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Granule cell - Wikipedia

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class="vector-toc-list"> <li id="toc-Cerebellar_granule_cell" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cerebellar_granule_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Cerebellar granule cell</span> </div> </a> <ul id="toc-Cerebellar_granule_cell-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dentate_gyrus_granule_cell" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dentate_gyrus_granule_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Dentate gyrus granule cell</span> </div> </a> <ul id="toc-Dentate_gyrus_granule_cell-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dorsal_cochlear_nucleus_granule_cell" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dorsal_cochlear_nucleus_granule_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Dorsal cochlear nucleus granule cell</span> </div> </a> <ul id="toc-Dorsal_cochlear_nucleus_granule_cell-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Olfactory_bulb_granule_cell" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Olfactory_bulb_granule_cell"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Olfactory bulb granule cell</span> </div> </a> <ul id="toc-Olfactory_bulb_granule_cell-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Function" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Function"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Function</span> </div> </a> <button aria-controls="toc-Function-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Function subsection</span> 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id="toc-Shorter_granule_cell_dendrites-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>External links</span> </div> 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class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Type of neuron with a very small cell body</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">Not to be confused with <a href="/wiki/Granular_cell" class="mw-redirect" title="Granular cell">granular cells</a> in the kidney, or <a href="/wiki/Granulocyte" title="Granulocyte">granulocytes</a>.</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:PurkinjeCell.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/PurkinjeCell.jpg/200px-PurkinjeCell.jpg" decoding="async" width="200" height="234" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/PurkinjeCell.jpg/300px-PurkinjeCell.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/15/PurkinjeCell.jpg/400px-PurkinjeCell.jpg 2x" data-file-width="611" data-file-height="715" /></a><figcaption>Drawing of <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cells</a> (A) and granule cells (B) from pigeon cerebellum by <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a>, 1899. Instituto Santiago Ramón y Cajal, Madrid, Spain.</figcaption></figure> <p>The name <b>granule cell</b> has been used for a number of different types of <a href="/wiki/Neuron" title="Neuron">neurons</a> whose only common feature is that they all have very small <a href="/wiki/Soma_(biology)" title="Soma (biology)">cell bodies</a>. Granule cells are found within the <a href="/wiki/Anatomy_of_the_cerebellum#Granular_layer" title="Anatomy of the cerebellum">granular layer</a> of the <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>, the <a href="/wiki/Dentate_gyrus" title="Dentate gyrus">dentate gyrus</a> of the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, the superficial layer of the <a href="/wiki/Dorsal_cochlear_nucleus" title="Dorsal cochlear nucleus">dorsal cochlear nucleus</a>, the <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a>, and the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a>. </p><p><a href="/wiki/Cerebellar_granule_cell" title="Cerebellar granule cell">Cerebellar granule cells</a> account for the majority of <a href="/wiki/Neuron" title="Neuron">neurons</a> in the human brain.<sup id="cite_ref-Cerebellum_1-0" class="reference"><a href="#cite_note-Cerebellum-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> These granule cells receive excitatory input from <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">mossy fibers</a> originating from <a href="/wiki/Pons" title="Pons">pontine</a> <a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">nuclei</a>. Cerebellar granule cells project up through the <a href="/wiki/Anatomy_of_the_cerebellum#Purkinje_layer" title="Anatomy of the cerebellum">Purkinje layer</a> into the <a href="/wiki/Anatomy_of_the_cerebellum#Molecular_layer" title="Anatomy of the cerebellum">molecular layer</a> where they branch out into parallel fibers that spread through <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cell</a> <a href="/wiki/Dendrite" title="Dendrite">dendritic</a> arbors. These parallel fibers form thousands of excitatory <a href="/wiki/Granule-cell%E2%80%93Purkinje-cell_synapse" title="Granule-cell–Purkinje-cell synapse">granule-cell–Purkinje-cell synapses</a> onto the intermediate and distal dendrites of Purkinje cells using <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> as a <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a>. </p><p>Layer 4 granule cells of the cerebral cortex receive inputs from the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> and send projections to supragranular layers 2–3, but also to infragranular layers of the cerebral cortex. </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=Granule_cell&amp;action=edit&amp;section=1" title="Edit section: Structure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Granule cells in different brain regions are both functionally and anatomically diverse: the only thing they have in common is smallness. For instance, olfactory bulb granule cells are <a href="/wiki/GABAergic" title="GABAergic">GABAergic</a> and axonless, while granule cells in the dentate gyrus have <a href="/wiki/Glutamatergic" title="Glutamatergic">glutamatergic</a> projection <a href="/wiki/Axons" class="mw-redirect" title="Axons">axons</a>. These two populations of granule cells are also the only major neuronal populations that undergo <a href="/wiki/Adult_neurogenesis" title="Adult neurogenesis">adult neurogenesis</a>, while cerebellar and cortical granule cells do not. Granule cells (save for those of the olfactory bulb) have a structure typical of a neuron consisting of <a href="/wiki/Dendrite" title="Dendrite">dendrites</a>, a <a href="/wiki/Soma_(biology)" title="Soma (biology)">soma</a> (cell body) and an axon. </p><p><b>Dendrites</b>: Each granule cell has 3 – 4 stubby dendrites which end in a claw. Each of the dendrites are only about 15 <a href="/wiki/Micrometre" title="Micrometre">μm</a> in length. </p><p><b>Soma</b>: Granule cells all have a small soma diameter of approximately 10 μm. </p><p><b>Axon</b>: Each granule cell sends a single axon onto the Purkinje cell dendritic tree.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="Purkinje cells are part of the cerebellar cortex. It is highly unlikely that all granule cells, like those of the cerebral cortex , the dentate gyrus and the dorsal cochlear nucleus (save for those of the olfactory bulb) would synpapse onto Purkinje cells. (June 2020)">citation needed</span></a></i>&#93;</sup> The axon has an extremely narrow diameter: ½ micrometre. </p><p><b>Synapse</b>: 100–300,000 granule cell axons <a href="/wiki/Synapse" title="Synapse">synapse</a> onto a single <a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cell</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="Purkinje cells are part of the cerebellar cortex. It is highly unlikely that all granule cells, like those of the cerebral cortex , the dentate gyrus and the dorsal cochlear nucleus (save for those of the olfactory bulb) would synpapse onto Purkinje cells. (June 2020)">citation needed</span></a></i>&#93;</sup> </p><p>The existence of <a href="/wiki/Gap_junctions" class="mw-redirect" title="Gap junctions">gap junctions</a> between granule cells allows multiple <a href="/wiki/Neurons" class="mw-redirect" title="Neurons">neurons</a> to be coupled to one another, allowing multiple cells to act in synchrony, and allows signalling functions necessary for granule cell development to occur.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Cerebellar_granule_cell">Cerebellar granule cell</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=2" title="Edit section: Cerebellar granule cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cerebellar_granule_cell" title="Cerebellar granule cell">Cerebellar granule cell</a></div> <p>The granule cells, produced by the <a href="/wiki/Rhombic_lip" title="Rhombic lip">rhombic lip</a>, are found in the granule cell layer of the <a href="/wiki/Cerebellar_cortex" class="mw-redirect" title="Cerebellar cortex">cerebellar cortex</a>. They are small and numerous. They are characterized by a very small soma and several short dendrites which terminate with claw-shaped endings. In the <a href="/wiki/Transmission_electron_microscope" class="mw-redirect" title="Transmission electron microscope">transmission electron microscope</a>, these cells are characterized by a darkly stained nucleus surrounded by a thin rim of cytoplasm. The axon ascends into the molecular layer where it splits to form <a href="/wiki/Parallel_fibers" class="mw-redirect" title="Parallel fibers">parallel fibers</a>.<sup id="cite_ref-Cerebellum_1-1" class="reference"><a href="#cite_note-Cerebellum-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dentate_gyrus_granule_cell">Dentate gyrus granule cell</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=3" title="Edit section: Dentate gyrus granule cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The principal cell type of the <a href="/wiki/Dentate_gyrus" title="Dentate gyrus">dentate gyrus</a> is the granule cell. The dentate gyrus granule cell has an elliptical cell body with a width of approximately 10 μm and a height of 18μm.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> </p><p>The granule cell has a characteristic cone-shaped tree of spiny apical <a href="/wiki/Dendrites" class="mw-redirect" title="Dendrites">dendrites</a>. The dendrite branches project throughout the entire molecular layer, and the furthest tips of the dendritic tree end at the <a href="/wiki/Hippocampal_fissure" class="mw-redirect" title="Hippocampal fissure">hippocampal fissure</a> or at the ventricular surface.<sup id="cite_ref-dentategyrus_4-0" class="reference"><a href="#cite_note-dentategyrus-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> The granule cells are tightly packed in the granular cell layer of the dentate gyrus. </p> <div class="mw-heading mw-heading3"><h3 id="Dorsal_cochlear_nucleus_granule_cell">Dorsal cochlear nucleus granule cell</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=4" title="Edit section: Dorsal cochlear nucleus granule cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The granule cells in the <a href="/wiki/Dorsal_cochlear_nucleus" title="Dorsal cochlear nucleus">dorsal cochlear nucleus</a> are small neurons with two or three short dendrites that give rise to a few branches with expansions at the terminals. The dendrites are short with claw-like endings that form <a href="/wiki/Glomerulus_(cerebellum)" title="Glomerulus (cerebellum)">glomeruli</a> to receive <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">mossy fibers</a>, similar to cerebellar granule cells.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> Its axon projects to the molecular layer of the dorsal cochlear nucleus where it forms parallel fibers, also similar to cerebellar granule cells.<sup id="cite_ref-Cochlear_6-0" class="reference"><a href="#cite_note-Cochlear-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> The dorsal cochlear granule cells are small excitatory <a href="/wiki/Interneurons" class="mw-redirect" title="Interneurons">interneurons</a> which are developmentally related and thus resemble the cerebellar granule cell. </p> <div class="mw-heading mw-heading3"><h3 id="Olfactory_bulb_granule_cell">Olfactory bulb granule cell</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=5" title="Edit section: Olfactory bulb granule cell"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The main intrinsic granule cell in the vertebrate <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a> lacks an axon (as does the accessory neuron). Each cell gives rise to short central dendrites and a single long apical dendrite that expands into the granule cell layer and enters the <a href="/wiki/Mitral_cell" title="Mitral cell">mitral cell</a> body layer. The dendrite branches terminate within the outer plexiform layer among the dendrites in the <a href="/wiki/Olfactory_tract" title="Olfactory tract">olfactory tract</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> In the mammalian olfactory bulb, granule cells can process both synaptic input and output due to the presence of large spines.<sup id="cite_ref-olfactorybulbcalcium_8-0" class="reference"><a href="#cite_note-olfactorybulbcalcium-8"><span class="cite-bracket">&#91;</span>8<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=Granule_cell&amp;action=edit&amp;section=6" title="Edit section: Function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Neural_pathways_and_circuits_of_the_cerebellum">Neural pathways and circuits of the cerebellum</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=7" title="Edit section: Neural pathways and circuits of the cerebellum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Architecture_of_the_Cerebellar_Cortex.svg" class="mw-file-description"><img alt="Cartoon representation of the neural connections that exist between the different types of neurons in the cerebellar cortex. Including Purkinje cells, granule cells and interneurons." src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Architecture_of_the_Cerebellar_Cortex.svg/450px-Architecture_of_the_Cerebellar_Cortex.svg.png" decoding="async" width="450" height="315" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Architecture_of_the_Cerebellar_Cortex.svg/675px-Architecture_of_the_Cerebellar_Cortex.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Architecture_of_the_Cerebellar_Cortex.svg/900px-Architecture_of_the_Cerebellar_Cortex.svg.png 2x" data-file-width="512" data-file-height="358" /></a><figcaption><a href="/wiki/Neural_pathway" title="Neural pathway">Neural pathways</a> and <a href="/wiki/Neural_circuit" title="Neural circuit">circuits</a> in the cerebellum. (+) represent excitatory synapse, while (-) represent inhibitory synapses.</figcaption></figure> <p>Cerebellar granule cells receive excitatory input from 3 or 4 <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">mossy fibers</a> originating from <a href="/wiki/Pontine_nuclei" title="Pontine nuclei">pontine nuclei</a>. Mossy fibers make an excitatory connection onto granule cells, which causes the granule cells to fire an action potential. </p><p>The axon of a cerebellar granule cell splits to form a <a href="/wiki/Parallel_fiber" class="mw-redirect" title="Parallel fiber">parallel fiber</a> which innervates <a href="/wiki/Purkinje_cells" class="mw-redirect" title="Purkinje cells">Purkinje cells</a>. The vast majority of granule cell axonal synapses are found on the parallel fibers.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>The parallel fibers are sent up through the <a href="/wiki/Anatomy_of_the_cerebellum#Purkinje_layer" title="Anatomy of the cerebellum">Purkinje layer</a> into the <a href="/wiki/Anatomy_of_the_cerebellum#Molecular_layer" title="Anatomy of the cerebellum">molecular layer</a> where they branch out and spread through Purkinje cell dendritic arbors. These parallel fibers form thousands of excitatory <a href="/wiki/Granule-cell-Purkinje-cell_synapses" class="mw-redirect" title="Granule-cell-Purkinje-cell synapses">Granule-cell-Purkinje-cell synapses</a> onto the dendrites of Purkinje cells. </p><p>This connection is excitatory as <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> is released. </p><p>The parallel fibers and ascending axon synapses from the same granule cell fire in synchrony which results in excitatory signals. In the cerebellar cortex there are a variety of inhibitory neurons (<a href="/wiki/Interneurons" class="mw-redirect" title="Interneurons">interneurons</a>). The only excitatory neurons present in the cerebellar cortex are granule cells.<sup id="cite_ref-functionofgranulecells_10-0" class="reference"><a href="#cite_note-functionofgranulecells-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p><p>Plasticity of the synapse between a parallel fiber and a Purkinje cell is believed to be important for <a href="/wiki/Motor_learning" title="Motor learning">motor learning</a>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> The function of cerebellar circuits is entirely dependent on processes carried out by the granular layer. Therefore, the function of granule cells determines the cerebellar function as a whole.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Mossy_fiber_input_on_cerebellar_granule_cells">Mossy fiber input on cerebellar granule cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=8" title="Edit section: Mossy fiber input on cerebellar granule cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Granule cell dendrites also synapse with distinctive unmyelinated axons which <a href="/wiki/Santiago_Ram%C3%B3n_y_Cajal" title="Santiago Ramón y Cajal">Santiago Ramón y Cajal</a> called <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">mossy fibers</a><sup id="cite_ref-dentategyrus_4-1" class="reference"><a href="#cite_note-dentategyrus-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> Mossy fibers and <a href="/wiki/Golgi_cell" title="Golgi cell">golgi cells</a> both make synaptic connections with granule cells. Together these cells form the glomeruli.<sup id="cite_ref-functionofgranulecells_10-1" class="reference"><a href="#cite_note-functionofgranulecells-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p><p>Granule cells are subject to <a href="/wiki/Feed_forward_(control)" title="Feed forward (control)">feed-forward</a> inhibition: granule cells excite Purkinje cells but also excite GABAergic interneurons that inhibit Purkinje cells. </p><p>Granule cells are also subject to <a href="/wiki/Feedback_inhibition" class="mw-redirect" title="Feedback inhibition">feedback inhibition</a>: Golgi cells receive excitatory stimuli from granule cells and in turn send back inhibitory signals to the granule cell.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>Mossy fiber input codes are conserved during <a href="/wiki/Synaptic_transmission" class="mw-redirect" title="Synaptic transmission">synaptic transmission</a> between granule cells, suggesting that <a href="/wiki/Innervation" class="mw-redirect" title="Innervation">innervation</a> is specific to the input that is received.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> Granule cells do not just relay signals from mossy fibers, rather they perform various, intricate transformations which are required in the spatiotemporal domain.<sup id="cite_ref-functionofgranulecells_10-2" class="reference"><a href="#cite_note-functionofgranulecells-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p><p>Each granule cell is receiving an input from two different mossy fiber inputs. The input is thus coming from two different places as opposed to the granule cell receiving multiple inputs from the same source. </p><p>The differences in mossy fibers that are sending signals to the granule cells directly effects the type of information that granule cells translate to Purkinje cells. The reliability of this translation will depend on the reliability of <a href="/wiki/Synapse" title="Synapse">synaptic</a> activity in granule cells and on the nature of the stimulus being received.<sup id="cite_ref-mossy_15-0" class="reference"><a href="#cite_note-mossy-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> The signal a granule cell receives from a <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">Mossy fiber</a> depends on the function of the mossy fiber itself. Therefore, granule cells are able to integrate information from the different mossy fibers and generate new patterns of activity.<sup id="cite_ref-mossy_15-1" class="reference"><a href="#cite_note-mossy-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Climbing_fiber_input_on_cerebellar_granule_cells">Climbing fiber input on cerebellar granule cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=9" title="Edit section: Climbing fiber input on cerebellar granule cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Different patterns of mossy fiber input will produce unique patterns of activity in granule cells that can be modified by a teaching signal conveyed by the <a href="/wiki/Climbing_fiber" title="Climbing fiber">climbing fiber</a> input. <a href="/wiki/David_Marr_(neuroscientist)" title="David Marr (neuroscientist)">David Marr</a> and <a href="/wiki/James_Albus" class="mw-redirect" title="James Albus">James Albus</a> suggested that the cerebellum operates as an adaptive filter, altering motor behaviour based on the nature of the sensory input. </p><p>Since multiple (~200,000) granule cells synapse onto a single Purkinje cell, the effects of each parallel fiber can be altered in response to a “teacher signal” from the climbing fiber input. </p> <div class="mw-heading mw-heading3"><h3 id="Specific_functions_of_different_granule_cells">Specific functions of different granule cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=10" title="Edit section: Specific functions of different granule cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <dl><dt>Cerebellum granule cells</dt></dl> <p><a href="/wiki/David_Marr_(neuroscientist)" title="David Marr (neuroscientist)">David Marr</a> suggested that the granule cells encode <i>combinations</i> of mossy fiber inputs. In order for the granule cell to respond, it needs to receive active inputs from multiple mossy fibers. The combination of multiple inputs results in the cerebellum being able to make more precise distinctions between input patterns than a single mossy fiber would allow.<sup id="cite_ref-Marr_16-0" class="reference"><a href="#cite_note-Marr-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> The cerebellar granule cells also play a role in orchestrating the tonic conductances which control sleep in conjunction with the ambient levels of <a href="/wiki/GABA" title="GABA">GABA</a> which are found in the brain. </p> <dl><dt>Dentate granule cells</dt></dl> <p>Loss of dentate gyrus neurons from the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a> results in spatial memory deficits. Therefore, dentate granule cells are thought to function in the formation of spatial memories <sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> and of episodic memories.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> Immature and mature dentate granule cells have distinct roles in memory function. Adult-born granule cells are thought to be involved in pattern separation whereas old granule cells contribute to rapid pattern completion.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> </p> <dl><dt>Dorsal cochlear granule cells</dt></dl> <p>Pyramidal cells from the <a href="/wiki/Primary_auditory_cortex" class="mw-redirect" title="Primary auditory cortex">primary auditory cortex</a> project directly on to the cochlear nucleus. This is important in the acoustic <a href="/wiki/Startle_reflex" class="mw-redirect" title="Startle reflex">startle reflex</a>, in which the pyramidal cells modulate the secondary orientation reflex and the granule cell input is responsible for appropriate orientation.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> This is because the signals received by the granule cells contain information about the head position. Granule cells in the dorsal cochlear nucleus play a role in the perception and response to sounds in our environment. </p> <dl><dt>Olfactory bulb granule cells</dt></dl> <p>Inhibition generated by granule cells, the most common GABAergic cell type in the olfactory bulb, plays a critical role in shaping the output of the olfactory bulb.<sup id="cite_ref-olfactory_21-0" class="reference"><a href="#cite_note-olfactory-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> There are two types of excitatory inputs received by <a href="/wiki/GABAergic" title="GABAergic">GABAergic</a> granule cells; those activated by an <a href="/wiki/AMPA_receptor" title="AMPA receptor">AMPA receptor</a> and those activated by a <a href="/wiki/NMDA_receptor" title="NMDA receptor">NMDA receptor</a>. This allows the granule cells to regulate the processing of the sensory input in the olfactory bulb.<sup id="cite_ref-olfactory_21-1" class="reference"><a href="#cite_note-olfactory-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> The <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a> transmits smell information from the nose to the brain, and is thus necessary for a proper sense of smell. Granule cells in the olfactory bulb have also been found to be important in forming memories linked with scents.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Critical_factors_for_function">Critical factors for function</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=11" title="Edit section: Critical factors for function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <dl><dt>Calcium</dt></dl> <p>Calcium dynamics are essential for several functions of granule cells such as changing <a href="/wiki/Membrane_potential" title="Membrane potential">membrane potential</a>, <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a>, <a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>, and regulation of <a href="/wiki/Gene_transcription" class="mw-redirect" title="Gene transcription">gene transcription</a>.<sup id="cite_ref-functionofgranulecells_10-3" class="reference"><a href="#cite_note-functionofgranulecells-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> The nature of the calcium signals that control the presynaptic and postsynaptic function of the olfactory bulb granule cells spines is mostly unknown.<sup id="cite_ref-olfactorybulbcalcium_8-1" class="reference"><a href="#cite_note-olfactorybulbcalcium-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> </p> <dl><dt>Nitric oxide</dt></dl> <p>Granule neurons have high levels of the neuronal isoform of <a href="/wiki/Nitric_oxide_synthase" title="Nitric oxide synthase">nitric oxide synthase</a>. This enzyme is dependent on the presence of calcium and is responsible for the production of <a href="/wiki/Nitric_oxide" title="Nitric oxide">nitric oxide</a> (NO). This <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a> is a negative regulator of granule cell precursor proliferation which promotes the differentiation of different granule cells. NO regulates interactions between granule cells and <a href="/wiki/Glia" title="Glia">glia</a><sup id="cite_ref-functionofgranulecells_10-4" class="reference"><a href="#cite_note-functionofgranulecells-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> and is essential for protecting the granule cells from damage. NO is also responsible for <a href="/wiki/Neuroplasticity" title="Neuroplasticity">neuroplasticity</a> and <a href="/wiki/Motor_learning" title="Motor learning">motor learning</a>.<sup id="cite_ref-23" class="reference"><a href="#cite_note-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="Role_in_disease">Role in disease</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=12" title="Edit section: Role in disease"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Altered_morphology_of_dentate_granule_cells">Altered morphology of dentate granule cells</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=13" title="Edit section: Altered morphology of dentate granule cells"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/TrkB" class="mw-redirect" title="TrkB">TrkB</a> is responsible for the maintenance of normal synaptic connectivity of the dentate granule cells. TrkB also regulates the specific <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology (biology)</a> of the granule cells and is thus said to be important in regulating neuronal development, neuronal plasticity, learning, and the development of <a href="/wiki/Epilepsy" title="Epilepsy">epilepsy</a>.<sup id="cite_ref-dentateanddisease_24-0" class="reference"><a href="#cite_note-dentateanddisease-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> The TrkB regulation of granule cells is important in preventing memory deficits and limbic epilepsy. This is due to the fact that dentate granule cells play a critical role in the function of the entorhinal-hippocampal circuitry in health and disease. Dentate granule cells are situated to regulate the flow of information into the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>, a structure required for normal learning and memory.<sup id="cite_ref-dentateanddisease_24-1" class="reference"><a href="#cite_note-dentateanddisease-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Decreased_granule_cell_neurogenesis">Decreased granule cell neurogenesis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=14" title="Edit section: Decreased granule cell neurogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Both <a href="/wiki/Epilepsy" title="Epilepsy">epilepsy</a> and <a href="/wiki/Major_depressive_disorder" title="Major depressive disorder">depression</a> show a disrupted production of adult-born hippocampal granule cells.<sup id="cite_ref-epilepsy_25-0" class="reference"><a href="#cite_note-epilepsy-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> Epilepsy is associated with increased production - but aberrant integration - of new cells early in the disease and decreased production late in the disease.<sup id="cite_ref-epilepsy_25-1" class="reference"><a href="#cite_note-epilepsy-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> Aberrant integration of adult-generated cells during the development of epilepsy may impair the ability of the dentate gyrus to prevent excess excitatory activity from reaching hippocampal <a href="/wiki/Pyramidal_cells" class="mw-redirect" title="Pyramidal cells">pyramidal cells</a>, thereby promoting seizures.<sup id="cite_ref-epilepsy_25-2" class="reference"><a href="#cite_note-epilepsy-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> Long-lasting epileptic seizure stimulate dentate granule cell neurogenesis. These newly born dentate granule cells may result in aberrant connections that result in the hippocampal network plasticity associated with <a href="/wiki/Epileptogenesis" title="Epileptogenesis">epileptogenesis</a>.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Shorter_granule_cell_dendrites">Shorter granule cell dendrites</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=15" title="Edit section: Shorter granule cell dendrites"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Patients with <a href="/wiki/Alzheimer%27s" class="mw-redirect" title="Alzheimer&#39;s">Alzheimer's</a> have shorter granule cell dendrites. Furthermore, the dendrites were less branched and had fewer spines than those in patients not with Alzheimer's.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> However, granule cell dendrites are not an essential component of <a href="/wiki/Senile_plaques" class="mw-redirect" title="Senile plaques">senile plaques</a> and these plaques have no direct effect on granule cells in the dentate gyrus. The specific neurofibrillary changes of dentate granule cells occur in patients with Alzheimer's, <a href="/wiki/Lewy_body" title="Lewy body">Lewy body</a> variant and <a href="/wiki/Progressive_supranuclear_palsy" title="Progressive supranuclear palsy">progressive supranuclear palsy</a>.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=16" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/List_of_distinct_cell_types_in_the_adult_human_body" class="mw-redirect" title="List of distinct cell types in the adult human body">List of distinct cell types in the adult human body</a> </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=Granule_cell&amp;action=edit&amp;section=17" 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-Cerebellum-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Cerebellum_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Cerebellum_1-1"><sup><i><b>b</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="CITEREFLlinásWaltonLang2004" class="citation book cs1">Llinás, Rodolfo; Walton, Kerry; Lang, Eric (2004). "Chapter 7: Cerebellum". In Shepherd, Gordon (ed.). <i>The synaptic organization of the brain</i>. Oxford, New York: Oxford University Press. pp.&#160;271–309. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/9780195159561" title="Special:BookSources/9780195159561"><bdi>9780195159561</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=bookitem&amp;rft.atitle=Chapter+7%3A+Cerebellum&amp;rft.btitle=The+synaptic+organization+of+the+brain&amp;rft.place=Oxford%2C+New+York&amp;rft.pages=271-309&amp;rft.pub=Oxford+University+Press&amp;rft.date=2004&amp;rft.isbn=9780195159561&amp;rft.aulast=Llin%C3%A1s&amp;rft.aufirst=Rodolfo&amp;rft.au=Walton%2C+Kerry&amp;rft.au=Lang%2C+Eric&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AGranule+cell" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFC._ReyherJ_LiibkeW_LarsenG_Hendrix1991" class="citation journal cs1">C. Reyher; J Liibke; W Larsen; G Hendrix; M Shipley; H Baumgarten (1991). <a rel="nofollow" class="external text" href="http://www.jneurosci.org/content/11/6/1485.full.pdf">"Olfactory Bulb Granule Cell Aggregates: Morphological Evidence for lnterperikaryal&#32;&#91;<i>sic</i>&#93; Electrotonic Coupling via Gap Junctions"</a> <span class="cs1-format">(PDF)</span>. <i><a href="/wiki/The_Journal_of_Neuroscience" title="The Journal of Neuroscience">The Journal of Neuroscience</a></i>. <b>11</b> (6): 1465–495. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1523%2FJNEUROSCI.11-06-01485.1991">10.1523/JNEUROSCI.11-06-01485.1991</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&#160;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6575397">6575397</a></span>. <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/1904478">1904478</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210814000347/https://www.jneurosci.org/content/jneuro/11/6/1485.full.pdf">Archived</a> <span class="cs1-format">(PDF)</span> from the original on 14 August 2021<span class="reference-accessdate">. Retrieved <span class="nowrap">3 February</span> 2016</span>.</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=The+Journal+of+Neuroscience&amp;rft.atitle=Olfactory+Bulb+Granule+Cell+Aggregates%3A+Morphological+Evidence+for+lnterperikaryal%26%2332%3B%26%2391%3Bsic%26%2393%3B+Electrotonic+Coupling+via+Gap+Junctions&amp;rft.volume=11&amp;rft.issue=6&amp;rft.pages=1465-495&amp;rft.date=1991&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6575397%23id-name%3DPMC&amp;rft_id=info%3Apmid%2F1904478&amp;rft_id=info%3Adoi%2F10.1523%2FJNEUROSCI.11-06-01485.1991&amp;rft.au=C.+Reyher&amp;rft.au=J+Liibke&amp;rft.au=W+Larsen&amp;rft.au=G+Hendrix&amp;rft.au=M+Shipley&amp;rft.au=H+Baumgarten&amp;rft_id=http%3A%2F%2Fwww.jneurosci.org%2Fcontent%2F11%2F6%2F1485.full.pdf&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AGranule+cell" class="Z3988"></span></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFClaiborneAmaralCowan1990" class="citation journal cs1">Claiborne BJ, Amaral DG, Cowan WM (1990). 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href="/w/index.php?title=Granule_cell&amp;action=edit&amp;section=18" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://archive.today/20130414114629/http://www.biolcell.org/boc/099/0363/boc0990363f01.htm?resolution=HIGH/">Position of granule cells within layers of the cerebella cortex</a></li> <li><a rel="nofollow" class="external text" href="http://www.nature.com/neuro/journal/v1/n1/fig_tab/nn0598_8_F1.html">Architecture of the Cerebellum</a></li> <li><a rel="nofollow" class="external text" href="http://grants.hhp.uh.edu/clayne/6397/Unit5_files/cerebell.htm">The Cerebellum</a></li> <li><a rel="nofollow" class="external text" href="http://brainmaps.org/index.php?q=Granule%20cells">Stained brain slice images which include the "Granule cells"</a> at the <a href="/wiki/BrainMaps" title="BrainMaps">BrainMaps project</a></li></ul> <div class="navbox-styles"><style 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href="/wiki/Central_nervous_system" title="Central nervous system">CNS</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Tissue Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Grey_matter" title="Grey matter">Grey matter</a></li> <li><a href="/wiki/White_matter" title="White matter">White matter</a> <ul><li><a href="/wiki/Projection_fiber" title="Projection fiber">Projection fibers</a></li> <li><a href="/wiki/Association_fiber" title="Association fiber">Association fiber</a></li> <li><a href="/wiki/Commissural_fiber" title="Commissural fiber">Commissural fiber</a></li> <li><a href="/wiki/Lemniscus_(anatomy)" title="Lemniscus (anatomy)">Lemniscus</a></li> <li><a href="/wiki/Nerve_tract" title="Nerve tract">Nerve tract</a></li> <li><a href="/wiki/Decussation" title="Decussation">Decussation</a></li></ul></li> <li><a href="/wiki/Neuropil" title="Neuropil">Neuropil</a></li> <li><a href="/wiki/Meninges" title="Meninges">Meninges</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cell Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neuron" title="Neuron">Neuronal</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Pyramidal_cell" title="Pyramidal cell">Pyramidal</a></li> <li><a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje</a></li> <li><a class="mw-selflink selflink">Granule</a></li> <li><a href="/wiki/Von_Economo_neuron" title="Von Economo neuron">Von Economo</a></li> <li><a href="/wiki/Medium_spiny_neuron" title="Medium spiny neuron">Medium spiny</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Glial</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Astrocyte" title="Astrocyte">Astrocyte</a></li> <li><a href="/wiki/Ependyma" title="Ependyma">Ependymal cells</a> <ul><li><a href="/wiki/Tanycyte" title="Tanycyte">Tanycyte</a></li></ul></li> <li><a href="/wiki/Oligodendrocyte_progenitor_cell" title="Oligodendrocyte progenitor cell">Oligodendrocyte progenitor cell</a></li> <li><a href="/wiki/Oligodendrocyte" title="Oligodendrocyte">Oligodendrocyte</a></li> <li><a href="/wiki/Microglia" title="Microglia">Microglia</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Peripheral_nervous_system" title="Peripheral nervous system">PNS</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i>Dorsal</i> <ul><li><a href="/wiki/Dorsal_root_of_spinal_nerve" title="Dorsal root of spinal nerve">Root</a></li> <li><a href="/wiki/Dorsal_root_ganglion" title="Dorsal root ganglion">Ganglion</a></li> <li><a href="/wiki/Dorsal_ramus_of_spinal_nerve" title="Dorsal ramus of spinal nerve">Ramus</a></li></ul></li> <li><i>Ventral</i> <ul><li><a href="/wiki/Ventral_root_of_spinal_nerve" title="Ventral root of spinal nerve">Root</a></li> <li><a href="/wiki/Ventral_ramus_of_spinal_nerve" title="Ventral ramus of spinal nerve">Ramus</a></li></ul></li> <li><i><a href="/wiki/Ramus_communicans" title="Ramus communicans">Ramus communicans</a></i> <ul><li><a href="/wiki/Gray_ramus_communicans" title="Gray ramus communicans">Gray</a></li> <li><a href="/wiki/White_ramus_communicans" title="White ramus communicans">White</a></li></ul></li> <li><a href="/wiki/Autonomic_ganglion" title="Autonomic ganglion">Autonomic ganglion</a> (<a href="/wiki/Preganglionic_nerve_fibers" title="Preganglionic nerve fibers">Preganglionic nerve fibers</a></li> <li><a href="/wiki/Postganglionic_nerve_fibers" title="Postganglionic nerve fibers">Postganglionic nerve fibers</a>)</li> <li><a href="/wiki/Nerve_fascicle" title="Nerve fascicle">Nerve fascicle</a></li> <li><a href="/wiki/Funiculus_(neuroanatomy)" title="Funiculus (neuroanatomy)">Funiculus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissues</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epineurium" title="Epineurium">Epineurium</a></li> <li><a href="/wiki/Perineurium" title="Perineurium">Perineurium</a></li> <li><a href="/wiki/Endoneurium" title="Endoneurium">Endoneurium</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Glia" title="Glia">Neuroglia</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Myelin" title="Myelin">Myelination</a>: <a href="/wiki/Schwann_cell" title="Schwann cell">Schwann cell</a> <ul><li><a href="/wiki/Neurilemma" title="Neurilemma">Neurilemma</a></li> <li><a href="/wiki/Myelin_incisure" title="Myelin incisure">Myelin incisure</a></li> <li><a href="/wiki/Node_of_Ranvier" title="Node of Ranvier">Node of Ranvier</a></li> <li><a href="/wiki/Internodal_segment" title="Internodal segment">Internodal segment</a></li></ul></li> <li><a href="/wiki/Satellite_glial_cell" title="Satellite glial cell">Satellite glial cell</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neuron" title="Neuron">Neurons</a>/<br /><a href="/wiki/Nerve_fiber" class="mw-redirect" title="Nerve fiber">nerve fibers</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Parts</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Soma_(biology)" title="Soma (biology)">Soma</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Axon_hillock" title="Axon hillock">Axon hillock</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Axon" title="Axon">Axon</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Telodendron" class="mw-redirect" title="Telodendron">Telodendron</a></li> <li><a href="/wiki/Axon_terminal" title="Axon terminal">Axon terminals</a></li> <li><a href="/wiki/Axoplasm" title="Axoplasm">Axoplasm</a></li> <li><a href="/wiki/Axolemma" title="Axolemma">Axolemma</a></li> <li><a href="/wiki/Neurofilament" title="Neurofilament">Neurofibril/neurofilament</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dendrite" title="Dendrite">Dendrite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><ul><li><a href="/wiki/Nissl_body" title="Nissl body">Nissl body</a></li> <li><a href="/wiki/Dendritic_spine" title="Dendritic spine">Dendritic spine</a></li> <li><a href="/wiki/Apical_dendrite" title="Apical dendrite">Apical dendrite</a>/<a href="/wiki/Basal_dendrite" title="Basal dendrite">Basal dendrite</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bipolar_neuron" title="Bipolar neuron">Bipolar</a></li> <li><a href="/wiki/Unipolar_neuron" title="Unipolar neuron">Unipolar</a></li> <li><a href="/wiki/Pseudounipolar_neuron" title="Pseudounipolar neuron">Pseudounipolar</a></li> <li><a href="/wiki/Multipolar_neuron" title="Multipolar neuron">Multipolar</a></li> <li><a href="/wiki/Interneuron" title="Interneuron">Interneuron</a> <ul><li><a href="/wiki/Renshaw_cell" title="Renshaw cell">Renshaw</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Afferent_nerve_fiber" title="Afferent nerve fiber">Afferent nerve fiber</a>/<br /><a href="/wiki/Sensory_neuron" title="Sensory neuron">Sensory neuron</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/General_somatic_afferent_fibers" class="mw-redirect" title="General somatic afferent fibers">GSA</a></li> <li><a href="/wiki/General_visceral_afferent_fibers" class="mw-redirect" title="General visceral afferent fibers">GVA</a></li> <li><a href="/wiki/Special_somatic_afferent" class="mw-redirect" title="Special somatic afferent">SSA</a></li> <li><a href="/wiki/Special_visceral_afferent" class="mw-redirect" title="Special visceral afferent">SVA</a></li> <li><a href="/wiki/Axon#types" title="Axon">fibers</a> <ul><li><a href="/wiki/Type_Ia_sensory_fiber" title="Type Ia sensory fiber">Ia or Aα</a></li> <li><a href="/wiki/Golgi_tendon_organ" title="Golgi tendon organ">Ib or Golgi or Aα</a></li> <li><a href="/wiki/Type_II_sensory_fiber" title="Type II sensory fiber">II or Aβ and Aγ</a></li> <li><a href="/wiki/Group_A_nerve_fiber" title="Group A nerve fiber">III or Aδ or fast pain</a></li> <li><a href="/wiki/Group_C_nerve_fiber" title="Group C nerve fiber">IV or C or slow pain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Efferent_nerve_fiber" title="Efferent nerve fiber">Efferent nerve fiber</a>/<br /><a href="/wiki/Motor_neuron" title="Motor neuron">Motor neuron</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/General_somatic_efferent_fibers" class="mw-redirect" title="General somatic efferent fibers">GSE</a></li> <li><a href="/wiki/General_visceral_efferent_fibers" class="mw-redirect" title="General visceral efferent fibers">GVE</a></li> <li><a href="/wiki/Special_visceral_efferent" class="mw-redirect" title="Special visceral efferent">SVE</a></li> <li><a href="/wiki/Upper_motor_neuron" title="Upper motor neuron">Upper motor neuron</a></li> <li><a href="/wiki/Lower_motor_neuron" title="Lower motor neuron">Lower motor neuron</a> <ul><li><a href="/wiki/Alpha_motor_neuron" title="Alpha motor neuron">α motorneuron</a></li> <li><a href="/wiki/Beta_motor_neuron" title="Beta motor neuron">β motorneuron</a></li> <li><a href="/wiki/Gamma_motor_neuron" title="Gamma motor neuron">γ motorneuron</a></li></ul></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Termination</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Synapse" title="Synapse">Synapse</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Electrical_synapse" title="Electrical synapse">Electrical synapse</a>/<a href="/wiki/Gap_junction" title="Gap junction">Gap junction</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a> <ul><li><a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">Synaptic vesicle</a></li> <li><a href="/wiki/Active_zone" title="Active zone">Active zone</a></li> <li><a href="/wiki/Postsynaptic_density" title="Postsynaptic density">Postsynaptic density</a></li></ul></li> <li><a href="/wiki/Autapse" title="Autapse">Autapse</a></li> <li><a href="/wiki/Ribbon_synapse" title="Ribbon synapse">Ribbon synapse</a></li> <li><a href="/wiki/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sensory_receptor" class="mw-redirect" title="Sensory receptor">Sensory receptors</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Tactile_corpuscle" title="Tactile corpuscle">Meissner's corpuscle</a></li> <li><a href="/wiki/Merkel_nerve_ending" title="Merkel nerve ending">Merkel nerve ending</a></li> <li><a href="/wiki/Pacinian_corpuscle" title="Pacinian corpuscle">Pacinian corpuscle</a></li> <li><a href="/wiki/Bulbous_corpuscle" title="Bulbous corpuscle">Ruffini ending</a></li> <li><a href="/wiki/Muscle_spindle" title="Muscle spindle">Muscle spindle</a></li> <li><a href="/wiki/Free_nerve_ending" title="Free nerve ending">Free nerve ending</a></li> <li><a href="/wiki/Nociceptor" title="Nociceptor">Nociceptor</a></li> <li><a href="/wiki/Olfactory_receptor_neuron" title="Olfactory receptor neuron">Olfactory receptor neuron</a></li> <li><a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">Photoreceptor cell</a></li> <li><a href="/wiki/Hair_cell" title="Hair cell">Hair cell</a></li> <li><a href="/wiki/Taste_receptor" title="Taste receptor">Taste receptor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <div 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_cerebellum" 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:Cerebellum" title="Template:Cerebellum"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cerebellum" title="Template talk:Cerebellum"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cerebellum" title="Special:EditPage/Template:Cerebellum"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Anatomy_of_the_cerebellum" style="font-size:114%;margin:0 4em"><a href="/wiki/Anatomy_of_the_cerebellum" title="Anatomy of the cerebellum">Anatomy</a> of the <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Surface</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%">Lobes</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_lobe_of_cerebellum" title="Anterior lobe of cerebellum">Anterior lobe</a></li> <li><a href="/wiki/Posterior_lobe_of_cerebellum" title="Posterior lobe of cerebellum">Posterior lobe</a> <ul><li><a href="/wiki/Horizontal_fissure_of_cerebellum" title="Horizontal fissure of cerebellum">Horizontal fissure</a></li></ul></li> <li><a href="/wiki/Flocculonodular_lobe" title="Flocculonodular lobe">Flocculonodular lobe</a> <ul><li><a href="/wiki/Flocculus_(cerebellar)" class="mw-redirect" title="Flocculus (cerebellar)">Flocculus</a></li> <li><a href="/wiki/Nodule_of_vermis" title="Nodule of vermis">Nodule</a></li></ul></li> <li><a href="/wiki/Primary_fissure_of_cerebellum" title="Primary fissure of cerebellum">Primary fissure</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Medial/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><a href="/wiki/Cerebellar_vermis" title="Cerebellar vermis">Vermis</a>: <i>anterior</i> <ul><li><a href="/wiki/Central_lobule" title="Central lobule">Central lobule</a></li> <li><a href="/wiki/Culmen_(cerebellum)" title="Culmen (cerebellum)">Culmen</a></li> <li><a href="/wiki/Lingula_of_cerebellum" title="Lingula of cerebellum">Lingula</a></li></ul></li> <li><i>posterior</i> <ul><li><a href="/wiki/Folium_vermis" title="Folium vermis">Folium</a></li> <li><a href="/wiki/Tuber_of_vermis" title="Tuber of vermis">Tuber</a></li> <li><a href="/wiki/Uvula_of_cerebellum" title="Uvula of cerebellum">Uvula</a></li></ul></li> <li><a href="/wiki/Vallecula_of_cerebellum" title="Vallecula of cerebellum">Vallecula of cerebellum</a></li></ul> <ul><li><a href="/wiki/Cerebellar_hemisphere" title="Cerebellar hemisphere">Hemisphere</a>: <i>anterior</i> <ul><li><a href="/wiki/Alar_central_lobule" class="mw-redirect" title="Alar central lobule">Alar central lobule</a></li></ul></li> <li><i>posterior</i> <ul><li><a href="/wiki/Biventer_lobule" title="Biventer lobule">Biventer lobule</a></li> <li><a href="/wiki/Cerebellar_tonsil" title="Cerebellar tonsil">Cerebellar tonsil</a></li></ul></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 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/Deep_cerebellar_nuclei" title="Deep cerebellar nuclei">Deep cerebellar nuclei</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/Dentate_nucleus" title="Dentate nucleus">Dentate</a></li> <li><i><a href="/wiki/Interposed_nucleus" title="Interposed nucleus">interposed</a></i> <ul><li><a href="/wiki/Emboliform_nucleus" title="Emboliform nucleus">Emboliform</a></li> <li><a href="/wiki/Globose_nucleus" title="Globose nucleus">Globose</a></li></ul></li> <li><a href="/wiki/Fastigial_nucleus" title="Fastigial nucleus">Fastigial</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cerebellum" title="Cerebellum">Cerebellar cortex</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>Molecular layer</i> <ul><li><a href="/wiki/Stellate_cell" title="Stellate cell">Stellate cell</a></li> <li><a href="/wiki/Basket_cell" title="Basket cell">Basket cell</a></li></ul></li></ul> <ul><li><i>Purkinje cell layer</i> <ul><li><a href="/wiki/Purkinje_cell" title="Purkinje cell">Purkinje cell</a></li> <li><a href="/wiki/Astrocyte#Bergmann_glia" title="Astrocyte">Bergmann glia cell = Golgi epithelial cell</a></li> <li><a href="/wiki/Fa%C3%B1anas_cell" class="mw-redirect" title="Fañanas cell">Fañanas cell</a></li></ul></li></ul> <ul><li><i>Granule cell layer</i> <ul><li><a href="/wiki/Golgi_cell" title="Golgi cell">Golgi cell</a></li> <li><a class="mw-selflink selflink">Granule cell</a></li> <li><a href="/wiki/Unipolar_brush_cell" title="Unipolar brush cell">Unipolar brush cell</a></li></ul></li></ul> <ul><li>Fibers: <a href="/wiki/Mossy_fiber_(cerebellum)" title="Mossy fiber (cerebellum)">Mossy fibers</a></li> <li><a href="/wiki/Climbing_fiber" title="Climbing fiber">Climbing fiber</a></li> <li><a href="/wiki/Parallel_fiber" class="mw-redirect" title="Parallel fiber">Parallel fiber</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-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%">Internal</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/Arbor_vitae_(anatomy)" title="Arbor vitae (anatomy)">Arbor vitae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cerebellar_peduncle" class="mw-redirect" title="Cerebellar peduncle">Peduncles</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/Inferior_cerebellar_peduncle" title="Inferior cerebellar peduncle">Inferior</a> (medulla): <a href="/wiki/Dorsal_spinocerebellar_tract" class="mw-redirect" title="Dorsal spinocerebellar tract">Dorsal spinocerebellar tract</a></li> <li><a href="/wiki/Olivocerebellar_tract" title="Olivocerebellar tract">Olivocerebellar tract</a></li> <li><a href="/wiki/Posterior_external_arcuate_fibers" class="mw-redirect" title="Posterior external arcuate fibers">Cuneocerebellar tract</a></li> <li><a href="/wiki/Juxtarestiform_body" title="Juxtarestiform body">Juxtarestiform body</a> (<a href="/wiki/Vestibulocerebellar_tract" title="Vestibulocerebellar tract">Vestibulocerebellar tract</a>)</li> <li><a href="/wiki/Trigeminocerebellar_fibers" title="Trigeminocerebellar fibers">Trigeminocerebellar fibers</a></li></ul> <ul><li><a href="/wiki/Middle_cerebellar_peduncle" title="Middle cerebellar peduncle">Middle</a> (pons): <a href="/wiki/Pontocerebellar_fibers" title="Pontocerebellar fibers">Pontocerebellar fibers</a></li></ul> <ul><li><a href="/wiki/Superior_cerebellar_peduncle" title="Superior cerebellar peduncle">Superior</a> (midbrain): <a href="/wiki/Ventral_spinocerebellar_tract" class="mw-redirect" title="Ventral spinocerebellar tract">Ventral spinocerebellar tract</a></li> <li><a href="/wiki/Dentatothalamic_tract" title="Dentatothalamic tract">Dentatothalamic tract</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐kmvjg Cached time: 20241122165002 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, 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