>ipRGCs are both pre- and postsynaptic to dopaminergic <a href="/wiki/Amacrine_cell" title="Amacrine cell">amacrine cells</a> (DA cells) via reciprocal synapses, with ipRGCs sending excitatory signals to the DA cells, and the DA cells sending inhibitory signals to the ipRGCs. These inhibitory signals are mediated through <a href="/wiki/Gamma-aminobutyric_acid" class="mw-redirect" title="Gamma-aminobutyric acid">GABA</a>, which is co-released from the DA cells along with <a href="/wiki/Dopamine" title="Dopamine">dopamine</a>. Dopamine has functions in the light-adaptation process by up-regulating melanopsin transcription in ipRGCs and thus increasing the photoreceptor's sensitivity.<sup id="cite_ref-:1_3-3" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In parallel with the DA amacrine cell inhibition, somatostatin-releasing amacrine cells, themselves inhibited by DA amacrine cells, inhibit ipRGCs.<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> Other synaptic inputs to ipRGC dendrites include cone bipolar cells and rod bipolar cells.<sup id="cite_ref-:0_11-2" class="reference"><a href="#cite_note-:0-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Outputs">Outputs</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=7" title="Edit section: Outputs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>One postsynaptic target of ipRGCs is the suprachiasmatic nucleus (SCN) of the hypothalamus, which serves as the circadian clock in an organism. ipRGCs release both pituitary adenylyl cyclase-activating protein (PACAP) and <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamate</a> onto the SCN via a monosynaptic connection called the <a href="/wiki/Retinohypothalamic_tract" title="Retinohypothalamic tract">retinohypothalamic tract</a> (RHT).<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> Glutamate has an excitatory effect on SCN neurons, and PACAP appears to enhance the effects of glutamate in the hypothalamus.<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> </p><p>Other post synaptic targets of ipRGCs include: the intergenticulate leaflet (IGL), a cluster of neurons located in the thalamus, which play a role in circadian entrainment; the olivary pretectal nucleus (OPN), a cluster of neurons in the midbrain that controls the pupillary light reflex; the ventrolateral preoptic nucleus (VLPO), located in the hypothalamus and is a control center for sleep; <span class="clarify-content" style="padding-left:0.1em; padding-right:0.1em; color:var(--color-subtle, #54595d); border:1px solid var(--border-color-subtle, #c8ccd1);">as well as to</span><sup class="noprint Inline-Template Template-Clarify" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="This passage needs to be better explained. (March 2020)">clarify</span></a></i>]</sup> the amygdala.<sup id="cite_ref-:1_3-4" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<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=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=8" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Pupillary_light_reflex">Pupillary light reflex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=9" title="Edit section: Pupillary light reflex"><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:IpRGC_PLR.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/IpRGC_PLR.svg/220px-IpRGC_PLR.svg.png" decoding="async" width="220" height="291" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d3/IpRGC_PLR.svg/330px-IpRGC_PLR.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d3/IpRGC_PLR.svg/440px-IpRGC_PLR.svg.png 2x" data-file-width="195" data-file-height="258" /></a><figcaption>Inputs and outputs to ipRGCs involved in the pupillary light reflex</figcaption></figure> <p>Using various photoreceptor knockout mice, researchers have identified the role of ipRGCs in both the transient and sustained signaling of the <a href="/wiki/Pupillary_light_reflex" title="Pupillary light reflex">pupillary light reflex</a> (PLR).<sup id="cite_ref-:2_17-0" class="reference"><a href="#cite_note-:2-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Transient PLR occurs at dim to moderate light intensities and is a result of phototransduction occurring in <a href="/wiki/Rod_cell" title="Rod cell">rod cells</a>, which provide synaptic input onto ipRGCs, which in turn relay the information to the <a href="/wiki/Olivary_pretectal_nucleus" class="mw-redirect" title="Olivary pretectal nucleus">olivary pretectal nucleus</a> in the <a href="/wiki/Midbrain" title="Midbrain">midbrain</a>.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The neurotransmitter involved in the relay of information to the midbrain from the ipRGCs in the transient PLR is <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamate</a>. At brighter light intensities the sustained PLR occurs, which involves both phototransduction of the rod providing input to the ipRGCs and phototransduction of the ipRGCs themselves via melanopsin. Researchers have suggested that the role of melanopsin in the sustained PLR is due to its lack of adaptation to light stimuli in contrast to rod cells, which exhibit adaptation. The sustained PLR is maintained by <a href="/wiki/Pituitary_adenylate_cyclase-activating_peptide" title="Pituitary adenylate cyclase-activating peptide">PACAP</a> release from ipRGCs in a pulsatile manner.<sup id="cite_ref-:2_17-1" class="reference"><a href="#cite_note-:2-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Possible_role_in_conscious_sight">Possible role in conscious sight</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=10" title="Edit section: Possible role in conscious sight"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Experiments with rodless, coneless humans allowed another possible role for the receptor to be studied. In 2007, a new role was found for the photoreceptive ganglion cell. Zaidi and colleagues showed that in humans the retinal ganglion cell photoreceptor contributes to <a href="/wiki/Consciousness" title="Consciousness">conscious</a> sight as well as to non-image-forming functions like circadian rhythms, behaviour and pupillary reactions.<sup id="cite_ref-pmid18082405_9-1" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Since these cells respond mostly to <a href="/wiki/Blue" title="Blue">blue</a> light, it has been suggested that they have a role in <a href="/wiki/Mesopic_vision" title="Mesopic vision">mesopic vision</a><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. (November 2010)">citation needed</span></a></i>]</sup> and that the old theory of a purely <a href="/wiki/Duplex_retina" title="Duplex retina">duplex retina</a> with rod (dark) and cone (light) light vision was simplistic. Zaidi and colleagues' work with rodless, coneless human subjects hence has also opened the door into image-forming (visual) roles for the ganglion cell photoreceptor. </p><p>The discovery that there are parallel pathways for vision was made: one classic rod- and cone-based arising from the outer retina, the other a rudimentary visual brightness detector arising from the inner retina. The latter seems to be activated by light before the former.<sup id="cite_ref-pmid18082405_9-2" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Classic photoreceptors also feed into the novel photoreceptor system, and colour constancy may be an important role as suggested by Foster<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. (May 2018)">citation needed</span></a></i>]</sup>. </p><p>It has been suggested by the authors of the rodless, coneless human model that the receptor could be instrumental in understanding many diseases, including major causes of blindness worldwide such as <a href="/wiki/Glaucoma" title="Glaucoma">glaucoma</a>, a disease which affects ganglion cells. </p><p>In other mammals, photosensitive ganglia have proven to have a genuine role in conscious vision. Tests conducted by Jennifer Ecker et al. found that rats lacking rods and cones were able to learn to swim toward sequences of vertical bars rather than an equally luminescent gray screen.<sup id="cite_ref-Ecker_8-4" class="reference"><a href="#cite_note-Ecker-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Violet-to-blue_light">Violet-to-blue light</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=11" title="Edit section: Violet-to-blue light"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most work suggests that the peak spectral sensitivity of the receptor is between 460 and 484 nm. Lockley et al. in 2003<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> showed that 460 nm (blue) wavelengths of light suppress melatonin twice as much as 555 nm (green) light, the peak sensitivity of the photopic visual system. In work by Zaidi, Lockley and co-authors using a rodless, coneless human, it was found that a very intense 481 nm stimulus led to some conscious light perception, meaning that some rudimentary vision was realized.<sup id="cite_ref-pmid18082405_9-3" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Discovery">Discovery</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=12" title="Edit section: Discovery"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1923, <a href="/wiki/Clyde_E._Keeler" title="Clyde E. Keeler">Clyde E. Keeler</a> observed that the pupils in the eyes of blind mice he had accidentally bred still responded to light.<sup id="cite_ref-Keeler_2-1" class="reference"><a href="#cite_note-Keeler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The ability of the rodless, coneless mice to retain a <a href="/wiki/Pupillary_light_reflex" title="Pupillary light reflex">pupillary light reflex</a> was suggestive of an additional photoreceptor cell.<sup id="cite_ref-:0_11-3" class="reference"><a href="#cite_note-:0-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>In the 1980s, research in rod- and cone-deficient rats showed regulation of dopamine in the retina, a known neuromodulator for light adaptation and photoentrainment.<sup id="cite_ref-:1_3-5" class="reference"><a href="#cite_note-:1-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Research continued in 1991, when <a href="/wiki/Russell_G._Foster" class="mw-redirect" title="Russell G. Foster">Russell G. Foster</a> and colleagues, including <a href="/wiki/Ignacio_Provencio" title="Ignacio Provencio">Ignacio Provencio</a>, showed that rods and cones were not necessary for photoentrainment, the visual drive of the <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">circadian rhythm</a>, nor for the regulation of <a href="/wiki/Melatonin" title="Melatonin">melatonin</a> secretion from the <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland</a>, via rod- and cone-knockout mice.<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><sup id="cite_ref-:0_11-4" class="reference"><a href="#cite_note-:0-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Later work by Provencio and colleagues showed that this photoresponse was mediated by the photopigment <a href="/wiki/Melanopsin" title="Melanopsin">melanopsin</a>, present in the ganglion cell layer of the retina.<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>The photoreceptors were identified in 2002 by <a href="/wiki/Samer_Hattar" title="Samer Hattar">Samer Hattar</a>, David Berson and colleagues, where they were shown to be melanopsin expressing ganglion cells that possessed an intrinsic light response and projected to a number of brain areas involved in non-image-forming vision.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>In 2005, Panda, Melyan, Qiu, and colleagues demonstrated that the melanopsin photopigment was the phototransduction pigment in ganglion cells.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Dennis Dacey and colleagues showed in a species of Old World monkey that giant ganglion cells expressing melanopsin projected to the <a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">lateral geniculate nucleus</a> (LGN).<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><sup id="cite_ref-:3_6-1" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Previously only projections to the midbrain (pre-tectal nucleus) and hypothalamus (<a href="/wiki/Suprachiasmatic_nucleus" title="Suprachiasmatic nucleus">supra-chiasmatic nuclei</a>, SCN) had been shown. However, a visual role for the receptor was still unsuspected and unproven. </p> <div class="mw-heading mw-heading2"><h2 id="Research">Research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=13" title="Edit section: Research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Research_in_humans">Research in humans</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=14" title="Edit section: Research in humans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Attempts were made to hunt down the receptor in humans, but humans posed special challenges and demanded a new model. Unlike in other animals, researchers could not ethically induce rod and cone loss either genetically or with chemicals so as to directly study the ganglion cells. For many years, only inferences could be drawn about the receptor in humans, though these were at times pertinent. </p><p>In 2007, Zaidi and colleagues published their work on rodless, coneless humans, showing that these people retain normal responses to nonvisual effects of light.<sup id="cite_ref-pmid18082405_9-4" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Coghlan2007_27-0" class="reference"><a href="#cite_note-Coghlan2007-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The identity of the non-rod, non-cone photoreceptor in humans was found to be a ganglion cell in the inner retina as shown previously in rodless, coneless models in some other mammals. The work was done using patients with rare diseases that wiped out classic rod and cone photoreceptor function but preserved ganglion cell function.<sup id="cite_ref-pmid18082405_9-5" class="reference"><a href="#cite_note-pmid18082405-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Coghlan2007_27-1" class="reference"><a href="#cite_note-Coghlan2007-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Despite having no rods or cones, the patients continued to exhibit circadian photoentrainment, circadian behavioural patterns, melatonin suppression, and pupil reactions, with peak spectral sensitivities to environmental and experimental light that match the melanopsin photopigment. Their brains could also associate vision with light of this frequency. Clinicians and scientists are now seeking to understand the new receptor's role in human diseases and blindness.<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. (October 2022)">citation needed</span></a></i>]</sup> Intrinsically photosensitive RGCs have also been implicated in the exacerbation of headache by light during migraine attacks.<sup id="cite_ref-Noseda2010_28-0" class="reference"><a href="#cite_note-Noseda2010-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=15" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Bistratified_cell" title="Bistratified cell">Bistratified cell</a></li> <li><a href="/wiki/Melanopsin" title="Melanopsin">Melanopsin</a></li> <li><a href="/wiki/Midget_cell" title="Midget cell">Midget cell</a></li> <li><a href="/wiki/Parasol_cell" title="Parasol cell">Parasol cell</a></li> <li><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor</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=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=16" 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 reflist-columns references-column-width" style="column-width: 30em;"> <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="CITEREFKeeler1927" class="citation journal cs1">Keeler, Clyde E. (1927). "Iris movements in blind mice". <i>American Journal of Physiology</i>. <b>81</b> (1): 107–112. <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%2Fajplegacy.1927.81.1.107">10.1152/ajplegacy.1927.81.1.107</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Physiology&rft.atitle=Iris+movements+in+blind+mice&rft.volume=81&rft.issue=1&rft.pages=107-112&rft.date=1927&rft_id=info%3Adoi%2F10.1152%2Fajplegacy.1927.81.1.107&rft.aulast=Keeler&rft.aufirst=Clyde+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AIntrinsically+photosensitive+retinal+ganglion+cell" class="Z3988"></span></span> </li> <li id="cite_note-Keeler-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Keeler_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Keeler_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKeeler1928" class="citation journal cs1">Keeler CE (October 1928). "Blind Mice". <i>Journal of Experimental Zoology</i>. <b>51</b> (4): 495–508. <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/1928JEZ....51..495K">1928JEZ....51..495K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1002%2Fjez.1400510404">10.1002/jez.1400510404</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Experimental+Zoology&rft.atitle=Blind+Mice&rft.volume=51&rft.issue=4&rft.pages=495-508&rft.date=1928-10&rft_id=info%3Adoi%2F10.1002%2Fjez.1400510404&rft_id=info%3Abibcode%2F1928JEZ....51..495K&rft.aulast=Keeler&rft.aufirst=CE&rfr_id=info%3Asid%2Fen.wikipedia.org%3AIntrinsically+photosensitive+retinal+ganglion+cell" class="Z3988"></span></span> </li> <li 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=New+Scientist&rft.atitle=How+blind+people+see+sunrise+and+sunset&rft.volume=196&rft.issue=2635%E2%80%932636&rft.pages=9&rft.date=2007&rft_id=info%3Adoi%2F10.1016%2FS0262-4079%2807%2963172-8&rft.aulast=Coghlan&rft.aufirst=Andy&rft_id=https%3A%2F%2Fwww.newscientist.com%2Farticle%2Fmg19626354.100-blind-people-see-sunrise-and-sunset.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AIntrinsically+photosensitive+retinal+ganglion+cell" class="Z3988"></span></span> </li> <li id="cite_note-Noseda2010-28"><span class="mw-cite-backlink"><b><a href="#cite_ref-Noseda2010_28-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNoseda2010" class="citation journal cs1">Noseda R (2010). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2818758">"A neural mechanism for exacerbation of headache by light"</a>. <i>Nature Neuroscience</i>. <b>13</b> (2): 239–245. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnn.2475">10.1038/nn.2475</a>. <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/PMC2818758">2818758</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/20062053">20062053</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Neuroscience&rft.atitle=A+neural+mechanism+for+exacerbation+of+headache+by+light&rft.volume=13&rft.issue=2&rft.pages=239-245&rft.date=2010&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2818758%23id-name%3DPMC&rft_id=info%3Apmid%2F20062053&rft_id=info%3Adoi%2F10.1038%2Fnn.2475&rft.aulast=Noseda&rft.aufirst=Rodrigo&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2818758&rfr_id=info%3Asid%2Fen.wikipedia.org%3AIntrinsically+photosensitive+retinal+ganglion+cell" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Intrinsically_photosensitive_retinal_ganglion_cell&action=edit&section=17" title="Edit section: External links"><span>edit</span></a><span 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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 href="/wiki/Retina" title="Retina">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 href="/wiki/Retina" title="Retina">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="Nervous_tissue" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Nervous_tissue" title="Template:Nervous tissue"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Nervous_tissue" title="Template talk:Nervous tissue"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Nervous_tissue" title="Special:EditPage/Template:Nervous tissue"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Nervous_tissue" style="font-size:114%;margin:0 4em"><a href="/wiki/Nervous_tissue" title="Nervous tissue">Nervous tissue</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Central_nervous_system" title="Central nervous system">CNS</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Tissue Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></li> <li><a href="/wiki/White_matter" title="White matter">White matter</a> <ul><li><a href="/wiki/Projection_fiber" title="Projection fiber">Projection fibers</a></li> <li><a href="/wiki/Association_fiber" title="Association fiber">Association fiber</a></li> <li><a href="/wiki/Commissural_fiber" title="Commissural fiber">Commissural fiber</a></li> <li><a href="/wiki/Lemniscus_(anatomy)" title="Lemniscus (anatomy)">Lemniscus</a></li> <li><a href="/wiki/Nerve_tract" title="Nerve tract">Nerve tract</a></li> <li><a href="/wiki/Decussation" title="Decussation">Decussation</a></li></ul></li> <li><a href="/wiki/Neuropil" title="Neuropil">Neuropil</a></li> <li><a href="/wiki/Meninges" title="Meninges">Meninges</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cell Types</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/Neuron" title="Neuron">Neuronal</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/Pyramidal_cell" title="Pyramidal cell">Pyramidal</a></li> <li><a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje</a></li> <li><a href="/wiki/Granule_cell" title="Granule cell">Granule</a></li> <li><a href="/wiki/Von_Economo_neuron" title="Von Economo neuron">Von Economo</a></li> <li><a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">Medium spiny</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Glial</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/Astrocyte" title="Astrocyte">Astrocyte</a></li> <li><a href="/wiki/Ependyma" title="Ependyma">Ependymal cells</a> <ul><li><a href="/wiki/Tanycyte" title="Tanycyte">Tanycyte</a></li></ul></li> <li><a href="/wiki/Oligodendrocyte_progenitor_cell" title="Oligodendrocyte progenitor cell">Oligodendrocyte progenitor cell</a></li> <li><a href="/wiki/Oligodendrocyte" title="Oligodendrocyte">Oligodendrocyte</a></li> <li><a href="/wiki/Microglia" title="Microglia">Microglia</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Peripheral_nervous_system" title="Peripheral nervous system">PNS</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i>Dorsal</i> <ul><li><a href="/wiki/Dorsal_root_of_spinal_nerve" title="Dorsal root of spinal nerve">Root</a></li> <li><a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">Ganglion</a></li> <li><a href="/wiki/Dorsal_ramus_of_spinal_nerve" title="Dorsal ramus of spinal nerve">Ramus</a></li></ul></li> <li><i>Ventral</i> <ul><li><a href="/wiki/Ventral_root_of_spinal_nerve" title="Ventral root of spinal nerve">Root</a></li> <li><a href="/wiki/Ventral_ramus_of_spinal_nerve" title="Ventral ramus of spinal nerve">Ramus</a></li></ul></li> <li><i><a href="/wiki/Ramus_communicans" title="Ramus communicans">Ramus communicans</a></i> <ul><li><a href="/wiki/Gray_ramus_communicans" title="Gray ramus communicans">Gray</a></li> <li><a href="/wiki/White_ramus_communicans" title="White ramus communicans">White</a></li></ul></li> <li><a href="/wiki/Autonomic_ganglion" title="Autonomic ganglion">Autonomic ganglion</a> (<a href="/wiki/Preganglionic_nerve_fibers" title="Preganglionic nerve fibers">Preganglionic nerve fibers</a></li> <li><a href="/wiki/Postganglionic_nerve_fibers" title="Postganglionic nerve fibers">Postganglionic nerve fibers</a>)</li> <li><a href="/wiki/Nerve_fascicle" title="Nerve fascicle">Nerve fascicle</a></li> <li><a href="/wiki/Funiculus_(neuroanatomy)" title="Funiculus (neuroanatomy)">Funiculus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissues</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/Epineurium" title="Epineurium">Epineurium</a></li> <li><a href="/wiki/Perineurium" title="Perineurium">Perineurium</a></li> <li><a href="/wiki/Endoneurium" title="Endoneurium">Endoneurium</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Neuroglia</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/Myelin" title="Myelin">Myelination</a>: <a href="/wiki/Schwann_cell" title="Schwann cell">Schwann cell</a> <ul><li><a href="/wiki/Neurilemma" title="Neurilemma">Neurilemma</a></li> <li><a href="/wiki/Myelin_incisure" title="Myelin incisure">Myelin incisure</a></li> <li><a href="/wiki/Node_of_Ranvier" title="Node of Ranvier">Node of Ranvier</a></li> <li><a href="/wiki/Internodal_segment" title="Internodal segment">Internodal segment</a></li></ul></li> <li><a href="/wiki/Satellite_glial_cell" title="Satellite glial cell">Satellite glial cell</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neuron" title="Neuron">Neurons</a>/<br /><a href="/wiki/Nerve_fiber" class="mw-redirect" title="Nerve fiber">nerve fibers</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Parts</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/Soma_(biology)" title="Soma (biology)">Soma</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/Axon_hillock" title="Axon hillock">Axon hillock</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Axon" title="Axon">Axon</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/Telodendron" class="mw-redirect" title="Telodendron">Telodendron</a></li> <li><a href="/wiki/Axon_terminal" title="Axon terminal">Axon terminals</a></li> <li><a href="/wiki/Axoplasm" title="Axoplasm">Axoplasm</a></li> <li><a href="/wiki/Axolemma" title="Axolemma">Axolemma</a></li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofibril/neurofilament</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dendrite" title="Dendrite">Dendrite</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><ul><li><a href="/wiki/Nissl_body" title="Nissl body">Nissl body</a></li> <li><a href="/wiki/Dendritic_spine" title="Dendritic spine">Dendritic spine</a></li> <li><a href="/wiki/Apical_dendrite" title="Apical dendrite">Apical dendrite</a>/<a href="/wiki/Basal_dendrite" title="Basal dendrite">Basal dendrite</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</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/Bipolar_neuron" title="Bipolar neuron">Bipolar</a></li> <li><a href="/wiki/Unipolar_neuron" title="Unipolar neuron">Unipolar</a></li> <li><a href="/wiki/Pseudounipolar_neuron" title="Pseudounipolar neuron">Pseudounipolar</a></li> <li><a href="/wiki/Multipolar_neuron" title="Multipolar neuron">Multipolar</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a> <ul><li><a href="/wiki/Renshaw_cell" title="Renshaw cell">Renshaw</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">Afferent nerve fiber</a>/<br /><a href="/wiki/Sensory_neuron" title="Sensory neuron">Sensory neuron</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/General_somatic_afferent_fibers" class="mw-redirect" title="General somatic afferent fibers">GSA</a></li> <li><a href="/wiki/General_visceral_afferent_fibers" class="mw-redirect" title="General visceral afferent fibers">GVA</a></li> <li><a href="/wiki/Special_somatic_afferent" class="mw-redirect" title="Special somatic afferent">SSA</a></li> <li><a href="/wiki/Special_visceral_afferent" class="mw-redirect" title="Special visceral afferent">SVA</a></li> <li><a href="/wiki/Axon#types" title="Axon">fibers</a> <ul><li><a href="/wiki/Type_Ia_sensory_fiber" title="Type Ia sensory fiber">Ia or Aα</a></li> <li><a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Ib or Golgi or Aα</a></li> <li><a href="/wiki/Type_II_sensory_fiber" title="Type II sensory fiber">II or Aβ and Aγ</a></li> <li><a href="/wiki/Group_A_nerve_fiber" title="Group A nerve fiber">III or Aδ or fast pain</a></li> <li><a href="/wiki/Group_C_nerve_fiber" title="Group C nerve fiber">IV or C or slow pain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">Efferent nerve fiber</a>/<br /><a href="/wiki/Motor_neuron" title="Motor neuron">Motor neuron</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/General_somatic_efferent_fibers" class="mw-redirect" title="General somatic efferent fibers">GSE</a></li> <li><a href="/wiki/General_visceral_efferent_fibers" class="mw-redirect" title="General visceral efferent fibers">GVE</a></li> <li><a href="/wiki/Special_visceral_efferent" class="mw-redirect" title="Special visceral efferent">SVE</a></li> <li><a href="/wiki/Upper_motor_neuron" title="Upper motor neuron">Upper motor neuron</a></li> <li><a href="/wiki/Lower_motor_neuron" title="Lower motor neuron">Lower motor neuron</a> <ul><li><a href="/wiki/Alpha_motor_neuron" title="Alpha motor neuron">α motorneuron</a></li> <li><a href="/wiki/Beta_motor_neuron" title="Beta motor neuron">β motorneuron</a></li> <li><a href="/wiki/Gamma_motor_neuron" title="Gamma motor neuron">γ motorneuron</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Termination</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Synapse" title="Synapse">Synapse</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/Electrical_synapse" title="Electrical synapse">Electrical synapse</a>/<a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a> <ul><li><a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">Synaptic vesicle</a></li> <li><a href="/wiki/Active_zone" title="Active zone">Active zone</a></li> <li><a href="/wiki/Postsynaptic_density" title="Postsynaptic density">Postsynaptic density</a></li></ul></li> <li><a href="/wiki/Autapse" title="Autapse">Autapse</a></li> <li><a href="/wiki/Ribbon_synapse" title="Ribbon synapse">Ribbon synapse</a></li> <li><a href="/wiki/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sensory_receptor" class="mw-redirect" title="Sensory receptor">Sensory receptors</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/Tactile_corpuscle" title="Tactile corpuscle">Meissner's corpuscle</a></li> <li><a href="/wiki/Merkel_nerve_ending" title="Merkel nerve ending">Merkel nerve ending</a></li> <li><a href="/wiki/Pacinian_corpuscle" title="Pacinian corpuscle">Pacinian corpuscle</a></li> <li><a href="/wiki/Bulbous_corpuscle" title="Bulbous corpuscle">Ruffini ending</a></li> <li><a href="/wiki/Muscle_spindle" title="Muscle spindle">Muscle spindle</a></li> <li><a href="/wiki/Free_nerve_ending" title="Free nerve ending">Free nerve ending</a></li> <li><a href="/wiki/Nociceptor" title="Nociceptor">Nociceptor</a></li> <li><a href="/wiki/Olfactory_receptor_neuron" title="Olfactory receptor neuron">Olfactory receptor neuron</a></li> <li><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor cell</a></li> <li><a href="/wiki/Hair_cell" title="Hair cell">Hair cell</a></li> <li><a href="/wiki/Taste_receptor" title="Taste receptor">Taste receptor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" 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