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Subthalamic nucleus - Wikipedia
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<li id="toc-Structure" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Structure"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Structure</span> </div> </a> <ul id="toc-Structure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Afferent_axons" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Afferent_axons"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Afferent axons</span> </div> </a> <ul id="toc-Afferent_axons-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Efferent_targets" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Efferent_targets"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Efferent targets</span> </div> </a> <ul id="toc-Efferent_targets-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Physiology" class="vector-toc-list-item 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class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:lemonchiffon">Subthalamic nucleus</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Basal-ganglia-coronal-sections-large.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Basal-ganglia-coronal-sections-large.png/250px-Basal-ganglia-coronal-sections-large.png" decoding="async" width="250" height="109" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Basal-ganglia-coronal-sections-large.png/375px-Basal-ganglia-coronal-sections-large.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/33/Basal-ganglia-coronal-sections-large.png/500px-Basal-ganglia-coronal-sections-large.png 2x" data-file-width="800" data-file-height="350" /></a></span><div class="infobox-caption">Coronal slices of human brain showing the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> (<a href="/wiki/External_globus_pallidus" title="External globus pallidus">external globus pallidus</a> (GPe) and <a href="/wiki/Internal_globus_pallidus" title="Internal globus pallidus">internal globus pallidus</a> (GPi)), <b>subthalamic nucleus</b> (STN) and <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> (SN).</div></td></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:DA-loops_in_PD.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/DA-loops_in_PD.svg/250px-DA-loops_in_PD.svg.png" decoding="async" width="250" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/DA-loops_in_PD.svg/375px-DA-loops_in_PD.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/57/DA-loops_in_PD.svg/500px-DA-loops_in_PD.svg.png 2x" data-file-width="1050" data-file-height="834" /></a></span><div class="infobox-caption">DA-loops in <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a></div></td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Details</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Part of</th><td class="infobox-data"><a href="/wiki/Subthalamus" title="Subthalamus">Subthalamus</a> (physically); <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> (functionally)</td></tr><tr><th colspan="2" class="infobox-header" style="background-color: #efefef">Identifiers</th></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Latin" title="Latin">Latin</a></th><td class="infobox-data"><i>nucleus subthalamicus</i></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em">Acronym(s)</th><td class="infobox-data">STN</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=D020531">D020531</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=435">435</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/nlx_anat_1010002">nlx_anat_1010002</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/14.1.08.702%20Entity%20TA98%20EN.htm">A14.1.08.702</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=5709">5709</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/Foundational_Model_of_Anatomy" title="Foundational Model of Anatomy">FMA</a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://bioportal.bioontology.org/ontologies/FMA/?p=classes&conceptid=http%3A%2F%2Fpurl.org%2Fsig%2Font%2Ffma%2Ffma62035">62035</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">[<a href="https://www.wikidata.org/wiki/Q2270424" class="extiw" title="d:Q2270424">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>The <b>subthalamic nucleus</b> (<b>STN</b>) is a small lens-shaped <a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">nucleus</a> in the <a href="/wiki/Brain" title="Brain">brain</a> where it is, from a functional point of view, part of the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> system. In terms of anatomy, it is the major part of the <a href="/wiki/Subthalamus" title="Subthalamus">subthalamus</a>. As suggested by its name, the subthalamic nucleus is located <a href="/wiki/Anatomical_terms_of_location" title="Anatomical terms of location">ventral</a> to the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>. It is also dorsal to the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> and medial to the <a href="/wiki/Internal_capsule" title="Internal capsule">internal capsule</a>. It was first described by <a href="/wiki/Jules_Bernard_Luys" title="Jules Bernard Luys">Jules Bernard Luys</a> in 1865,<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> and the term <i>corpus Luysi</i> or <i>Luys' body</i> is still sometimes used. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Anatomy">Anatomy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=1" title="Edit section: Anatomy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv/220px--Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="193" data-durationhint="18" data-mwtitle="Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv" data-mwprovider="wikimediacommons" resource="/wiki/File:Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv.480p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="480p.vp9.webm" data-width="546" data-height="480" /><source src="//upload.wikimedia.org/wikipedia/commons/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv" type="video/ogg; codecs="theora"" data-width="686" data-height="602" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="274" data-height="240" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="410" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/0/0d/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv/Ultra-High-Field-MRI-Post-Mortem-Structural-Connectivity-of-the-Human-Subthalamic-Nucleus-Video1.ogv.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="410" data-height="360" /></video></span><figcaption>Structural connectivity of the human subthalamic nucleus as visualized through <a href="/wiki/Diffusion-weighted_MRI" class="mw-redirect" title="Diffusion-weighted MRI">diffusion-weighted MRI</a>.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Structure">Structure</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=2" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The principal type of <a href="/wiki/Neuron" title="Neuron">neuron</a> found in the subthalamic nucleus has rather long, sparsely spiny <a href="/wiki/Dendrite" title="Dendrite">dendrites</a>.<sup id="cite_ref-Afsharpour1985_2-0" class="reference"><a href="#cite_note-Afsharpour1985-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> In the more centrally located neurons, the dendritic arbors have a more <a href="/wiki/Ellipsoid" title="Ellipsoid">ellipsoidal</a> shape.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The dimensions of these arbors (1200 μm, 600 μm, and 300 μm) are similar across many species—including rat, cat, monkey and human—which is unusual. However, the number of neurons increases with brain size as well as the external dimensions of the nucleus. The principal neurons are <a href="/wiki/Glutamatergic" title="Glutamatergic">glutamatergic</a>, which give them a particular functional position in the basal ganglia system. In humans there are also a small number (about 7.5%) of <a href="/wiki/GABA" title="GABA">GABAergic</a> <a href="/wiki/Interneuron" title="Interneuron">interneurons</a> that participate in the local circuitry; however, the dendritic arbors of subthalamic neurons shy away from the border and primarily interact with one another.<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> </p><p>The structure of the subthalamic nucleus has not yet been fully explored and understood, but it is likely composed of several internal domains. The primate subthalamic nucleus is often divided in three internal anatomical-functional domains. However, this so-called <i>tripartite model</i> has been debated because it does not fully explain the complexity of the subthalamic nucleus in brain function.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Afferent_axons">Afferent axons</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=3" title="Edit section: Afferent axons"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The subthalamic nucleus receives its main input from the <a href="/wiki/External_globus_pallidus" title="External globus pallidus">external globus pallidus</a> (GPe),<sup id="cite_ref-pmid2350684_8-0" class="reference"><a href="#cite_note-pmid2350684-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> not so much through the <a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">ansa lenticularis</a> as often said but by radiating 'comb' fibers crossing the medial pallidum first and the internal capsule (forming part of <a href="/wiki/Ludwig_Edinger" title="Ludwig Edinger">Edinger</a>'s comb system, see figure), as well as the ansa subthalamica.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> These <a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">afferents</a> are GABAergic, inhibiting neurons in the subthalamic nucleus. Excitatory, glutamatergic inputs come from the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a> (entire frontal cortex with a predominance for motor, premotor and oculomotor input to the posterolateral part of the nucleus), and from the pars <a href="/w/index.php?title=Parafascicularis&action=edit&redlink=1" class="new" title="Parafascicularis (page does not exist)">parafascicularis</a> of the <a href="/w/index.php?title=Central_complex&action=edit&redlink=1" class="new" title="Central complex (page does not exist)">central complex</a>. The subthalamic nucleus also receives <a href="/wiki/Neuromodulator" class="mw-redirect" title="Neuromodulator">neuromodulatory</a> inputs, notably <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> axons from the <a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> pars compacta.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> It also receives inputs from the <a href="/wiki/Pedunculopontine_nucleus" title="Pedunculopontine nucleus">pedunculopontine nucleus</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Efferent_targets">Efferent targets</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=4" title="Edit section: Efferent targets"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The axons of subthalamic nucleus neurons leave the nucleus dorsally. The efferent axons are glutamatergic (excitatory). Except for the connection to the striatum (17.3% in macaques), most of the subthalamic principal neurons are multitargets and directed to the other elements of the core of the basal ganglia.<sup id="cite_ref-pmid418083_11-0" class="reference"><a href="#cite_note-pmid418083-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Some send axons to the substantia nigra medially and to the medial and lateral nuclei of the pallidum laterally (3-target, 21.3%). Some are 2-target with the lateral pallidum and the substantia nigra (2.7%) or the lateral pallidum and the medial (48%). Less are single target for the lateral pallidum. In the pallidum, subthalamic terminals end in bands parallel to the pallidal border.<sup id="cite_ref-pmid418083_11-1" class="reference"><a href="#cite_note-pmid418083-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Smith1990_12-0" class="reference"><a href="#cite_note-Smith1990-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> When all axons reaching this target are added, the main efference of the subthalamic nucleus is, in 82.7% of the cases, clearly the <a href="/wiki/Internal_globus_pallidus" title="Internal globus pallidus">internal globus pallidus</a> (GPi). </p><p>Some researchers have reported internal <a href="/wiki/Axon" title="Axon">axon</a> collaterals.<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> However, there is little functional evidence for this. </p> <div class="mw-heading mw-heading2"><h2 id="Physiology">Physiology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=5" title="Edit section: Physiology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Basal_ganglia_circuits.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/320px-Basal_ganglia_circuits.svg.png" decoding="async" width="320" height="460" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/480px-Basal_ganglia_circuits.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Basal_ganglia_circuits.svg/640px-Basal_ganglia_circuits.svg.png 2x" data-file-width="1010" data-file-height="1453" /></a><figcaption>Anatomical overview of the main circuits of the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>. Subthalamic nucleus is shown in red. Picture shows 2 coronal slices that have been superimposed to include the involved basal ganglia structures. + and - signs at the point of the arrows indicate respectively whether the pathway is excitatory or inhibitory in effect. <style data-mw-deduplicate="TemplateStyles:r1239334494">@media screen{html.skin-theme-clientpref-night .mw-parser-output div:not(.notheme)>.tmp-color,html.skin-theme-clientpref-night .mw-parser-output p>.tmp-color,html.skin-theme-clientpref-night .mw-parser-output table:not(.notheme) .tmp-color{color:inherit!important}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output div:not(.notheme)>.tmp-color,html.skin-theme-clientpref-os .mw-parser-output p>.tmp-color,html.skin-theme-clientpref-os .mw-parser-output table:not(.notheme) .tmp-color{color:inherit!important}}</style><span class="tmp-color" style="color:green">Green arrows</span> refer to excitatory <a href="/wiki/Glutamic_acid" title="Glutamic acid">glutamatergic</a> pathways, <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239334494"><span class="tmp-color" style="color:red">red arrows</span> refer to inhibitory <a href="/wiki/Gamma-Aminobutyric_acid" class="mw-redirect" title="Gamma-Aminobutyric acid">GABAergic</a> pathways and <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239334494"><span class="tmp-color" style="color:turquoise">turquoise arrows</span> refer to <a href="/wiki/Dopamine" title="Dopamine">dopaminergic</a> pathways that are excitatory on the direct pathway and inhibitory on the indirect pathway.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Subthalamic_nucleus">Subthalamic nucleus</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=6" title="Edit section: Subthalamic nucleus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first intracellular electrical recordings of subthalamic neurons were performed using sharp electrodes in a rat slice preparation.<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 2010)">citation needed</span></a></i>]</sup> In these recordings three key observations were made, all three of which have dominated subsequent reports of subthalamic firing properties. The first observation was that, in the absence of current injection or synaptic stimulation, the majority of cells were spontaneously firing. The second observation is that these cells are capable of transiently firing at very high frequencies. The third observation concerns non-linear behaviors when cells are transiently depolarized after being hyperpolarized below –65mV. They are then able to engage voltage-gated calcium and sodium currents to fire bursts of action potentials. </p><p>Several recent studies have focused on the autonomous pacemaking ability of subthalamic neurons. These cells are often referred to as "fast-spiking pacemakers",<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> since they can generate spontaneous <a href="/wiki/Action_potential" title="Action potential">action potentials</a> at rates of 80 to 90 Hz in primates. </p><p>Oscillatory and synchronous activity<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><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> is likely to be a typical pattern of discharge in subthalamic neurons recorded from patients and animal models characterized by the loss of dopaminergic cells in the <a href="/wiki/Substantia_nigra_pars_compacta" class="mw-redirect" title="Substantia nigra pars compacta">substantia nigra pars compacta</a>, which is the principal pathology that underlies <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Lateropallido-subthalamic_system">Lateropallido-subthalamic system</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=7" title="Edit section: Lateropallido-subthalamic system"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Strong reciprocal connections link the subthalamic nucleus and the external segment of the <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a>. Both are fast-spiking pacemakers. Together, they are thought to constitute the "central pacemaker of the basal ganglia"<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> with synchronous bursts. </p><p>The connection of the lateral pallidum with the subthalamic nucleus is also the one in the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> system where the reduction between emitter/receiving elements is likely the strongest. In terms of volume, in humans, the lateral pallidum measures 808 mm<sup>3</sup>, the subthalamic nucleus only 158 mm<sup>3</sup>.<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> This translated in numbers of neurons represents a strong compression with loss of map precision. </p><p>Some axons from the lateral pallidum go to the striatum.<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> The activity of the medial pallidum is influenced by afferences from the lateral pallidum and from the subthalamic nucleus.<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> The same for the <a href="/wiki/Substantia_nigra_pars_reticulata" class="mw-redirect" title="Substantia nigra pars reticulata">substantia nigra pars reticulata</a>.<sup id="cite_ref-Smith1990_12-1" class="reference"><a href="#cite_note-Smith1990-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The subthalamic nucleus sends axons to another regulator: the pedunculo-pontine complex (id). </p><p>The lateropallido-subthalamic system is thought to play a key role in the generation of the patterns of activity seen in <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson's disease</a>.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Pathophysiology">Pathophysiology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=8" title="Edit section: Pathophysiology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Lesioning the STN leads to alleviation of motor symptoms such as <a href="/wiki/Hypokinesia" title="Hypokinesia">akinesia</a>, <a href="/wiki/Rigidity_(neurology)" class="mw-redirect" title="Rigidity (neurology)">rigidity</a>, and <a href="/wiki/Tremor" title="Tremor">tremor</a> in <a href="/wiki/Parkinson%27s_disease" title="Parkinson's disease">Parkinson disease</a>. This was first shown in the <a href="/wiki/MPTP" title="MPTP">MPTP</a> primate model in a paper by <a href="/wiki/Hagai_Bergman" title="Hagai Bergman">Bergman</a> and colleagues.<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> This inspired Benazzouz and colleagues to probe deep brain stimulation of the nucleus, which was known to exert similar effects as ablative lesions.<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> Soon after, the team of <a href="/wiki/Alim_Louis_Benabid" title="Alim Louis Benabid">Alim Louis Benabid</a> showed that deep brain stimulation of the nucleus leads to symptom relief in human patients with Parkinson disease, as well,<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> which led to the establishment of the currently <a href="/wiki/Food_and_Drug_Administration" title="Food and Drug Administration">FDA</a> approved and widely applied form of <a href="/wiki/Deep_brain_stimulation" title="Deep brain stimulation">deep brain stimulation</a>. The first to be stimulated are the terminal arborisations of afferent axons, which modify the activity of subthalamic neurons. However, it has been shown in thalamic slices from mice,<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> that the stimulus also causes nearby astrocytes to release <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP), a precursor to <a href="/wiki/Adenosine" title="Adenosine">adenosine</a> (through a catabolic process). In turn, adenosine A1 receptor activation depresses excitatory transmission in the thalamus, thus mimicking <a href="/wiki/Ablation" title="Ablation">ablation</a> of the subthalamic nucleus. </p><p>Before the <a href="/wiki/Hagai_Bergman" title="Hagai Bergman">Bergman</a> paper, the stereotactic field avoided lesioning the nucleus, since it was known that unilateral destruction or disruption of the subthalamic nucleus — which may result from naturally occurring strokes — may lead to <a href="/wiki/Hemiballismus" title="Hemiballismus">hemiballismus</a>. While this remains generally true, iatrogenic lesioning of the STN has been carried out numerous times and has recently gained new wind with the advent of <a href="/wiki/HIFU" class="mw-redirect" title="HIFU">MR guided focused ultrasound</a>, which has also been probed for subthalamic nucleotomies to treat Parkinson disease.<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> Curiously, a team around <a href="/wiki/Michael_D._Fox" title="Michael D. Fox">Michael Fox</a> could recently show that, while some lesions that led to hemiballism were indeed in and around the STN, the majority of reported cases were in other regions of the brain.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>As one of the STN's suspected functions is in impulse control, dysfunction in this region has been implicated in <a href="/wiki/Obsessive%E2%80%93compulsive_disorder" title="Obsessive–compulsive disorder">obsessive–compulsive disorder</a>.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Application of high frequency pulses by <a href="/wiki/Deep_brain_stimulation" title="Deep brain stimulation">deep brain stimulation</a> has shown some promise in correcting severe impulsive behavior and has been <a href="/wiki/Food_and_Drug_Administration" title="Food and Drug Administration">FDA</a> approved for treatment resistant cases with the disorder.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </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=Subthalamic_nucleus&action=edit&section=9" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The function of the STN is unknown, but current theories place it as a component of the <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> control system that may perform action selection. It is plays a part in the so-called "hyperdirect" and "indirect" pathways of motor control, contrasting with the direct pathway which is thought to bypass the STN on its way from Striatum to internal pallidum. STN dysfunction has been implicated in motor symptoms such as rigidity, bradykinesia and tremor,<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> behavioral features such as stopping of ongoing movements<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> or impulsivity in individuals presented with two equally rewarding stimuli.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>The physiological role of the STN has been for long hidden by its pathological role. But lately, the research on the physiology of the STN led to the discovery that the STN is required to achieve intended movement, including locomotion, balance and motor coordination. It is involved in stopping or interrupting on-going motor tasks. Moreover, STN excitation was generally correlated with significant reduction in locomotor activity, while in contrast, STN inhibition enhanced locomotion.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Additional_images">Additional images</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=10" title="Edit section: Additional images"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul class="gallery mw-gallery-traditional"> <li class="gallerybox" style="width: 155px"> <div class="thumb" style="width: 150px; height: 150px;"><span typeof="mw:File"><a href="/wiki/File:Gray717.png" class="mw-file-description" title="Coronal section of brain immediately in front of pons. Subthalamic nucleus labeled as "Nucleus of Luys"."><img alt="Coronal section of brain immediately in front of pons. Subthalamic nucleus labeled as "Nucleus of Luys"." src="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Gray717.png/106px-Gray717.png" decoding="async" width="106" height="120" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Gray717.png/158px-Gray717.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/20/Gray717.png/211px-Gray717.png 2x" data-file-width="500" data-file-height="568" /></a></span></div> <div class="gallerytext">Coronal section of brain immediately in front of pons. Subthalamic nucleus labeled as "Nucleus of Luys".</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Subthalamic_nucleus&action=edit&section=11" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Subthalamic_nucleus" class="extiw" title="commons:Category:Subthalamic nucleus">Subthalamic nucleus</a></span>.</div></div> </div> <ul><li><a href="/wiki/Primate_basal_ganglia" title="Primate basal ganglia">Primate basal ganglia</a></li> <li><a href="/wiki/Mesencephalic_locomotor_region" title="Mesencephalic locomotor region">Mesencephalic locomotor region</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=Subthalamic_nucleus&action=edit&section=12" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFLuys1865" class="citation book cs1 cs1-prop-foreign-lang-source"><a href="/wiki/Jules_Bernard_Luys" title="Jules Bernard Luys">Luys JB</a> (1865). <i>Recherches sur le système cérébro-spinal, sa structure, ses fonctions et ses maladies</i> (in French). Paris: Baillière.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Recherches+sur+le+syst%C3%A8me+c%C3%A9r%C3%A9bro-spinal%2C+sa+structure%2C+ses+fonctions+et+ses+maladies&rft.place=Paris&rft.pub=Bailli%C3%A8re&rft.date=1865&rft.aulast=Luys&rft.aufirst=JB&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASubthalamic+nucleus" class="Z3988"></span></span> </li> <li id="cite_note-Afsharpour1985-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-Afsharpour1985_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAfsharpour1985" class="citation journal cs1">Afsharpour S (June 1985). 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Brain+Research&rft.atitle=Experimental+investigation+into+the+role+of+the+subthalamic+nucleus+%28STN%29+in+motor+control+using+optogenetics+in+mice&rft.volume=1755&rft.pages=147226&rft.date=2021-03&rft_id=info%3Adoi%2F10.1016%2Fj.brainres.2020.147226&rft_id=info%3Apmid%2F33358727&rft.aulast=Guillaumin&rft.aufirst=A&rft.au=Serra%2C+GP&rft.au=Georges%2C+F&rft.au=Wall%C3%A9n-Mackenzie%2C+%C3%85&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.brainres.2020.147226&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASubthalamic+nucleus" class="Z3988"></span></span> </li> </ol></div></div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output 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.navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="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"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template: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 href="/wiki/Subcommissural_organ" title="Subcommissural organ">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 class="mw-selflink selflink">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" aria-labelledby="Brain_and_spinal_cord:_neural_tracts_and_fasciculi" 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:Neural_tracts" title="Template:Neural tracts"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Neural_tracts" title="Template talk:Neural tracts"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Neural_tracts" title="Special:EditPage/Template:Neural tracts"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Brain_and_spinal_cord:_neural_tracts_and_fasciculi" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_brain" title="Human brain">Brain</a> and <a href="/wiki/Spinal_cord" title="Spinal cord">spinal cord</a>: <a href="/wiki/Neural_pathway" title="Neural pathway">neural tracts</a> and <a href="/wiki/Nerve_fascicle" title="Nerve fascicle">fasciculi</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Sensory</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/Dorsal_column-medial_lemniscus_pathway" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">DCML</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%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#First_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">1°</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/Lamellar_corpuscle" class="mw-redirect" title="Lamellar corpuscle">Pacinian corpuscle</a>/<a href="/wiki/Tactile_corpuscle" title="Tactile corpuscle">Meissner's corpuscle</a> → <b><a href="/wiki/Posterior_column" class="mw-redirect" title="Posterior column">Posterior column</a></b> (<a href="/wiki/Gracile_fasciculus" class="mw-redirect" title="Gracile fasciculus">Gracile fasciculus</a>/<a href="/wiki/Cuneate_fasciculus" class="mw-redirect" title="Cuneate fasciculus">Cuneate fasciculus</a>) → <a href="/wiki/Gracile_nucleus" class="mw-redirect" title="Gracile nucleus">Gracile nucleus</a>/<a href="/wiki/Cuneate_nucleus" class="mw-redirect" title="Cuneate nucleus">Cuneate nucleus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#Second_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">2°</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/Sensory_decussation" title="Sensory decussation">sensory decussation</a>/arcuate fibers</i> (<a href="/wiki/Posterior_external_arcuate_fibers" class="mw-redirect" title="Posterior external arcuate fibers">Posterior external arcuate fibers</a>, <a href="/wiki/Internal_arcuate_fibers" title="Internal arcuate fibers">Internal arcuate fibers</a>) → <a href="/wiki/Medial_lemniscus" title="Medial lemniscus">Medial lemniscus</a>/<a href="/wiki/Trigeminal_lemniscus" title="Trigeminal lemniscus">Trigeminal lemniscus</a> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_posterolateral_nucleus" title="Ventral posterolateral nucleus">VPL</a>, <a href="/wiki/Ventral_posteromedial_nucleus" title="Ventral posteromedial nucleus">VPM</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;background: transparent;"><a href="/wiki/Dorsal_column-medial_lemniscus_pathway#Third_neuron" class="mw-redirect" title="Dorsal column-medial lemniscus pathway">3°</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_limb_of_internal_capsule" class="mw-redirect" title="Posterior limb of internal capsule">Posterior limb of internal capsule</a> → <a href="/wiki/Postcentral_gyrus" title="Postcentral gyrus">Postcentral gyrus</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/Anterolateral_system" class="mw-redirect" title="Anterolateral system">Anterolateral</a>/<br /><a href="/wiki/Pain" title="Pain">pain</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/Nociception#Neospinothalamic_tract" title="Nociception">Fast</a>/lateral</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1° (<a href="/wiki/Free_nerve_ending" title="Free nerve ending">Free nerve ending</a> → <a href="/wiki/A_delta_fiber" class="mw-redirect" title="A delta fiber">A delta fiber</a>) → 2° (<a href="/wiki/Anterior_white_commissure" title="Anterior white commissure">Anterior white commissure</a> → <b><a href="/wiki/Lateral_spinothalamic_tract" class="mw-redirect" title="Lateral spinothalamic tract">Lateral</a> and <a href="/wiki/Anterior_spinothalamic_tract" class="mw-redirect" title="Anterior spinothalamic tract">Anterior</a> <a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinothalamic tract</a></b> → <a href="/wiki/Spinothalamic_tract" title="Spinothalamic tract">Spinal lemniscus</a> → <a href="/wiki/Ventral_posterolateral_nucleus" title="Ventral posterolateral nucleus">VPL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 3° (<a href="/wiki/Postcentral_gyrus" title="Postcentral gyrus">Postcentral gyrus</a>) → 4° (<a href="/wiki/Posterior_parietal_cortex" title="Posterior parietal cortex">Posterior parietal cortex</a>)</li></ul> <p>2° (<b><a href="/wiki/Spinomesencephalic_tract" class="mw-redirect" title="Spinomesencephalic tract">Spinomesencephalic tract</a></b> → <a href="/wiki/Superior_colliculus" title="Superior colliculus">Superior colliculus</a> of <a href="/wiki/Midbrain_tectum" class="mw-redirect" title="Midbrain tectum">Midbrain tectum</a>) </p> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Nociception#Paleospinothalamic_tract" title="Nociception">Slow</a>/medial</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li>1° (<a href="/wiki/Group_C_nerve_fiber" title="Group C nerve fiber">Group C nerve fiber</a> → <b><a href="/wiki/Spinoreticular_tract" title="Spinoreticular tract">Spinoreticular tract</a></b> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a>) → 2° (<a href="/wiki/Medial_dorsal_nucleus" title="Medial dorsal nucleus">MD</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 3° (<a href="/wiki/Cingulate_cortex" title="Cingulate cortex">Cingulate cortex</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%">Motor</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/Pyramidal_tracts" title="Pyramidal tracts">Pyramidal</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>flexion:</i> <a href="/wiki/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a> → <a href="/wiki/Posterior_limb_of_internal_capsule" class="mw-redirect" title="Posterior limb of internal capsule">Posterior limb of internal capsule</a> → <a href="/wiki/Decussation_of_pyramids" class="mw-redirect" title="Decussation of pyramids">Decussation of pyramids</a> → <b><a href="/wiki/Pyramidal_tracts" title="Pyramidal tracts">Corticospinal tract</a> (<a href="/wiki/Lateral_corticospinal_tract" title="Lateral corticospinal tract">Lateral</a>, <a href="/wiki/Anterior_corticospinal_tract" title="Anterior corticospinal tract">Anterior</a>)</b> → <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/Extrapyramidal_system" title="Extrapyramidal system">Extrapyramidal</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%"><span class="nobold"><i>flexion:</i></span></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/Primary_motor_cortex" title="Primary motor cortex">Primary motor cortex</a> → <a href="/wiki/Genu_of_internal_capsule" class="mw-redirect" title="Genu of internal capsule">Genu of internal capsule</a> → <b><a href="/wiki/Corticobulbar_tract" title="Corticobulbar tract">Corticobulbar tract</a></b> → <a href="/wiki/Facial_motor_nucleus" title="Facial motor nucleus">Facial motor nucleus</a> → <a href="/wiki/Facial_muscles" title="Facial muscles">Facial muscles</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>flexion:</i></span></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/Red_nucleus" title="Red nucleus">Red nucleus</a> → <b><a href="/wiki/Rubrospinal_tract" title="Rubrospinal tract">Rubrospinal tract</a></b></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>extension:</i></span></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/Anatomy_of_the_cerebellum#Phylogenetic_and_functional_divisions" title="Anatomy of the cerebellum"> Vestibulocerebellum</a> → <a href="/wiki/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a> → <b><a href="/wiki/Vestibulospinal_tract" title="Vestibulospinal tract">Vestibulospinal tract</a></b></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i>extension:</i></span></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/Anatomy_of_the_cerebellum#Phylogenetic_and_functional_divisions" title="Anatomy of the cerebellum"> Vestibulocerebellum</a> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a> → <b><a href="/wiki/Reticular_formation#Descending_reticulospinal_tracts" title="Reticular formation">Reticulospinal tract</a></b></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Midbrain_tectum" class="mw-redirect" title="Midbrain tectum">Midbrain tectum</a> → <b><a href="/wiki/Tectospinal_tract" title="Tectospinal tract">Tectospinal tract</a></b> → muscles of neck</li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Basal_ganglia" title="Basal ganglia">Basal ganglia</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%"><span class="nobold"><a href="/wiki/Direct_pathway_of_movement" class="mw-redirect" title="Direct pathway of movement"><i>direct:</i></a></span></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">1° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a> → <a href="/wiki/Striatum" title="Striatum">Striatum</a>) → 2° (<a href="/wiki/Medial_globus_pallidus" class="mw-redirect" title="Medial globus pallidus">GPi</a>) → 3° (<a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a>/<a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a> → <a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a> → <a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 4° (<a href="/wiki/Thalamocortical_radiations" title="Thalamocortical radiations">Thalamocortical radiations</a> → <a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor area</a>) → 5° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a>)</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><a href="/wiki/Indirect_pathway_of_movement" class="mw-redirect" title="Indirect pathway of movement"><i>indirect:</i></a></span></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">1° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a> → <a href="/wiki/Striatum" title="Striatum">Striatum</a>) → 2° (<a href="/wiki/Lateral_globus_pallidus" class="mw-redirect" title="Lateral globus pallidus">GPe</a>) → 3° (<a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a> → <a class="mw-selflink selflink">Subthalamic nucleus</a>) → 4° (<a href="/wiki/Subthalamic_fasciculus" title="Subthalamic fasciculus">Subthalamic fasciculus</a> → <a href="/wiki/Medial_globus_pallidus" class="mw-redirect" title="Medial globus pallidus">GPi</a>) → 5° (<a href="/wiki/Lenticular_fasciculus" title="Lenticular fasciculus">Lenticular fasciculus</a>/<a href="/wiki/Ansa_lenticularis" title="Ansa lenticularis">Ansa lenticularis</a> → <a href="/wiki/Thalamic_fasciculus" title="Thalamic fasciculus">Thalamic fasciculus</a> → <a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a> of <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a>) → 6° (<a href="/wiki/Thalamocortical_radiations" title="Thalamocortical radiations">Thalamocortical radiations</a> → <a href="/wiki/Supplementary_motor_area" title="Supplementary motor area">Supplementary motor area</a>) → 7° (<a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a>)</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><span class="nobold"><i><a href="/wiki/Nigrostriatal_pathway" title="Nigrostriatal pathway">nigrostriatal pathway</a>:</i></span></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/Substantia_nigra_pars_compacta" class="mw-redirect" title="Substantia nigra pars compacta">Pars compacta</a> → <a href="/wiki/Striatum" title="Striatum">Striatum</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/Cerebellum" title="Cerebellum">Cerebellar</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%">Afferent</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/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a> → <b><a href="/wiki/Vestibulocerebellar_tract" title="Vestibulocerebellar tract">Vestibulocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellum" title="Cerebellum">Cerebellum</a> → <a href="/wiki/Cerebellum_granule_cell" class="mw-redirect" title="Cerebellum granule cell">Granule cell</a></li></ul> <ul><li><a href="/wiki/Pontine_nuclei" title="Pontine nuclei">Pontine nuclei</a> → <b><a href="/wiki/Pontocerebellar_fibers" title="Pontocerebellar fibers">Pontocerebellar fibers</a></b> → <a href="/wiki/Middle_cerebellar_peduncle" title="Middle cerebellar peduncle">MCP</a> → <a href="/wiki/Deep_cerebellar_nuclei" title="Deep cerebellar nuclei">Deep cerebellar nuclei</a> → <a href="/wiki/Cerebellum_granule_cell" class="mw-redirect" title="Cerebellum granule cell">Granule cell</a></li></ul> <ul><li><a href="/wiki/Inferior_olivary_nucleus" title="Inferior olivary nucleus">Inferior olivary nucleus</a> → <b><a href="/wiki/Olivocerebellar_tract" title="Olivocerebellar tract">Olivocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Hemisphere</a> → <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cell</a> → <a href="/wiki/Deep_cerebellar_nuclei" title="Deep cerebellar nuclei">Deep cerebellar nuclei</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Efferent</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/Dentate_nucleus" title="Dentate nucleus">Dentate nucleus</a> in <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Lateral hemisphere/pontocerebellum</a> → <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <b><a href="/wiki/Dentatothalamic_tract" title="Dentatothalamic tract">Dentatothalamic tract</a></b> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a>) → <a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a></li></ul> <ul><li><a href="/wiki/Interposed_nucleus" title="Interposed nucleus">Interposed nucleus</a> in <a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Intermediate hemisphere/spinocerebellum</a> → <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <a href="/wiki/Reticular_formation" title="Reticular formation">Reticular formation</a>, or → <b><a href="/wiki/Cerebellothalamic_tract" title="Cerebellothalamic tract">Cerebellothalamic tract</a></b> → <a href="/wiki/Red_nucleus" title="Red nucleus">Red nucleus</a> → <a href="/wiki/Thalamus" title="Thalamus">Thalamus</a> (<a href="/wiki/Ventral_lateral_nucleus" title="Ventral lateral nucleus">VL</a>) → <a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a></li></ul> <ul><li><a href="/wiki/Fastigial_nucleus" title="Fastigial nucleus">Fastigial nucleus</a> in <a href="/wiki/Flocculonodular_lobe" title="Flocculonodular lobe">Flocculonodular lobe/vestibulocerebellum</a> → <b><a href="/wiki/Vestibulocerebellar_tract" title="Vestibulocerebellar tract">Vestibulocerebellar tract</a></b> → <a href="/wiki/Vestibular_nuclei" title="Vestibular nuclei">Vestibular nuclei</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Bidirectional:<br /><a href="/wiki/Spinocerebellar_tract" class="mw-redirect" title="Spinocerebellar tract">Spinocerebellar</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/Proprioception#Conscious_and_unconscious_proprioception" title="Proprioception">Unconscious<br /> proprioception</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/Lower_limb" class="mw-redirect" title="Lower limb">lower limb</a> → 1° (<a href="/wiki/Muscle_spindle" title="Muscle spindle">muscle spindles</a> → <a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">DRG</a>) → 2° (<a href="/wiki/Posterior_thoracic_nucleus" title="Posterior thoracic nucleus">Posterior thoracic nucleus</a> → <b><a href="/wiki/Dorsal_spinocerebellar_tract" class="mw-redirect" title="Dorsal spinocerebellar tract">Dorsal/posterior spinocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellar_vermis" title="Cerebellar vermis">Cerebellar vermis</a>)</li></ul> <ul><li><a href="/wiki/Upper_limb" title="Upper limb">upper limb</a> → 1° (<a href="/wiki/Muscle_spindle" title="Muscle spindle">muscle spindles</a> → <a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">DRG</a>) → 2° (<a href="/wiki/Accessory_cuneate_nucleus" title="Accessory cuneate nucleus">Accessory cuneate nucleus</a> → <b><a href="/wiki/Posterior_external_arcuate_fibers" class="mw-redirect" title="Posterior external arcuate fibers">Cuneocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Anterior_lobe_of_cerebellum" title="Anterior lobe of cerebellum">Anterior lobe of cerebellum</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Reflex_arc" title="Reflex arc">Reflex arc</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/Lower_limb" class="mw-redirect" title="Lower limb">lower limb</a> → 1° (<a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organ</a>) → 2° (<b><a href="/wiki/Ventral_spinocerebellar_tract" class="mw-redirect" title="Ventral spinocerebellar tract">Ventral/anterior spinocerebellar tract</a></b>→ <a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">SCP</a> → <a href="/wiki/Cerebellar_vermis" title="Cerebellar vermis">Cerebellar vermis</a>)</li></ul> <ul><li><a href="/wiki/Upper_limb" title="Upper limb">upper limb</a> → 1° (<a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Golgi tendon organ</a>) → 2° (<b><a href="/wiki/Rostral_spinocerebellar_tract" class="mw-redirect" title="Rostral spinocerebellar tract">Rostral spinocerebellar tract</a></b> → <a href="/wiki/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">ICP</a> → <a href="/wiki/Cerebellum" title="Cerebellum">Cerebellum</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" 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