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<li id="toc-Lateral_habenula" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lateral_habenula"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Lateral habenula</span> </div> </a> <ul id="toc-Lateral_habenula-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Medial_habenula" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Medial_habenula"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Medial habenula</span> </div> </a> <ul id="toc-Medial_habenula-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Asymmetry" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Asymmetry"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Asymmetry</span> </div> </a> <ul id="toc-Asymmetry-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Olfactory_coding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Olfactory_coding"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Olfactory coding</span> </div> </a> <ul id="toc-Olfactory_coding-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Functions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Functions"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Functions</span> </div> </a> <ul id="toc-Functions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Depression" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Depression"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Depression</span> </div> </a> <ul id="toc-Depression-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Motivation_and_addiction" class="vector-toc-list-item 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href="https://bs.wikipedia.org/wiki/Mo%C5%BEdana_mamuza" title="Moždana mamuza – Bosnian" lang="bs" hreflang="bs" data-title="Moždana mamuza" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Habenula" title="Habenula – Czech" lang="cs" hreflang="cs" data-title="Habenula" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Habenulae" title="Habenulae – German" lang="de" hreflang="de" data-title="Habenulae" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Hab%C3%A9nula" title="Habénula – Spanish" lang="es" hreflang="es" data-title="Habénula" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Habenula" title="Habenula – French" lang="fr" hreflang="fr" data-title="Habenula" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Abenula" title="Abenula – Italian" lang="it" hreflang="it" data-title="Abenula" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Habenula" title="Habenula – Latin" lang="la" hreflang="la" data-title="Habenula" 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.infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox"><tbody><tr><th colspan="2" class="infobox-above" style="background-color:lemonchiffon">Habenula</th></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Epithalamus.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Epithalamus.png/250px-Epithalamus.png" decoding="async" width="250" height="185" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Epithalamus.png/375px-Epithalamus.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cb/Epithalamus.png/500px-Epithalamus.png 2x" data-file-width="600" data-file-height="443" /></a></span><div class="infobox-caption">Medial aspect of <a href="/wiki/Human_brain" title="Human brain">human brain</a> showing location of the habenula in front of the <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland or body</a> in the <a href="/wiki/Epithalamus" title="Epithalamus">epithalamus</a> shown in red The <a href="/wiki/Habenular_commissure" title="Habenular commissure">habenular commissure</a> is labelled shown connecting the habenula.</div></td></tr><tr><td colspan="2" class="infobox-image"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Brainstem_and_thalamus_ja_ja_135.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Brainstem_and_thalamus_ja_ja_135.png/250px-Brainstem_and_thalamus_ja_ja_135.png" decoding="async" width="250" height="161" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Brainstem_and_thalamus_ja_ja_135.png/375px-Brainstem_and_thalamus_ja_ja_135.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/83/Brainstem_and_thalamus_ja_ja_135.png/500px-Brainstem_and_thalamus_ja_ja_135.png 2x" data-file-width="1000" data-file-height="645" /></a></span><div class="infobox-caption">Habenula shown in blue just in front of the <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland</a> shown in red</div></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/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=D019262">D019262</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=294">294</a></td></tr><tr><th scope="row" class="infobox-label" style="padding-right:0.25em"><a href="/wiki/NeuroLex" title="NeuroLex">NeuroLex</a> ID</th><td class="infobox-data"><a rel="nofollow" class="external text" href="http://uri.neuinfo.org/nif/nifstd/birnlex_1611">birnlex_1611</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/14.1.08.003%20Entity%20TA98%20EN.htm">A14.1.08.003</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=5662">5662</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%2Ffma62032">62032</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/Q267641" class="extiw" title="d:Q267641">edit on Wikidata</a>]</small></div></td></tr></tbody></table> <p>The <b>habenula</b> (diminutive of Latin <span title="Latin-language text"><i lang="la">habena</i></span> meaning <a href="/wiki/Rein" title="Rein">rein</a>) is a small bilateral <a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">neuronal structure</a> in the <a href="/wiki/Brain" title="Brain">brain</a> of vertebrates, that has also been called a microstructure since it is no bigger than a pea. The naming as little rein describes its elongated shape in the <a href="/wiki/Epithalamus" title="Epithalamus">epithalamus</a>, where it borders the <a href="/wiki/Third_ventricle" title="Third ventricle">third ventricle</a>, and lies in front of the <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland</a>.<sup id="cite_ref-Antolin-Fontes_1-0" class="reference"><a href="#cite_note-Antolin-Fontes-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Although it is a microstructure each habenular nucleus is divided into two distinct regions of nuclei, a medial habenula (MHb), and a lateral habenula (LHb) both having different neuronal populations, inputs, and outputs.<sup id="cite_ref-Antunes_2-0" class="reference"><a href="#cite_note-Antunes-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Boulos_3-0" class="reference"><a href="#cite_note-Boulos-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The medial habenula can be subdivided into five subnuclei, the lateral habenula into four subnuclei.<sup id="cite_ref-Ables2023_4-0" class="reference"><a href="#cite_note-Ables2023-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Research has shown morphological complexity in the MHb and LHb. Different inputs to the MHb are discriminated between the different subnuclei.<sup id="cite_ref-Juárez-Leal_5-0" class="reference"><a href="#cite_note-Juárez-Leal-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In the two regions of nuclei there is a difference in <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> giving different functions to each.<sup id="cite_ref-Viswanath_6-0" class="reference"><a href="#cite_note-Viswanath-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The habenula is a conserved structure across vertebrates. In mammals it is highly symmetric, and in <a href="/wiki/Fish" title="Fish">fish</a>, <a href="/wiki/Amphibian" title="Amphibian">amphibians</a> and <a href="/wiki/Reptile" title="Reptile">reptiles</a> it is highly asymmetric in size, molecular composition, and connections.<sup id="cite_ref-Antolin-Fontes_1-1" class="reference"><a href="#cite_note-Antolin-Fontes-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The habenular nuclei are a major component in the <a href="/wiki/Limbic_system" title="Limbic system">limbic system</a> pathways.<sup id="cite_ref-Antolin-Fontes_1-2" class="reference"><a href="#cite_note-Antolin-Fontes-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Fasciculus_retroflexus" title="Fasciculus retroflexus">fasciculus retroflexus</a> pathway between the habenula and the <a href="/wiki/Interpeduncular_nucleus" title="Interpeduncular nucleus">interpeduncular nucleus</a> is one of the first major <a href="/wiki/Nerve_tract" title="Nerve tract">nerve tracts</a> to form in the developing brain.<sup id="cite_ref-Antolin-Fontes_1-3" class="reference"><a href="#cite_note-Antolin-Fontes-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>The habenula is a central structure that connects forebrain regions to midbrain regions, and acts as a hub or node for the integration of emotional and sensory processing.<sup id="cite_ref-Antunes_2-1" class="reference"><a href="#cite_note-Antunes-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> It integrates information from the limbic system, sensory and <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> to guide appropriate and effective responses.<sup id="cite_ref-Juárez-Leal_5-1" class="reference"><a href="#cite_note-Juárez-Leal-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The habenula is involved in the regulation of monoamine neurotransmitters notably dopamine and serotonin.<sup id="cite_ref-Antunes_2-2" class="reference"><a href="#cite_note-Antunes-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Boulos_3-1" class="reference"><a href="#cite_note-Boulos-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Both of these neurotransmitters are strongly associated with anxiety disorders, and avoidance behaviours.<sup id="cite_ref-Antunes_2-3" class="reference"><a href="#cite_note-Antunes-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The functions of the habenula are also involved in motivation, emotion, learning, and <a href="/wiki/Nociception" title="Nociception">pain</a>.<sup id="cite_ref-Antunes_2-4" class="reference"><a href="#cite_note-Antunes-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The MHb plays an important role in depression, stress, memory, and nicotine withdrawal, as well as a role in cocaine, methamphetamine and alcohol addiction.<sup id="cite_ref-Viswanath_6-1" class="reference"><a href="#cite_note-Viswanath-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The MHb shows a high level of <a href="/wiki/Nicotinic_acetylcholine_receptor" title="Nicotinic acetylcholine receptor">nicotinic acetylcholine receptors</a> (nAChRs), that are involved in many forms of <a href="/wiki/Addiction" title="Addiction">addiction</a>. Previously their expression was only noted in other structures associated with addiction. Their expression in the MHb has become a later focus of research.<sup id="cite_ref-Viswanath_6-2" class="reference"><a href="#cite_note-Viswanath-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </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=Habenula&action=edit&section=1" title="Edit section: Anatomy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Each habenular nucleus has two divisions, a medial habenular nucleus (MHb), and a lateral habenular nucleus (LHb). Studies have shown that the medial habenula can be subdivided into five subnuclei, and the lateral habenula into four subnuclei.<sup id="cite_ref-Ables2023_4-1" class="reference"><a href="#cite_note-Ables2023-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The right and left habenular nuclei are connected to each other by the <a href="/wiki/Habenular_commissure" title="Habenular commissure">habenular commissure</a>. The <a href="/wiki/Pineal_gland" title="Pineal gland">pineal gland</a> is attached to the brain in this region.<sup id="cite_ref-Guglielmotti_Cristino_2006_7-0" class="reference"><a href="#cite_note-Guglielmotti_Cristino_2006-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The medial habenula (MHb) receives connections from posterior <a href="/wiki/Septum_pellucidum" title="Septum pellucidum">septum pellucidum</a> and <a href="/wiki/Diagonal_band_of_Broca" title="Diagonal band of Broca">diagonal band of Broca</a>; the lateral habenula receives afferents from the lateral <a href="/wiki/Hypothalamus" title="Hypothalamus">hypothalamus</a>, <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a>, <a href="/wiki/Internal_globus_pallidus" title="Internal globus pallidus">internal globus pallidus</a>, <a href="/wiki/Ventral_pallidum" title="Ventral pallidum">ventral pallidum</a>, and diagonal band of Broca.<sup id="cite_ref-Geisler_8-0" class="reference"><a href="#cite_note-Geisler-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> As a whole, this complexly interconnected region is part of the <a href="/wiki/Dorsal_diencephalic_conduction_system" class="mw-redirect" title="Dorsal diencephalic conduction system">dorsal diencephalic conduction system</a> (DDCS), responsible for relaying information from the limbic system to the <a href="/wiki/Midbrain" title="Midbrain">midbrain</a>, <a href="/wiki/Hindbrain" title="Hindbrain">hindbrain</a>, and <a href="/wiki/Forebrain" title="Forebrain">medial forebrain</a>.<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><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> </p> <div class="mw-heading mw-heading3"><h3 id="Lateral_habenula">Lateral habenula</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=2" title="Edit section: Lateral habenula"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The primary input regions to the lateral habenula (LHb) are the lateral <a href="/wiki/Preoptic_area" title="Preoptic area">preoptic area</a> (bringing input from the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a> and lateral septum), the ventral <a href="/wiki/Pallidum" class="mw-redirect" title="Pallidum">pallidum</a> (bringing input from the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a> and mediodorsal nucleus of the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>), the lateral <a href="/wiki/Hypothalamus" title="Hypothalamus">hypothalamus</a>, the medial habenula, and the internal segment of the <a href="/wiki/Globus_pallidus" title="Globus pallidus">globus pallidus</a> (bringing input from other <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a> structures).<sup id="cite_ref-Geisler_8-1" class="reference"><a href="#cite_note-Geisler-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Neurons in the lateral habenula are 'reward-negative' as they are activated by stimuli associated with unpleasant events, the absence of the reward or the presence of punishment especially when this is unpredictable.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Reward information to the lateral habenula comes from the internal part of the globus pallidus.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>The outputs of the lateral habenula target dopaminergic regions (<a href="/wiki/Substantia_nigra" title="Substantia nigra">substantia nigra</a> <a href="/wiki/Pars_compacta" title="Pars compacta">pars compacta</a> and the <a href="/wiki/Ventral_tegmental_area" title="Ventral tegmental area">ventral tegmental area</a>), serotonergic regions (<a href="/wiki/Median_raphe_nucleus" title="Median raphe nucleus">median raphe</a> and <a href="/wiki/Dorsal_raphe_nucleus" title="Dorsal raphe nucleus">dorsal raphe nuclei</a>), and a cholinergic region (the <a href="/wiki/Laterodorsal_tegmental_nucleus" title="Laterodorsal tegmental nucleus">laterodorsal tegmental nucleus</a>).<sup id="cite_ref-Geisler_8-2" class="reference"><a href="#cite_note-Geisler-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> This output inhibits dopamine neurons in substantia nigra pars compacta and the ventral tegmental area, with activation in the lateral habenula linking to deactivation in them, and vice versa, deactivation in the lateral habenula with their activation.<sup id="cite_ref-Matsumoto_2007_13-0" class="reference"><a href="#cite_note-Matsumoto_2007-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The lateral habenula functions to oppose the action of the laterodorsal tegmental nucleus in the acquisition of <a href="/wiki/Avoidance_response" title="Avoidance response">avoidance responses</a> but not the processing of avoidance later on when it is a memory, motivation or its execution.<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> Research suggests that lateral habenula may play a crucial role in decision making.<sup id="cite_ref-pmid24270185_15-0" class="reference"><a href="#cite_note-pmid24270185-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>There has also shown to be an association with aberrant LHb activity and depression.<sup id="cite_ref-Yang_16-0" class="reference"><a href="#cite_note-Yang-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Medial_habenula">Medial habenula</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=3" title="Edit section: Medial habenula"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The medial habenula (MHb) receives connections from the posterior <a href="/wiki/Septum_pellucidum" title="Septum pellucidum">septum pellucidum</a> and <a href="/wiki/Diagonal_band_of_Broca" title="Diagonal band of Broca">diagonal band of Broca</a>.<sup id="cite_ref-Geisler_8-3" class="reference"><a href="#cite_note-Geisler-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Input to the medial habenula comes from a variety of regions and carries a number of different chemicals. Most neuronal projections to the MHb come from the <a href="/wiki/Septal_area" title="Septal area">septal area</a>.<sup id="cite_ref-Juárez-Leal_5-2" class="reference"><a href="#cite_note-Juárez-Leal-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Input regions include <a href="/wiki/Septal_nuclei" class="mw-redirect" title="Septal nuclei">septal nuclei</a> (the nucleus fimbrialis septi and the nucleus triangularis septi); <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> inputs from the interfascicular nucleus of the ventral tegmental area, noradrenergic inputs from the <a href="/wiki/Locus_ceruleus" class="mw-redirect" title="Locus ceruleus">locus ceruleus</a>, and <a href="/wiki/GABAergic" title="GABAergic">GABAergic</a> inputs from the <a href="/wiki/Diagonal_band_of_Broca" title="Diagonal band of Broca">diagonal band of Broca</a>. The medial habenula sends outputs of <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>, <a href="/wiki/Substance_P" title="Substance P">substance P</a> and <a href="/wiki/Acetylcholine" title="Acetylcholine">acetylcholine</a> to the <a href="/wiki/Periaqueductal_gray" title="Periaqueductal gray">periaqueductal gray</a> via the <a href="/wiki/Interpeduncular_nucleus" title="Interpeduncular nucleus">interpeduncular nucleus</a> as well as to the pineal gland.<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><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> </p> <div class="mw-heading mw-heading3"><h3 id="Asymmetry">Asymmetry</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=4" title="Edit section: Asymmetry"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Asymmetry in the habenula was first noted in 1883 by Nikolaus Goronowitsch.<sup id="cite_ref-Guglielmotti_Cristino_2006_7-1" class="reference"><a href="#cite_note-Guglielmotti_Cristino_2006-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Various species exhibit left-right asymmetric differentiation of habenular neurons. In many <a href="/wiki/Fish" title="Fish">fishes</a> and <a href="/wiki/Amphibian" title="Amphibian">amphibians</a>, the habenula on one side is significantly larger and better organized into distinct nuclei in the dorsal diencephalon than its smaller pair. The sidedness of such differentiation (whether the left or the right is more developed) varies with the species. In <a href="/wiki/Bird" title="Bird">birds</a> and <a href="/wiki/Mammal" title="Mammal">mammals</a>, however, both habenulae are more symmetrical (although not entirely) and consist of a medial and a lateral nucleus on each side which is in fish and amphibians equivalent to dorsal habenula and the ventral habenula, respectively.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Geisler_8-4" class="reference"><a href="#cite_note-Geisler-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1a_20-0" class="reference"><a href="#cite_note-:1a-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Olfactory_coding">Olfactory coding</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=5" title="Edit section: Olfactory coding"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In some <a href="/wiki/Fish" title="Fish">fish</a> (<a href="/wiki/Lamprey" title="Lamprey">lampreys</a> and <a href="/wiki/Teleosts" class="mw-redirect" title="Teleosts">teleosts</a>), <a href="/wiki/Mitral_cell" title="Mitral cell">mitral cell</a> (principal olfactory neurons) axons project exclusively to the right hemisphere of the habenula in an asymmetric manner. It is reported that the dorsal habenula (DHb) are functionally asymmetric with predominantly odor responses in the right hemisphere. It was also shown that DHb neurons are spontaneously active even in the absence of olfactory stimulation. These spontaneously-active DHb neurons are organized into functional clusters which were proposed to govern olfactory responses.<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="Functions">Functions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=6" title="Edit section: Functions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>These nuclei are hypothesized to be involved in regulation of monoamines, such as <a href="/wiki/Dopamine" title="Dopamine">dopamine</a> and <a href="/wiki/Serotonin" title="Serotonin">serotonin</a>.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_23-0" class="reference"><a href="#cite_note-:2-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>The habenular nuclei are involved in pain processing, reproductive behavior, nutrition, <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">sleep-wake cycles</a>, stress responses, and learning. Recent demonstrations using <a href="/wiki/FMRI" class="mw-redirect" title="FMRI">fMRI</a><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> and single unit electrophysiology<sup id="cite_ref-Matsumoto_2007_13-1" class="reference"><a href="#cite_note-Matsumoto_2007-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> have closely linked the function of the lateral habenula with reward processing, in particular with regard to encoding negative feedback or negative rewards. Matsumoto and Hikosaka suggested in 2007 that this reward and reward-negative information in the brain might "be elaborated through the interplay among the lateral habenula, the basal ganglia, and monoaminergic (dopaminergic and serotonergic) systems" and that the lateral habenula may play a pivotal role in this "integrative function".<sup id="cite_ref-Matsumoto_2007_13-2" class="reference"><a href="#cite_note-Matsumoto_2007-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Then, Bromberg-Martin et al. (2011) highlighted that neurons in the lateral habenula signal positive and negative information-prediction errors in addition to positive and negative reward-prediction errors.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Depression">Depression</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=7" title="Edit section: Depression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Both the medial and lateral habenula show reduced volume in those with <a href="/wiki/Major_depressive_disorder" title="Major depressive disorder">depression</a>. Neuron cell numbers were also reduced on the right side.<sup id="cite_ref-Ranft_26-0" class="reference"><a href="#cite_note-Ranft-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Such changes are not seen in those with <a href="/wiki/Schizophrenia" title="Schizophrenia">schizophrenia</a>.<sup id="cite_ref-Ranft_26-1" class="reference"><a href="#cite_note-Ranft-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Deep_brain_stimulation" title="Deep brain stimulation">Deep brain stimulation</a> of the major afferent bundle (i.e., stria medullaris thalami) of the lateral habenula has been used for treatment of depression where it is severe, protracted and therapy-resistant.<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><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> </p><p><a href="/wiki/NMDA_receptor" title="NMDA receptor">N-Methyl-D-aspartate (NMDA) receptor</a>-dependent <a href="/wiki/Bursting" title="Bursting">burst firing</a> in the lateral habenula has been associated with depression in animal studies,<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> and it has been shown that the general anesthetic <a href="/wiki/Ketamine" title="Ketamine">ketamine</a> blocks this firing by acting as a <a href="/wiki/Receptor_antagonist" title="Receptor antagonist">receptor antagonist</a>.<sup id="cite_ref-YangCui2018_30-0" class="reference"><a href="#cite_note-YangCui2018-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Ketamine has been the subject of numerous studies after having shown fast-acting antidepressant effects in humans (in a 0.5 mg/bw kg dose).<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> </p> <div class="mw-heading mw-heading2"><h2 id="Motivation_and_addiction">Motivation and addiction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=8" title="Edit section: Motivation and addiction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Recent exploration of the habenular nuclei has begun to associate the structure with an organism's current mood, feeling of motivation, and reward recognition.<sup id="cite_ref-:3_32-0" class="reference"><a href="#cite_note-:3-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Previously, the LHb has been identified as an "anti-reward" signal, but recent research suggests that the LHb helps identify preference, helping the brain to discriminate between potential actions and subsequent motivation decisions.<sup id="cite_ref-Stopper_33-0" class="reference"><a href="#cite_note-Stopper-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> In a study using a <a href="/wiki/Pavlovian_conditioning" class="mw-redirect" title="Pavlovian conditioning">Pavlovian conditioning</a> model, results showed an increase in the habenula response.<sup id="cite_ref-:5_34-0" class="reference"><a href="#cite_note-:5-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> This increase coincided with conditioned stimuli associated with more aversive punishments (i.e. electric shock).<sup id="cite_ref-:5_34-1" class="reference"><a href="#cite_note-:5-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Therefore, researchers speculate that inhibition or damage to the LHb resulting in a failure to process such information may lead to random motivation behavior.<sup id="cite_ref-Stopper_33-1" class="reference"><a href="#cite_note-Stopper-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_34-2" class="reference"><a href="#cite_note-:5-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>LHb is especially important in understanding the reward and motivation relationship as it relates to addictive behaviors.<sup id="cite_ref-:3_32-1" class="reference"><a href="#cite_note-:3-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> The LHb inhibits <a href="/wiki/Dopaminergic" title="Dopaminergic">dopaminergic</a> neurons, decreasing the release of dopamine.<sup id="cite_ref-:4_35-0" class="reference"><a href="#cite_note-:4-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> It was determined by several animal studies that receiving a reward coincided with elevated dopamine levels, but once the learned association was learned by the animal, dopamine levels remain elevated, only decreasing when the reward is removed.<sup id="cite_ref-:1a_20-1" class="reference"><a href="#cite_note-:1a-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:2_23-1" class="reference"><a href="#cite_note-:2-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:3_32-2" class="reference"><a href="#cite_note-:3-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:4_35-1" class="reference"><a href="#cite_note-:4-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Therefore, dopamine levels only increase with unpredicted rewards and with a "positive prediction error".<sup id="cite_ref-:1a_20-2" class="reference"><a href="#cite_note-:1a-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Moreover, it was determined that removal of an anticipated award activated LHb, inhibited dopamine levels.<sup id="cite_ref-:1a_20-3" class="reference"><a href="#cite_note-:1a-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> This finding helps explain why addictive drugs are associated with elevated dopamine levels.<sup id="cite_ref-:1a_20-4" class="reference"><a href="#cite_note-:1a-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Nicotine_and_nAChRs">Nicotine and nAChRs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=9" title="Edit section: Nicotine and nAChRs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>According to the <a href="/wiki/National_Institute_on_Drug_Abuse" title="National Institute on Drug Abuse">National Institute on Drug Abuse</a>, one in five <a href="/wiki/Preventable_deaths" class="mw-redirect" title="Preventable deaths">preventable deaths</a> in the United States is caused by <a href="/wiki/Tobacco_smoking" title="Tobacco smoking">tobacco</a> use.<sup id="cite_ref-:6_36-0" class="reference"><a href="#cite_note-:6-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Nicotine" title="Nicotine">Nicotine</a> is the addictive drug found in most tobacco products and is easily absorbed by the bloodstream of the body.<sup id="cite_ref-:6_36-1" class="reference"><a href="#cite_note-:6-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Despite common misconceptions regarding the relaxing effects of tobacco and nicotine use, behavioral testing in animals has demonstrated nicotine to have an anxiogenic effect.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Nicotinic_acetylcholine_receptor" title="Nicotinic acetylcholine receptor">Nicotinic acetylcholine receptors</a> (nAChRs) have been identified as the primary site for nicotine activity and regulate consequent cellular polarization.<sup id="cite_ref-:7_38-0" class="reference"><a href="#cite_note-:7-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> nAChRs are made up a number of α and β subunits and are found in both the LHb and MHb, where research suggests they may play a key role in addiction and withdrawal behaviors.<sup id="cite_ref-:7_38-1" class="reference"><a href="#cite_note-:7-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=10" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The habenular is a well conserved structure that appeared in vertebrates more than 360 million years ago.<sup id="cite_ref-Ables2023_4-2" class="reference"><a href="#cite_note-Ables2023-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The habenular commissure was described for the first time in 1555 by <a href="/wiki/Andreas_Vesalius" title="Andreas Vesalius">Andreas Vesalius</a><sup id="cite_ref-Turliuc_Turliuc_Cucu_Dumitrescu_2015_pp._88–100_40-0" class="reference"><a href="#cite_note-Turliuc_Turliuc_Cucu_Dumitrescu_2015_pp._88–100-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> and the habenula nuclei in 1872 by <a href="/wiki/Theodor_Meynert" title="Theodor Meynert">Theodor Hermann Meynert</a>.<sup id="cite_ref-Turliuc_Turliuc_Cucu_Dumitrescu_2016_pp._127–133_41-0" class="reference"><a href="#cite_note-Turliuc_Turliuc_Cucu_Dumitrescu_2016_pp._127–133-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=11" 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-Antolin-Fontes-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Antolin-Fontes_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Antolin-Fontes_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Antolin-Fontes_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Antolin-Fontes_1-3"><sup><i><b>d</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="CITEREFAntolin-FontesAblesGörlichIbañez-Tallon2015" class="citation journal cs1">Antolin-Fontes, B; Ables, JL; Görlich, A; Ibañez-Tallon, I (September 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4452453">"The habenulo-interpeduncular pathway in nicotine aversion and withdrawal"</a>. <i>Neuropharmacology</i>. <b>96</b> (Pt B): 213–22. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuropharm.2014.11.019">10.1016/j.neuropharm.2014.11.019</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4452453">4452453</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/25476971">25476971</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuropharmacology&rft.atitle=The+habenulo-interpeduncular+pathway+in+nicotine+aversion+and+withdrawal.&rft.volume=96&rft.issue=Pt+B&rft.pages=213-22&rft.date=2015-09&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4452453%23id-name%3DPMC&rft_id=info%3Apmid%2F25476971&rft_id=info%3Adoi%2F10.1016%2Fj.neuropharm.2014.11.019&rft.aulast=Antolin-Fontes&rft.aufirst=B&rft.au=Ables%2C+JL&rft.au=G%C3%B6rlich%2C+A&rft.au=Iba%C3%B1ez-Tallon%2C+I&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4452453&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHabenula" class="Z3988"></span></span> </li> <li id="cite_note-Antunes-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Antunes_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Antunes_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Antunes_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Antunes_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Antunes_2-4"><sup><i><b>e</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAntunesCamposMartinsGouveia2023" class="citation journal cs1">Antunes, GF; Campos, ACP; Martins, DO; Gouveia, FV; Rangel Junior, MJ; Pagano, RL; Martinez, RCR (27 June 2023). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10342060">"Unravelling the Role of Habenula Subnuclei on Avoidance Response: Focus on Activation and Neuroinflammation"</a>. <i>International Journal of Molecular Sciences</i>. <b>24</b> (13): 10693. <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/10.3390%2Fijms241310693">10.3390/ijms241310693</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10342060">10342060</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/37445871">37445871</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=International+Journal+of+Molecular+Sciences&rft.atitle=Unravelling+the+Role+of+Habenula+Subnuclei+on+Avoidance+Response%3A+Focus+on+Activation+and+Neuroinflammation.&rft.volume=24&rft.issue=13&rft.pages=10693&rft.date=2023-06-27&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC10342060%23id-name%3DPMC&rft_id=info%3Apmid%2F37445871&rft_id=info%3Adoi%2F10.3390%2Fijms241310693&rft.aulast=Antunes&rft.aufirst=GF&rft.au=Campos%2C+ACP&rft.au=Martins%2C+DO&rft.au=Gouveia%2C+FV&rft.au=Rangel+Junior%2C+MJ&rft.au=Pagano%2C+RL&rft.au=Martinez%2C+RCR&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC10342060&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHabenula" class="Z3988"></span></span> </li> <li id="cite_note-Boulos-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Boulos_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Boulos_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBoulosDarcqKieffer2017" class="citation journal cs1">Boulos, LJ; Darcq, E; Kieffer, BL (15 February 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5143215">"Translating the Habenula-From Rodents to Humans"</a>. <i>Biological Psychiatry</i>. <b>81</b> (4): 296–305. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.biopsych.2016.06.003">10.1016/j.biopsych.2016.06.003</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5143215">5143215</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27527822">27527822</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biological+Psychiatry&rft.atitle=Translating+the+Habenula-From+Rodents+to+Humans.&rft.volume=81&rft.issue=4&rft.pages=296-305&rft.date=2017-02-15&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5143215%23id-name%3DPMC&rft_id=info%3Apmid%2F27527822&rft_id=info%3Adoi%2F10.1016%2Fj.biopsych.2016.06.003&rft.aulast=Boulos&rft.aufirst=LJ&rft.au=Darcq%2C+E&rft.au=Kieffer%2C+BL&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5143215&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHabenula" class="Z3988"></span></span> </li> <li id="cite_note-Ables2023-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Ables2023_4-0"><sup><i><b>a</b></i></sup></a> <a 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class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC11081310">11081310</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/36933754">36933754</a>.</cite><span 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"A review of analogies between some neuroanatomical terms and roman household objects". <i>Annals of Anatomy - Anatomischer Anzeiger</i>. <b>204</b>: 127–133. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.aanat.2015.07.001">10.1016/j.aanat.2015.07.001</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0940-9602">0940-9602</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26337365">26337365</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+of+Anatomy+-+Anatomischer+Anzeiger&rft.atitle=A+review+of+analogies+between+some+neuroanatomical+terms+and+roman+household+objects&rft.volume=204&rft.pages=127-133&rft.date=2016&rft.issn=0940-9602&rft_id=info%3Apmid%2F26337365&rft_id=info%3Adoi%2F10.1016%2Fj.aanat.2015.07.001&rft.aulast=Turliuc&rft.aufirst=Dana&rft.au=Turliuc%2C+%C5%9Eerban&rft.au=Cucu%2C+Andrei&rft.au=Dumitrescu%2C+Gabriela+Floren%C5%A3a&rft.au=C%C4%83r%C4%83uleanu%2C+Alexandru&rft.au=Buzdug%C4%83%2C+C%C4%83t%C4%83lin&rft.au=Tama%C5%9F%2C+Camelia&rft.au=Sava%2C+Anca&rft.au=Costea%2C+Claudia+Florida&rfr_id=info%3Asid%2Fen.wikipedia.org%3AHabenula" class="Z3988"></span></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Habenula&action=edit&section=12" 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="http://brainmaps.org/index.php?q=Habenula">Stained brain slice images which include the "Habenula"</a> at the <a href="/wiki/BrainMaps" title="BrainMaps">BrainMaps project</a></li> <li><a rel="nofollow" class="external text" href="https://www.neuinfo.org/mynif/search.php?q=Habenula&t=data&s=cover&b=0&r=20">NIF Search - Habenula</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160304060146/https://www.neuinfo.org/mynif/search.php?q=Habenula&t=data&s=cover&b=0&r=20">Archived</a> 2016-03-04 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> via the <a href="/wiki/Neuroscience_Information_Framework" title="Neuroscience Information Framework">Neuroscience Information Framework</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080504165606/http://isc.temple.edu/neuroanatomy/lab/atlas/pdhn/">Neuroanatomy lab sectional atlas</a></li></ul> <p><br /> </p> <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 .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist 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.navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .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 class="mw-selflink selflink">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 href="/wiki/Subthalamic_nucleus" title="Subthalamic nucleus">Subthalamic nucleus</a></li> <li><a href="/wiki/Zona_incerta" title="Zona incerta">Zona incerta</a></li> <li><a href="/wiki/Nuclei_campi_perizonalis" class="mw-redirect" title="Nuclei campi perizonalis">Nuclei campi perizonalis</a> (<a href="/wiki/Fields_of_Forel" title="Fields of Forel">Fields of Forel</a>)</li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Smell" 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:Olfactory_system" title="Template:Olfactory system"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Olfactory_system" title="Template talk:Olfactory system"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Olfactory_system" title="Special:EditPage/Template:Olfactory system"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Smell" style="font-size:114%;margin:0 4em"><a href="/wiki/Sense_of_smell" title="Sense of smell">Smell</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Microanatomy</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Olfactory_epithelium" title="Olfactory epithelium">Epithelium</a> <ul><li><a href="/wiki/Olfactory_glands" title="Olfactory glands">glands</a></li> <li><a href="/wiki/Olfactory_mucosa" title="Olfactory mucosa">mucosa</a></li></ul></li> <li><a href="/wiki/Sustentacular_cell" title="Sustentacular cell">Sustentacular cell</a></li> <li><a href="/wiki/Tufted_cell" title="Tufted cell">Tufted cell</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Olfactory_nerve" title="Olfactory nerve">Olfactory nerve</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Olfactory_receptor" title="Olfactory receptor">Olfactory receptor</a> <ul><li><a href="/wiki/Olfactory_receptor_neuron" title="Olfactory receptor neuron">neuron</a></li></ul></li> <li><a href="/wiki/Olfactory_bulb" title="Olfactory bulb">Olfactory bulb</a> <ul><li><a href="/wiki/Glomerulus_(olfaction)" title="Glomerulus (olfaction)">glomeruli</a></li> <li><a href="/wiki/Olfactory_bulb_mitral_cell" class="mw-redirect" title="Olfactory bulb mitral cell">mitral cells</a></li></ul></li> <li><a href="/wiki/Olfactory_tract" title="Olfactory tract">Olfactory tract</a></li> <li><a href="/wiki/Olfactory_trigone" title="Olfactory trigone">Olfactory trigone</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Brain" title="Brain">Brain</a> areas involved in smell</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><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><a href="/wiki/Primary_olfactory_cortex" title="Primary olfactory cortex">Primary olfactory cortex</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Lateral_olfactory_stria" class="mw-redirect" title="Lateral olfactory stria">Lateral olfactory stria</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%">1</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/Piriform_cortex" title="Piriform cortex">Piriform cortex</a></li> <li><a href="/wiki/EC-hippocampus_system" title="EC-hippocampus system">EC-hippocampus system</a> <ul><li><a href="/wiki/Entorhinal_cortex" title="Entorhinal cortex">Entorhinal cortex</a></li> <li><a href="/wiki/Hippocampal_formation" title="Hippocampal formation">Hippocampal formation</a></li></ul></li> <li><a href="/wiki/Prepyriform_area" title="Prepyriform area">Prepyriform area</a></li> <li><a href="/wiki/Periamygdaloid_cortex" title="Periamygdaloid cortex">Periamygdaloid cortex</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">2</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> → <a href="/wiki/Habenular_nuclei" class="mw-redirect" title="Habenular nuclei">Habenular nuclei</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">3</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/Amygdala" title="Amygdala">Amygdala</a> → <a href="/wiki/Stria_terminalis" title="Stria terminalis">Stria terminalis</a> → <a href="/wiki/Hypothalamus" title="Hypothalamus">Hypothalamus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">4</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/Medial_forebrain_bundle" title="Medial forebrain bundle">Medial forebrain bundle</a> → <a href="/wiki/Hypothalamus" title="Hypothalamus">Hypothalamus</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/Medial_olfactory_stria" class="mw-redirect" title="Medial olfactory stria">Medial olfactory stria</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/Anterior_olfactory_nucleus" title="Anterior olfactory nucleus">Anterior olfactory nucleus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">General</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Odor" title="Odor">Odor</a></li> <li><a href="/wiki/Olfactory_system" title="Olfactory system">Olfactory system</a></li> <li><a href="/wiki/Evolution_of_olfaction" title="Evolution of olfaction">Evolution of olfaction</a></li></ul> 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