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Subcommissural organ - Wikipedia

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<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-Reissner&#039;s_fiber" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reissner&#039;s_fiber"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Reissner's fiber</span> </div> </a> <ul id="toc-Reissner&#039;s_fiber-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-SCO-spondin,_a_glycoprotein_of_the_SCO/RF_complex" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#SCO-spondin,_a_glycoprotein_of_the_SCO/RF_complex"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>SCO-spondin, a glycoprotein of the SCO/RF complex</span> </div> </a> <button aria-controls="toc-SCO-spondin,_a_glycoprotein_of_the_SCO/RF_complex-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 SCO-spondin, a glycoprotein of the SCO/RF complex subsection</span> </button> <ul id="toc-SCO-spondin,_a_glycoprotein_of_the_SCO/RF_complex-sublist" class="vector-toc-list"> <li id="toc-Multidomain_organization" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Multidomain_organization"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Multidomain organization</span> </div> </a> <ul id="toc-Multidomain_organization-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Development" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Development"> <div 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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:lemonchiffon">Subcommissural organ</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Gray715.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Gray715.png/250px-Gray715.png" decoding="async" width="250" height="185" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Gray715.png/375px-Gray715.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Gray715.png/500px-Gray715.png 2x" data-file-width="600" data-file-height="443" /></a></span><div class="infobox-caption">Right midline aspect of a human brain sectioned in the median <a href="/wiki/Sagittal_plane" title="Sagittal plane">sagittal plane</a>. The subcommissural organ is not labeled, but the region is visible near the <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland</a>.</div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></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>organum subcommissurale</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=D013351">D013351</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/NeuroNames" title="NeuroNames">NeuroNames</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://braininfo.rprc.washington.edu/centraldirectory.aspx?ID=483">483</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/NeuroLex" title="NeuroLex">NeuroLex</a> ID</th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://uri.neuinfo.org/nif/nifstd/birnlex_1028">birnlex_1028</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/14.1.08.511%20Entity%20TA98%20EN.htm">A14.1.08.511</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=5783">5783</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&amp;conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma72414">72414</a></td></tr><tr><td colspan="2" class="infobox-below"><a href="/wiki/Anatomical_terms_of_neuroanatomy" title="Anatomical terms of neuroanatomy">Anatomical terms of neuroanatomy</a><div style="text-align: right;"><small class="noprint">&#91;<a href="https://www.wikidata.org/wiki/Q1423193" class="extiw" title="d:Q1423193">edit on Wikidata</a>]</small></div></td></tr></tbody></table><p> The <b>subcommissural organ</b> (SCO) is one of the <a href="/wiki/Circumventricular_organs" title="Circumventricular organs">circumventricular organs</a> of the <a href="/wiki/Brain" title="Brain">brain</a>. It is a small glandular structure<sup id="cite_ref-Rod1998_1-0" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> that is located in the posterior region of the <a href="/wiki/Third_ventricle" title="Third ventricle">third ventricle</a>, near the entrance of the <a href="/wiki/Cerebral_aqueduct" title="Cerebral aqueduct">cerebral aqueduct</a>. </p><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Subcommissural_organ_of_the_mouse_brain.jpg" class="mw-file-description"><img alt="The subcommissural organ of a mouse." src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Subcommissural_organ_of_the_mouse_brain.jpg/220px-Subcommissural_organ_of_the_mouse_brain.jpg" decoding="async" width="220" height="192" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Subcommissural_organ_of_the_mouse_brain.jpg/330px-Subcommissural_organ_of_the_mouse_brain.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4c/Subcommissural_organ_of_the_mouse_brain.jpg/440px-Subcommissural_organ_of_the_mouse_brain.jpg 2x" data-file-width="1500" data-file-height="1309" /></a><figcaption>A <a href="/wiki/Photomicrograph" class="mw-redirect" title="Photomicrograph">photomicrograph</a> of the subcommissural organ in a mouse brain (view from the front). The lightly stained columns projecting toward the center are the apical extensions of the <a href="/wiki/Ependymal_cells" class="mw-redirect" title="Ependymal cells">ependymal cells</a>. The posterior commissure is at the top of the photo, and the space below the SCO is part of the third ventricle, which is lined with ependymal cells. <a href="/wiki/Cell_nuclei" class="mw-redirect" title="Cell nuclei">Cell nuclei</a> are stained blue. The scale bar = 50 microns (0.05 mm).</figcaption></figure><p> The name of the SCO comes from its location beneath the <a href="/wiki/Posterior_commissure" title="Posterior commissure">posterior commissure</a>, a bundle of nerve fibers interconnecting parts of the two hemispheres of the brain. The SCO is one of the first differentiated brain structures to develop.<sup id="cite_ref-Rod1998_1-1" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> Although it is evolutionarily an ancient structure that is present throughout the <a href="/wiki/Chordate" title="Chordate">chordate</a> phylum, its arrangement varies somewhat among species.<sup id="cite_ref-Rod1998_1-2" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> Functions of the SCO are unknown; some evidence indicates it may participate in <a href="/wiki/Clearance_(pharmacology)" title="Clearance (pharmacology)">clearance</a> of certain compounds from the <a href="/wiki/Cerebrospinal_fluid" title="Cerebrospinal fluid">cerebrospinal fluid</a>, and possibly in <a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenetic</a> mechanisms, such as development of the <a href="/wiki/Posterior_commissure" title="Posterior commissure">posterior commissure</a>.<sup id="cite_ref-Rod1998_1-3" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-0" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</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=Subcommissural_organ&amp;action=edit&amp;section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Cells of the subcommissural organ, which are specialized in the secretion of <a href="/wiki/Glycoprotein" title="Glycoprotein">glycoproteins</a> (see below), are arranged into two layers: a superficial layer called the <a href="/wiki/Ependyma" title="Ependyma">ependyma</a> and an underlying layer called the hypendyma.<sup id="cite_ref-Rod1998_1-4" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> The ependyma consists of long, columnar cells that release their secretions into the ventricular cerebrospinal fluid. The hypendyma (which is most prominent in mammals) is characterized by numerous capillaries and glial cells. Hypendymal cells and ependymal cells both are secretory in nature; their processes project to local blood vessels and also to the subarachnoidal space.<sup id="cite_ref-Rod1998_1-5" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-1" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> </p><p>The ependymal cell bodies present a clear zonation that is especially marked in certain species: 1) In the perinuclear region, large and dilated cisternae of the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">rough endoplasmic reticulum (RER)</a> are the most distinctive ultrastructural feature of virtually all species; 2) the intermediate region consists mainly of RER and <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a>; 3) the subapical region is relatively narrow, and includes microtubules, mitochondria and smooth endoplasmic reticulum; and 4) the apical region comprises a large protrusion into the ventricle.<sup id="cite_ref-Rod1998_1-6" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-2" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> There are no neuronal cell bodies within the subcommissural organ, although the specialized cells receive some innervation from outside neurons.<sup id="cite_ref-Pax1995_3-0" class="reference"><a href="#cite_note-Pax1995-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</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=Subcommissural_organ&amp;action=edit&amp;section=2" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ependymal cells secrete high molecular mass glycoproteins into the cerebrospinal fluid, in which the bulk of them condense to form a filamentous structure named Reissner's fiber.<sup id="cite_ref-Ortloff2013_4-0" class="reference"><a href="#cite_note-Ortloff2013-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> The subcommissural organ/<a href="/wiki/Reissner%27s_fiber" title="Reissner&#39;s fiber">Reissner's fiber</a> complex is thought to be involved in the reabsorption and circulation of the cerebrospinal fluid, and with functions related to electrolyte and <a href="/wiki/Osmoregulation" title="Osmoregulation">water balance</a>.<sup id="cite_ref-Ortloff2013_4-1" class="reference"><a href="#cite_note-Ortloff2013-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-PerezFigares2001_5-0" class="reference"><a href="#cite_note-PerezFigares2001-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> </p><p>One of the proteins secreted by the subcommissural organ, and which is present in Reissner's fiber, is spondin. SCO-spondin is a "giant" (5000 amino acids) glycoprotein (<a href="/wiki/Thrombospondin" title="Thrombospondin">thrombospondin</a> <a href="/wiki/Protein_superfamily" title="Protein superfamily">superfamily</a>) found in <a href="/wiki/Vertebrate" title="Vertebrate">Vertebrata</a>. This glycoprotein shares molecular domains with <a href="/wiki/Axon_guidance" title="Axon guidance">axonal pathfinding</a> molecules.<sup id="cite_ref-PerezFigares2001_5-1" class="reference"><a href="#cite_note-PerezFigares2001-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> The <a href="/wiki/Ependyma" title="Ependyma">ependymal cells</a> and the <a href="/wiki/SSPO" class="mw-redirect" title="SSPO">SCO–spondin</a> secretion are thought to play a role in <a href="/wiki/Homeostasis" title="Homeostasis">homeostasis</a>.<sup id="cite_ref-pmid17150405_6-0" class="reference"><a href="#cite_note-pmid17150405-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> </p><p>The ependymal cells of the SCO are also involved in the production of brain <a href="/wiki/Transthyretin" title="Transthyretin">transthyretin</a>, a protein involved in the transport of thyroid hormones in blood.<sup id="cite_ref-Mon2005_7-0" class="reference"><a href="#cite_note-Mon2005-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> </p><p>Some studies indicate the presence of both tyrosine-hydroxylase-immunoreactive <a href="/wiki/Axon" title="Axon">nerve fibers</a> and <a href="/wiki/Dopamine" title="Dopamine">dopamine</a> receptors in the SCO ependyma.<sup id="cite_ref-pmid15197646_8-0" class="reference"><a href="#cite_note-pmid15197646-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> In addition, there is evidence suggesting that the SCO activity in adult animals may be regulated by <a href="/wiki/Serotonin" title="Serotonin">serotonin</a>.<sup id="cite_ref-pmid15469891_9-0" class="reference"><a href="#cite_note-pmid15469891-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>All capillaries in the <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> with a functional blood-brain barrier express glucose transporters (<a href="/wiki/GLUT1" title="GLUT1">GLUT1</a>). These transporters are generally absent in leaky barrier structures. The circumventricular organs that are known to have leaky barrier capillaries were stained by fibronectin antibodies but not by GLUT1 antibodies. The subcommissural organ appears to be unique in that it shows neither GLUT1 nor capillary.<sup id="cite_ref-pmid7790418_10-0" class="reference"><a href="#cite_note-pmid7790418-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Reissner's_fiber"><span id="Reissner.27s_fiber"></span>Reissner's fiber</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=3" title="Edit section: Reissner&#039;s fiber"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Reissner%27s_fiber" title="Reissner&#39;s fiber">Reissner's fiber</a></div> <p>Reissner's fiber is also thought to be important in morphogenetic neuronal processes, being involved in neuronal survival, aggregation and neurite extension. In vitro studies demonstrated that the presence of RF, in conjunction with glial cells, is essential to the survival of neuronal cells. The studies seem to point that the RF might bind some of the growth factors produced by glial cells and transport them to the neurons. On the process of neuronal aggregation, RF seems to serve as a control factor in direct cell-to-cell communication, favoring neuronal aggregation when the density of neurons is low and preventing this aggregation when the density gets higher. Although the mechanism behind this is not well understood, it is known to be linked to the different domains in SCO-spondin that are related to coagulation factors and TSRs, as referred above. Furthermore, the RF as a part on the neurite extension, promoting neurite outgrowth from both spinal and cortical neurons, in cell cultures, which may also be connected to the TSR domains of SCO-spondin.<sup id="cite_ref-pmid11241859_11-0" class="reference"><a href="#cite_note-pmid11241859-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="SCO-spondin,_a_glycoprotein_of_the_SCO/RF_complex"><span id="SCO-spondin.2C_a_glycoprotein_of_the_SCO.2FRF_complex"></span>SCO-spondin, a glycoprotein of the SCO/RF complex</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=4" title="Edit section: SCO-spondin, a glycoprotein of the SCO/RF complex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The primary structure of the major constituent of bovine RF, SCO-spondin, has been fully established as a large N-glycosylated protein (450 kDa).<sup id="cite_ref-pmid11008217_12-0" class="reference"><a href="#cite_note-pmid11008217-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pmid11241859_11-1" class="reference"><a href="#cite_note-pmid11241859-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Many lines of evidence denote that SCO-spondin plays a role in CNS development.<sup id="cite_ref-pmid18218138_13-0" class="reference"><a href="#cite_note-pmid18218138-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> This molecule belongs to a protein superfamily exhibiting conserved motifs of the thrombospondin type 1 repeat. Proteins of this family are strongly expressed during mammalian CNS development, being involved in mechanisms of cellular adhesion and axonal pathfinding (a process by which neurons send out axons to reach the correct targets during neural development).<sup id="cite_ref-pmid18218138_13-1" class="reference"><a href="#cite_note-pmid18218138-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>Numerous investigations have been directed towards the identification and characterization of the secretory compounds of the SCO, clarifying partially its function. Immunoblot analyses of bovine SCO using antibodies against RF glycoproteins allowed the identification of high molecular weight glycoproteins of 540, 450, 320 and 190 kDa.<sup id="cite_ref-pmid18218138_13-2" class="reference"><a href="#cite_note-pmid18218138-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The 540 and the 320 kDa compounds would correspond to precursor forms.<sup id="cite_ref-pmid1661820_14-0" class="reference"><a href="#cite_note-pmid1661820-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Multidomain_organization">Multidomain organization</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=5" title="Edit section: Multidomain organization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The main SCO-spondin isoform consists of multiple domains. This multidomain organization is a special feature of the Chordate Phylum, and there is a high degree of conservation in the amino acids composition in mammals.<sup id="cite_ref-pmid17126404_15-0" class="reference"><a href="#cite_note-pmid17126404-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> The complete sequence and modular organization of SCO-spondin was first characterized in Bos taurus.<sup id="cite_ref-pmid11008217_12-1" class="reference"><a href="#cite_note-pmid11008217-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> The structure of this protein is unique as it presents a mosaic arrangement of these domains along the backbone. </p><p>The putative function of SCO-spondin in neuronal differentiation is discussed regarding these features and homologies with other developmental molecules of the central nervous system exhibiting TSR domains, and involved in axonal guidance.<sup id="cite_ref-pmid11241859_11-2" class="reference"><a href="#cite_note-pmid11241859-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Peptides corresponding to SCO-spondin TSR domains strongly increased adhesivity and neuritic outgrowth of cortical neurons and induced disaggregation of spinal cord neurons. Therefore, it is a candidate to interfere with neuronal development and/or axonal guidance during ontogenesis of the central nervous system in the modulation of side-to-side and side-to-substratum interactions, and also in promoting neurite outgrowth.<sup id="cite_ref-pmid11241859_11-3" class="reference"><a href="#cite_note-pmid11241859-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p>The identification of conserved domains including Emilin (EMI), von Willebrand factor D (vWD) low-density lipoprotein receptor type A repeats (LDLrA) domains, SCO repeats (SCORs), 26 thrombospondin type 1 repeats (TSRs), a coagulation factor 5/8 type C (FA5-8C) or discoidin motif and a C-terminal cystin knot (CTCK) domain provides a wider insight into the putative function of this protein. Similar types of arrangement was encountered in zonadhesins and immunoglobulin G (IgG) FC binding fragment which may account for SCO-spondin functional aspect on promoting cell-to-substratum adhesivity.<sup id="cite_ref-pmid17126404_15-1" class="reference"><a href="#cite_note-pmid17126404-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> </p><p>The presence of low-density lipoprotein receptor type A (LDLrA) domains repeated ten times in the consensus sequence could provide a hint as to the function of SCORs, since LDLrA are known to interact with proteases or protease inhibitors.<sup id="cite_ref-pmid11301018_16-0" class="reference"><a href="#cite_note-pmid11301018-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> There may be a functional link between LDLrAs and SCORs, which could both be involved in the regulation of either protease activation or protease inhibition.<sup id="cite_ref-pmid17126404_15-2" class="reference"><a href="#cite_note-pmid17126404-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> The motifs coagulation factor 5/8 type C or discoidin and thrombospondin type 1 repeat (TSR) present in SCO-spondin consensus were initially described in blood proteins, where they were shown to play a role in coagulation or platelet aggregation. SCO-spondin and F-spondin share a similar pattern of expression in the floor plate, flexural organ and subcommissural organ and could have a redundant activity. The biological function of F-spondin and SCO-spondin on the deflection of commissural axons in the neural tube was assessed respectively by experiments of gain and loss of function<sup id="cite_ref-pmid10399931_17-0" class="reference"><a href="#cite_note-pmid10399931-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> and by analyses of mutants with defective floor plate. F-spondin and SCO-spondin were both shown to promote neurite outgrowth of various neuronal cell populations, in cell culture.<sup id="cite_ref-pmid14692680_18-0" class="reference"><a href="#cite_note-pmid14692680-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> </p><p>SCO-spondin may interfere with several biological events during early ontogenetical development of the CNS. Nevertheless, SCO-spondin is also present during the adult life, and similarly to thrombospondins, which act on various biological systems, i.e., neuronal differentiation, angiogenesis and platelet aggregation.<sup id="cite_ref-The_Thrombospondin_Gene_Family_19-0" class="reference"><a href="#cite_note-The_Thrombospondin_Gene_Family-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Development">Development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=6" title="Edit section: Development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="SCO">SCO</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=7" title="Edit section: SCO"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Despite being a much conserved structure throughout evolution, there are some differences on the SCO from different mammals. It is the first secretory structure to differentiate and remains fully developed and functional during the life of almost every vertebrate, excluding bats, anthropoid apes and humans. More specifically, in humans, the SCO development has a regressive nature. It reaches its apex development in fetus from 3 to 5 month old, functioning as a fully active secretory structure of the brain during this time span, and extending from the pineal recess over the posterior commissure to the mesocoelic recess. It is composed by a characteristic high columnar epithelium, which is not found in the adult SCO. Following this maxed developed state, the SCO starts regressing and in children from 3 to 4 years old it already has a vestigial character, being reduced to islet like structures on the adult. Although the remaining cells can possess some secretory material the SCO is truly vestigial in both structure and secretory function, in adults.<sup id="cite_ref-pmid11241867_20-0" class="reference"><a href="#cite_note-pmid11241867-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="SCO-spondin">SCO-spondin</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=8" title="Edit section: SCO-spondin"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>As part of the embryonic cerebrospinal fluid (eCSF), SCO-spondin is of the uttermost importance in the development of the neuronal system, being a key protein in the balancing of differentiation and proliferation of the neuroepithelium. It starts being secreted by the diencephalic floor plate in the first embryonic stages playing an important part in the development and differentiation of structures such as the pineal gland.<sup id="cite_ref-pmid_23761733_21-0" class="reference"><a href="#cite_note-pmid_23761733-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> In particular, the SCO-spondin appears to have a major role on the growth of the posterior commissure (PC), which was proved when mutants lacking SCO, and hence having no SCO-spondin, were unable to form a functional PC. On early stages of development the axonal growth is stimulated, being inhibited afterwards.<sup id="cite_ref-pmid20012322_22-0" class="reference"><a href="#cite_note-pmid20012322-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> A steep gradient of spondin expression in the neuroepithelium signals the need for different processes to take place, favoring the fasciculation on the cephalic region and the incorporation of new neurons on the caudal region. As such, the lower concentrations of SCO-spondin in the caudal region favor the axonal outgrowth and incorporation of new axons on the posterior commissure and the higher concentrations in the cephalic region promotes the interactions between the neighboring axons.<sup id="cite_ref-pmid_23761733_21-1" class="reference"><a href="#cite_note-pmid_23761733-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> In conjugation with the secretion of SCO-spondin, the midline positioning of the SCO assumes a great importance on the axon guidance process. This positioning facilitates the signaling of the turning points for the axons, through the spreading of spondin.<sup id="cite_ref-pmid20012322_22-1" class="reference"><a href="#cite_note-pmid20012322-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> In addition to the functions in axon guidance and related growth of the posterior commissure, the SCO-spondin also appears to have a role on the adhesion of the trophoblast to the uterine walls. There is a slightly different SCO-spondin produced in the trophoblast, most likely due to alternative splicing. This spondin may recognize the classic protein on the uterine wall, facilitating the adhesion.<sup id="cite_ref-pmid15053980_23-0" class="reference"><a href="#cite_note-pmid15053980-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinical_significance">Clinical significance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=9" title="Edit section: Clinical significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Hydrocephalus">Hydrocephalus</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=10" title="Edit section: Hydrocephalus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Given that the subcommissural organ is not highly permeable and does not possess <a href="/wiki/Fenestra_(histology)" class="mw-redirect" title="Fenestra (histology)">fenestrated</a> capillaries like other subventricular organs, it has emerged as a major site of congenital <a href="/wiki/Hydrocephalus" title="Hydrocephalus">hydrocephalus</a>.<sup id="cite_ref-pmid19364914_24-0" class="reference"><a href="#cite_note-pmid19364914-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-patho_25-0" class="reference"><a href="#cite_note-patho-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> It is suggested that this is related to immunological blockage of SCO secretions and Sylvian's aqueduct malformation<sup id="cite_ref-vitamin_26-0" class="reference"><a href="#cite_note-vitamin-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> and obliteration or turbulent cerebrospinal fluid flow due to the absence of Reissner's fibers.<sup id="cite_ref-pmid18218138_13-3" class="reference"><a href="#cite_note-pmid18218138-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> There is evidence that in transgenic mice the overexpression of <a href="/wiki/Sox3" class="mw-redirect" title="Sox3">Sox3</a> in the dorsal midline of the diencephalon in a dose-dependent manner<sup id="cite_ref-sox3_27-0" class="reference"><a href="#cite_note-sox3-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> and that the conditional inactivation of <a href="/wiki/Presenilin" title="Presenilin">presenilin-1</a><sup id="cite_ref-pmid21262207_28-0" class="reference"><a href="#cite_note-pmid21262207-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> or the lack of <a href="/wiki/Huntingtin" title="Huntingtin">huntingtin</a><sup id="cite_ref-pmid_3298867_29-0" class="reference"><a href="#cite_note-pmid_3298867-29"><span class="cite-bracket">&#91;</span>29<span class="cite-bracket">&#93;</span></a></sup> in wnt cell lineages leads to congenital hydrocephalus, which highlights the role of these proteins mediating the relation between the SCO and the condition (see also: <a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt signaling pathway</a> for more information). A more recent study using HTx rats reinforced the idea that the abnormal and dysfunction of the SCO precedes the development of the hydrocephalus.<sup id="cite_ref-Ortloff2013_4-2" class="reference"><a href="#cite_note-Ortloff2013-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Other_disease">Other disease</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=11" title="Edit section: Other disease"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>It is reported that in spontaneously hypertensive rats there is a relation between SCO and hypertension due to changing in its secretor activity and protein composition.<sup id="cite_ref-pmid_9792610_30-0" class="reference"><a href="#cite_note-pmid_9792610-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pmid16329042_31-0" class="reference"><a href="#cite_note-pmid16329042-31"><span class="cite-bracket">&#91;</span>31<span class="cite-bracket">&#93;</span></a></sup> </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=Subcommissural_organ&amp;action=edit&amp;section=12" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1860, <a href="/wiki/Ernst_Reissner" title="Ernst Reissner">Ernst Reissner</a>, anatomist at the University of Dorpat, published a monograph on the microscopic structure of the spinal cord of <i>Petromyzon fluviatilis</i>. He described a string of 1.5 μm in diameter characterized by its high refringence, its extremely regular shape, and its lying free within the central canal. In 1866, Karl Kutschin confirmed Reissner's observations and named the fibrous structure Reissner's fiber.<sup id="cite_ref-Rod1998_1-7" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-3" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> </p><p>Edinger (1892) described, in sharks, what later was known as "the subcommissural organ". Studnicka (1900), called attention to uncommonly tall ependymal cells covering the posterior comissure of <i>P. fluviatilis</i>. Sargent, also in 1900, establishes the basis of what is presently regarded as the subcomissural organ – Reissner fiber complex. Finally, in 1910, Dendy and Nicholls introduce the term "subcommissural organ" to describe this brain gland.<sup id="cite_ref-Rod1998_1-8" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-4" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> </p><p>The rich vascularization of the SCO was first reported by Pesonen (1940). In 1958, Helmut Hofer postulated that this organ, despite its structural and functional differences from other circumventricular organs, is a highly secretory component of the circumventricular organ system.<sup id="cite_ref-Morphologie_32-0" class="reference"><a href="#cite_note-Morphologie-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Rod1998_1-9" class="reference"><a href="#cite_note-Rod1998-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Oks1993_2-5" class="reference"><a href="#cite_note-Oks1993-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subcommissural_organ&amp;action=edit&amp;section=13" 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 reflist-columns-2"> <ol class="references"> <li id="cite_note-Rod1998-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Rod1998_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Rod1998_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Rod1998_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Rod1998_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Rod1998_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Rod1998_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Rod1998_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Rod1998_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Rod1998_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Rod1998_1-9"><sup><i><b>j</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFRodríguezRodríguezHein1998" class="citation journal cs1">Rodríguez EM, Rodríguez S, Hein S (April 1998). "The subcommissural organ". <i>Microscopy Research and Technique</i>. <b>41</b> (2): 98–123. <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%2F%28SICI%291097-0029%2819980415%2941%3A2%3C98%3A%3AAID-JEMT2%3E3.0.CO%3B2-M">10.1002/(SICI)1097-0029(19980415)41:2&#60;98::AID-JEMT2&#62;3.0.CO&#59;2-M</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9579598">9579598</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&#160;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:358861">358861</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Microscopy+Research+and+Technique&amp;rft.atitle=The+subcommissural+organ&amp;rft.volume=41&amp;rft.issue=2&amp;rft.pages=98-123&amp;rft.date=1998-04&amp;rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A358861%23id-name%3DS2CID&amp;rft_id=info%3Apmid%2F9579598&amp;rft_id=info%3Adoi%2F10.1002%2F%28SICI%291097-0029%2819980415%2941%3A2%3C98%3A%3AAID-JEMT2%3E3.0.CO%3B2-M&amp;rft.aulast=Rodr%C3%ADguez&amp;rft.aufirst=EM&amp;rft.au=Rodr%C3%ADguez%2C+S&amp;rft.au=Hein%2C+S&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ASubcommissural+organ" class="Z3988"></span></span> </li> <li id="cite_note-Oks1993-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Oks1993_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Oks1993_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Oks1993_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Oks1993_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Oks1993_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Oks1993_2-5"><sup><i><b>f</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOkscheRodríguezLlebrez1993" class="citation book cs1">Oksche A, Rodríguez EM, Llebrez PF (1993). Oksche A, Rodríguez EM, Fernández-Llebrez P (eds.). <i>The Subcommissural Organ: An Ependymal Brain Gland</i>. Berlin: Springer Verlag. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-3-642-78013-4">10.1007/978-3-642-78013-4</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/978-3-540-56336-5" title="Special:BookSources/978-3-540-56336-5"><bdi>978-3-540-56336-5</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a>&#160;<a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/27681500">27681500</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&#160;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:36028700">36028700</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=The+Subcommissural+Organ%3A+An+Ependymal+Brain+Gland&amp;rft.place=Berlin&amp;rft.pub=Springer+Verlag&amp;rft.date=1993&amp;rft_id=info%3Aoclcnum%2F27681500&amp;rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A36028700%23id-name%3DS2CID&amp;rft_id=info%3Adoi%2F10.1007%2F978-3-642-78013-4&amp;rft.isbn=978-3-540-56336-5&amp;rft.aulast=Oksche&amp;rft.aufirst=A&amp;rft.au=Rodr%C3%ADguez%2C+EM&amp;rft.au=Llebrez%2C+PF&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3ASubcommissural+organ" class="Z3988"></span><sup class="noprint Inline-Template" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (March 2014)">page&#160;needed</span></a></i>&#93;</sup></span> </li> <li id="cite_note-Pax1995-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Pax1995_3-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOldfieldMckinley1995" class="citation book cs1">Oldfield BJ, Mckinley MJ (1995). Paxinos G (ed.). <i>The Rat Nervous System</i>. 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II; Rodríguez, E. (June 2013). 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ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Mesencephalon" title="Template:Mesencephalon"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Mesencephalon" title="Template talk:Mesencephalon"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Mesencephalon" title="Special:EditPage/Template:Mesencephalon"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Anatomy_of_the_midbrain" style="font-size:114%;margin:0 4em">Anatomy of the <a href="/wiki/Midbrain" title="Midbrain">midbrain</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Midbrain_tectum" class="mw-redirect" title="Midbrain tectum">Tectum</a><br />(Dorsal)</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/Corpora_quadrigemina" title="Corpora quadrigemina">Corpora quadrigemina</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/Inferior_colliculus" title="Inferior colliculus">Inferior colliculus</a> <ul><li><a href="/wiki/Inferior_colliculus#Input_and_output_connections_of_IC" title="Inferior colliculus">Inferior brachium</a></li></ul></li> <li><a href="/wiki/Superior_colliculus" title="Superior colliculus">Superior colliculus</a> <ul><li><a href="/wiki/Superior_colliculus#Structure" title="Superior colliculus">Superior brachium</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</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/Pretectal_area" title="Pretectal area">Pretectal area</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Sensory / ascending</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/Spinotectal_tract" class="mw-redirect" title="Spinotectal tract">Spinotectal tract</a></li> <li><a href="/wiki/Central_tegmental_tract" title="Central tegmental tract">Central tegmental tract</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Motor / descending</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/Tectospinal_tract" title="Tectospinal tract">Tectospinal tract</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/Ventricular_system" title="Ventricular system">CSF</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cerebral_aqueduct" title="Cerebral aqueduct">Cerebral aqueduct</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cerebral_peduncle" title="Cerebral peduncle">Peduncle</a><br />(Ventral)</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/Midbrain_tegmentum" title="Midbrain tegmentum">Tegmentum</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Sensory / ascending</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>Lemnisci <ul><li><a href="/wiki/Medial_lemniscus" title="Medial lemniscus">Medial</a></li> <li><a href="/wiki/Lateral_lemniscus" title="Lateral lemniscus">Lateral</a></li></ul></li> <li><a href="/wiki/Medial_longitudinal_fasciculus" title="Medial longitudinal fasciculus">Ascending MLF</a> <ul><li><a href="/wiki/Vestibulo-oculomotor_fibers" title="Vestibulo-oculomotor fibers">Vestibulo-oculomotor fibers</a></li></ul></li> <li><a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinothalamic tract</a></li> <li><a href="/wiki/Ventral_trigeminal_tract" title="Ventral trigeminal tract">Ventral trigeminal tract</a></li> <li><a href="/wiki/Dentatothalamic_tract" title="Dentatothalamic tract">Dentatothalamic tract</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Motor / descending</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/Rubrospinal_tract" title="Rubrospinal tract">Rubrospinal tract</a></li> <li><a href="/wiki/Rubro-olivary_tract" title="Rubro-olivary tract">Rubro-olivary tract</a></li> <li><a href="/wiki/Medial_longitudinal_fasciculus" title="Medial longitudinal fasciculus">Descending MLF</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/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cranial_nerve_nucleus" title="Cranial nerve nucleus">cranial nuclei</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_afferent_fibers" class="mw-redirect" title="General somatic afferent fibers">GSA</a> <ul><li><a href="/wiki/Mesencephalic_nucleus_of_trigeminal_nerve" title="Mesencephalic nucleus of trigeminal nerve">Mesencephalic</a></li></ul></li> <li><a href="/wiki/General_somatic_efferent_fibers" class="mw-redirect" title="General somatic efferent fibers">GSE</a> <ul><li><a href="/wiki/Oculomotor_nucleus" title="Oculomotor nucleus">Oculomotor nucleus</a></li> <li><a href="/wiki/Trochlear_nucleus" title="Trochlear nucleus">Trochlear nucleus</a></li></ul></li> <li><a href="/wiki/General_visceral_efferent_fibers" class="mw-redirect" title="General visceral efferent fibers">GVE</a> <ul><li><a href="/wiki/Edinger%E2%80%93Westphal_nucleus" title="Edinger–Westphal nucleus">Edinger–Westphal nucleus</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ventral_tegmental_area" title="Ventral tegmental area">Ventral tegmental area</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/Rostromedial_tegmental_nucleus" title="Rostromedial tegmental nucleus">Rostromedial tegmental nucleus</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Periaqueductal_gray" title="Periaqueductal gray">Periaqueductal gray</a></li> <li><a href="/wiki/Red_nucleus" title="Red nucleus">Red nucleus</a></li> <li><a href="/wiki/Rostral_interstitial_nucleus_of_medial_longitudinal_fasciculus" title="Rostral interstitial nucleus of medial longitudinal fasciculus">Rostral interstitial nucleus of medial longitudinal fasciculus</a></li> <li><a href="/wiki/Parabrachial_area" class="mw-redirect" title="Parabrachial area">Parabrachial area</a></li> <li><a href="/wiki/Interpeduncular_nucleus" title="Interpeduncular nucleus">Interpeduncular nucleus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Midbrain_reticular_formation" title="Midbrain reticular formation">Midbrain reticular formation</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/Dorsal_tegmental_nucleus" title="Dorsal tegmental nucleus">Dorsal tegmental nucleus</a></li> <li><a href="/wiki/Raphe_nuclei" title="Raphe nuclei">Raphe nuclei</a> <ul><li><a href="/wiki/Dorsal_raphe_nucleus" title="Dorsal raphe nucleus">dorsal</a></li></ul></li> <li><a href="/wiki/Reticulotegmental_nucleus" title="Reticulotegmental nucleus">Reticulotegmental nucleus</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%">Base</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/White_matter" title="White matter">White</a> / <a href="/wiki/Cerebral_crus" title="Cerebral crus">Cerebral crus</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_tracts" title="Pyramidal tracts">Pyramidal tracts</a> <ul><li><a href="/wiki/Corticospinal_tract" title="Corticospinal tract">Corticospinal tract</a></li> <li><a href="/wiki/Corticobulbar_tract" title="Corticobulbar tract">Corticobulbar tract</a></li></ul></li> <li><a href="/wiki/Corticopontine_fibers" title="Corticopontine fibers">Corticopontine tract</a> <ul><li><a href="/wiki/Frontopontine_fibers" title="Frontopontine fibers">Frontopontine fibers</a></li> <li><a href="/wiki/Temporopontine_fibers" title="Temporopontine fibers">Temporopontine fibers</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey</a> / <a href="/wiki/Substantia_nigra" title="Substantia nigra">Substantia nigra</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/Pars_compacta" title="Pars compacta">Pars compacta</a></li> <li><a href="/wiki/Pars_reticulata" title="Pars reticulata">Pars reticulata</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Surface</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/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">Superior cerebellar peduncle</a> <ul><li><a href="/wiki/Decussation_of_superior_cerebellar_peduncles" class="mw-redirect" title="Decussation of superior cerebellar peduncles">Decussation</a></li></ul></li> <li><a href="/wiki/Interpeduncular_fossa" title="Interpeduncular fossa">Interpeduncular fossa</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Anatomy_of_the_diencephalon_of_the_human_brain" 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:Diencephalon" title="Template:Diencephalon"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Diencephalon" title="Template talk:Diencephalon"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Diencephalon" title="Special:EditPage/Template:Diencephalon"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Anatomy_of_the_diencephalon_of_the_human_brain" style="font-size:114%;margin:0 4em">Anatomy of the <a href="/wiki/Diencephalon" title="Diencephalon">diencephalon</a> of the <a href="/wiki/Human_brain" title="Human brain">human brain</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Epithalamus" title="Epithalamus">Epithalamus</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%">Surface</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/Pineal_gland" title="Pineal gland">Pineal gland</a></li> <li><a href="/wiki/Habenula" title="Habenula">Habenula</a></li> <li><a href="/wiki/Habenular_trigone" title="Habenular trigone">Habenular trigone</a></li> <li><a href="/wiki/Habenular_commissure" title="Habenular commissure">Habenular commissure</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</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/Pretectal_area" title="Pretectal area">Pretectal area</a></li> <li><a href="/wiki/Habenular_nuclei" class="mw-redirect" title="Habenular nuclei">Habenular nuclei</a></li> <li><a class="mw-selflink selflink">Subcommissural organ</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/Thalamus" title="Thalamus">Thalamus</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%">Surface</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/Stria_medullaris" title="Stria medullaris">Stria medullaris</a></li> <li><a href="/wiki/Thalamic_reticular_nucleus" title="Thalamic reticular nucleus">Thalamic reticular nucleus</a></li> <li><a href="/wiki/Taenia_thalami" title="Taenia thalami">Taenia thalami</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a>/<br /><a href="/wiki/List_of_thalamic_nuclei" title="List of thalamic nuclei">nuclei</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><i>paired:</i> <a href="/wiki/Anterior_nuclei_of_thalamus" title="Anterior nuclei of thalamus">AN</a></li> <li><a href="/wiki/Ventral_nuclear_group" title="Ventral nuclear group">Ventral</a> <ul><li><a href="/wiki/Ventral_anterior_nucleus" title="Ventral anterior nucleus">VA</a>/<a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a></li> <li><a href="/wiki/Ventral_posterior_nucleus" title="Ventral posterior nucleus">VP</a>/<a href="/wiki/Ventral_posteromedial_nucleus" title="Ventral posteromedial nucleus">VPM</a>/<a href="/wiki/Ventral_posterolateral_nucleus" title="Ventral posterolateral nucleus">VPL</a></li></ul></li> <li><a href="/wiki/Lateral_nuclear_group" title="Lateral nuclear group">Lateral</a> <ul><li><a href="/wiki/Lateral_dorsal_nucleus_of_thalamus" title="Lateral dorsal nucleus of thalamus">LD</a></li> <li><a href="/wiki/Lateral_posterior_nucleus_of_thalamus" title="Lateral posterior nucleus of thalamus">LP</a></li> <li><a href="/wiki/Pulvinar_nuclei" title="Pulvinar nuclei">Pulvinar nuclei</a></li></ul></li> <li><a href="/wiki/Metathalamus" class="mw-redirect" title="Metathalamus">Metathalamus</a> <ul><li><a href="/wiki/Medial_geniculate_nucleus" title="Medial geniculate nucleus">MG</a></li> <li><a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">LG</a> <ul><li><a href="/wiki/Parvocellular_cell" title="Parvocellular cell">P cell</a></li> <li><a href="/wiki/Magnocellular_cell" title="Magnocellular cell">M cell</a></li> <li><a href="/wiki/Koniocellular_cell" title="Koniocellular cell">K cell</a></li></ul></li></ul></li></ul> <ul><li><i>midline:</i> <a href="/wiki/Medial_dorsal_nucleus" title="Medial dorsal nucleus">MD</a></li> <li><a href="/wiki/Intralaminar_nuclei_of_thalamus" class="mw-redirect" title="Intralaminar nuclei of thalamus">Intralaminar</a> <ul><li><a href="/wiki/Centromedian_nucleus" title="Centromedian nucleus">Centromedian</a></li></ul></li> <li><a href="/wiki/Midline_nuclear_group" title="Midline nuclear group">Midline nuclear group</a></li> <li><a href="/wiki/Interthalamic_adhesion" title="Interthalamic adhesion">Interthalamic adhesion</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/White_matter" title="White matter">White matter</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/Mammillothalamic_tract" title="Mammillothalamic tract">Mammillothalamic tract</a></li> <li><a href="/wiki/Pallidothalamic_tracts" title="Pallidothalamic tracts">Pallidothalamic tracts</a> <ul><li><a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a></li> <li><a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a></li> <li><a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a></li></ul></li> <li><i><a href="/wiki/Dorsal_column-medial_lemniscus_pathway" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">DCML</a></i> <ul><li><a href="/wiki/Medial_lemniscus" title="Medial lemniscus">Medial lemniscus</a></li> <li><a href="/wiki/Trigeminal_lemniscus" title="Trigeminal lemniscus">Trigeminal lemniscus</a></li></ul></li> <li><a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinothalamic tract</a></li> <li><a href="/wiki/Lateral_lemniscus" title="Lateral lemniscus">Lateral lemniscus</a></li> <li><a href="/wiki/Dentatothalamic_tract" title="Dentatothalamic tract">Dentatothalamic tract</a></li> <li><a href="/wiki/Acoustic_radiation" title="Acoustic radiation">Acoustic radiation</a></li> <li><a href="/wiki/Optic_radiation" title="Optic radiation">Optic radiation</a></li> <li><a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a></li> <li><a href="/wiki/Ventral_trigeminal_tract" title="Ventral trigeminal tract">Ventral trigeminal tract</a></li></ul> <ul><li><a href="/wiki/Medullary_laminae_of_thalamus" title="Medullary laminae of thalamus">Medullary laminae</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/Hypothalamus" title="Hypothalamus">Hypothalamus</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%">Surface</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/Median_eminence" title="Median eminence">Median eminence</a>/<a href="/wiki/Tuber_cinereum" title="Tuber cinereum">Tuber cinereum</a></li> <li><a href="/wiki/Mammillary_body" title="Mammillary body">Mammillary body</a></li> <li><a href="/wiki/Pituitary_stalk" title="Pituitary stalk">Infundibulum</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Autonomic_nervous_system" title="Autonomic nervous system">Autonomic</a> zones</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/Anterior_hypothalamic_nucleus" title="Anterior hypothalamic nucleus">Anterior (parasympathetic/heat loss)</a></li> <li><a href="/wiki/Posterior_nucleus_of_hypothalamus" title="Posterior nucleus of hypothalamus">Posterior (sympathetic/heat conservation)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Endocrine_system" title="Endocrine system">Endocrine</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><i><a href="/wiki/Posterior_pituitary" title="Posterior pituitary">posterior pituitary</a>:</i> <a href="/wiki/Paraventricular_nucleus_of_hypothalamus" title="Paraventricular nucleus of hypothalamus">Paraventricular</a> <ul><li><a href="/wiki/Magnocellular_neurosecretory_cell" title="Magnocellular neurosecretory cell">Magnocellular neurosecretory cell</a></li> <li><a href="/wiki/Parvocellular_neurosecretory_cell" title="Parvocellular neurosecretory cell">Parvocellular neurosecretory cell</a></li></ul></li> <li><a href="/wiki/Supraoptic_nucleus" title="Supraoptic nucleus">Supraoptic</a> <ul><li><a href="/wiki/Oxytocin" title="Oxytocin">oxytocin</a>/<a href="/wiki/Vasopressin" title="Vasopressin">vasopressin</a></li></ul></li></ul> <ul><li><i>other:</i> <a href="/wiki/Arcuate_nucleus_(hypothalamus)" title="Arcuate nucleus (hypothalamus)">Arcuate (dopamine/GHRH)</a></li> <li><a href="/wiki/Preoptic_area" title="Preoptic area">Preoptic (GnRH)</a></li> <li><a href="/wiki/Suprachiasmatic_nucleus" title="Suprachiasmatic nucleus">Suprachiasmatic (melatonin)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Emotion" title="Emotion">Emotion</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/Lateral_hypothalamus" title="Lateral hypothalamus">Lateral</a></li> <li><a href="/wiki/Ventromedial_nucleus" class="mw-redirect" title="Ventromedial nucleus">Ventromedial</a></li> <li><a href="/wiki/Dorsomedial_hypothalamic_nucleus" title="Dorsomedial hypothalamic nucleus">Dorsomedial</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/White_matter" title="White matter">White matter</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><i>afferent</i> <ul><li><a href="/wiki/Stria_terminalis" title="Stria terminalis">Stria terminalis</a></li> <li><a href="/wiki/Medial_forebrain_bundle" title="Medial forebrain bundle">Medial forebrain bundle</a></li> <li><a href="/wiki/Retinohypothalamic_tract" title="Retinohypothalamic tract">Retinohypothalamic tract</a></li></ul></li> <li><i>efferent</i> <ul><li><a href="/wiki/Mammillothalamic_tract" title="Mammillothalamic tract">Mammillothalamic tract</a></li> <li><a href="/wiki/Dorsal_longitudinal_fasciculus" title="Dorsal longitudinal fasciculus">Dorsal longitudinal fasciculus</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pituitary_gland" title="Pituitary gland">Pituitary</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/Posterior_pituitary" title="Posterior pituitary">Posterior</a> is diencephalon, but <a href="/wiki/Anterior_pituitary" title="Anterior pituitary">anterior</a> is glandular</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Subthalamus" title="Subthalamus">Subthalamus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Subthalamic_nucleus" title="Subthalamic nucleus">Subthalamic nucleus</a></li> <li><a href="/wiki/Zona_incerta" title="Zona incerta">Zona incerta</a></li> <li><a href="/wiki/Nuclei_campi_perizonalis" class="mw-redirect" title="Nuclei campi perizonalis">Nuclei campi perizonalis</a> (<a href="/wiki/Fields_of_Forel" title="Fields of Forel">Fields of Forel</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 authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a 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