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<span>Synthesis</span> </div> </a> <ul id="toc-Synthesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanism" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mechanism"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Mechanism</span> </div> </a> <ul id="toc-Mechanism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Receptor_targets" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Receptor_targets"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Receptor targets</span> </div> </a> <ul id="toc-Receptor_targets-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Actions" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Actions"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Actions</span> </div> </a> <button aria-controls="toc-Actions-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 Actions subsection</span> </button> <ul id="toc-Actions-sublist" class="vector-toc-list"> <li id="toc-Co-release" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Co-release"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Co-release</span> </div> </a> <ul id="toc-Co-release-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Evolution_of_Neuropeptide_Signaling" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Evolution_of_Neuropeptide_Signaling"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Evolution of Neuropeptide Signaling</span> </div> </a> <ul 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Available in 26 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-26" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">26 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A8%D8%A8%D8%AA%D9%8A%D8%AF_%D8%B9%D8%B5%D8%A8%D9%8A" title="ببتيد عصبي – Arabic" lang="ar" hreflang="ar" data-title="ببتيد عصبي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Neuropeptid" title="Neuropeptid – Bosnian" lang="bs" hreflang="bs" data-title="Neuropeptid" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Neurop%C3%A8ptid" title="Neuropèptid – Catalan" lang="ca" hreflang="ca" data-title="Neuropèptid" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Neuropeptid" title="Neuropeptid – Czech" lang="cs" hreflang="cs" data-title="Neuropeptid" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Neuropeptid" title="Neuropeptid – Danish" lang="da" hreflang="da" data-title="Neuropeptid" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Neuropeptide" title="Neuropeptide – German" lang="de" hreflang="de" data-title="Neuropeptide" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Neuropeptiidid" title="Neuropeptiidid – Estonian" lang="et" hreflang="et" data-title="Neuropeptiidid" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Neurop%C3%A9ptido" title="Neuropéptido – Spanish" lang="es" hreflang="es" data-title="Neuropéptido" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Ne%C5%ADrona_peptido" title="Neŭrona peptido – Esperanto" lang="eo" hreflang="eo" data-title="Neŭrona peptido" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%86%D9%88%D8%B1%D9%88%D9%BE%D9%BE%D8%AA%DB%8C%D8%AF" title="نوروپپتید – Persian" lang="fa" hreflang="fa" data-title="نوروپپتید" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Neuropeptide" title="Neuropeptide – French" lang="fr" hreflang="fr" data-title="Neuropeptide" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Neurop%C3%A9ptido" title="Neuropéptido – Galician" lang="gl" hreflang="gl" data-title="Neuropéptido" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Neuropeptidi" title="Neuropeptidi – Croatian" lang="hr" hreflang="hr" data-title="Neuropeptidi" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Neuropeptide" title="Neuropeptide – Italian" lang="it" hreflang="it" data-title="Neuropeptide" 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/Neuropeptidis" title="Neuropeptidis – Latin" lang="la" hreflang="la" data-title="Neuropeptidis" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Neuropeptydy" title="Neuropeptydy – Polish" lang="pl" hreflang="pl" data-title="Neuropeptydy" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9D%D0%B5%D0%B9%D1%80%D0%BE%D0%BF%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D1%8B" title="Нейропептиды – Russian" lang="ru" hreflang="ru" data-title="Нейропептиды" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Neuropeptide" title="Neuropeptide – Simple English" lang="en-simple" hreflang="en-simple" data-title="Neuropeptide" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AF%DB%95%D9%85%D8%A7%D8%B1%DB%95%D9%BE%DB%8E%D9%BE%D8%AA%D8%A7%DB%8C%D8%AF" title="دەمارەپێپتاید – Central Kurdish" lang="ckb" hreflang="ckb" data-title="دەمارەپێپتاید" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/Neuropeptidi" title="Neuropeptidi – Serbian" lang="sr" hreflang="sr" data-title="Neuropeptidi" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Neuropeptid" title="Neuropeptid – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Neuropeptid" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Neuropeptidi" title="Neuropeptidi – Finnish" lang="fi" hreflang="fi" data-title="Neuropeptidi" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Neuropeptider" title="Neuropeptider – Swedish" lang="sv" hreflang="sv" data-title="Neuropeptider" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/N%C3%B6ropeptid" title="Nöropeptid – Turkish" lang="tr" hreflang="tr" data-title="Nöropeptid" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9D%D0%B5%D0%B9%D1%80%D0%BE%D0%BF%D0%B5%D0%BF%D1%82%D0%B8%D0%B4%D0%B8" title="Нейропептиди – Ukrainian" 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href="/wiki/File:Neuropeptide_Y.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/Neuropeptide_Y.png/220px-Neuropeptide_Y.png" decoding="async" width="220" height="291" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/1/13/Neuropeptide_Y.png 1.5x" data-file-width="274" data-file-height="362" /></a><figcaption>Neuropeptide Y</figcaption></figure> <p><b>Neuropeptides</b> are chemical messengers made up of small chains of <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> that are synthesized and released by <a href="/wiki/Neuron" title="Neuron">neurons</a>. Neuropeptides typically bind to <a href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">G protein-coupled receptors</a> (GPCRs) to modulate neural activity and other tissues like the gut, muscles, and heart. </p><p>Neuropeptides are synthesized from large precursor proteins which are cleaved and post-translationally processed then packaged into dense core <a href="/wiki/Vesicle_(biology_and_chemistry)" title="Vesicle (biology and chemistry)">vesicles</a>. Neuropeptides are often co-released with other neuropeptides and <a href="/wiki/Neurotransmitters" class="mw-redirect" title="Neurotransmitters">neurotransmitters</a> in a single neuron, yielding a multitude of effects. Once released, neuropeptides can diffuse widely to affect a broad range of targets. </p><p>Neuropeptides are extremely ancient and highly diverse chemical messengers. <a href="/wiki/Placozoa" title="Placozoa">Placozoans</a> such as <i><a href="/wiki/Trichoplax" title="Trichoplax">Trichoplax</a></i>, extremely basal animals which do not possess neurons, use peptides for cell-to-cell communication in a way similar to the neuropeptides of higher animals. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Examples">Examples</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=1" title="Edit section: Examples"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Unreferenced_section plainlinks metadata ambox ambox-content ambox-Unreferenced" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>does not <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">cite</a> any <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">sources</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Neuropeptide" title="Special:EditPage/Neuropeptide">improve this section</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a>. Unsourced material may be challenged and <a href="/wiki/Wikipedia:Verifiability#Burden_of_evidence" title="Wikipedia:Verifiability">removed</a>.</span> <span class="date-container"><i>(<span class="date">February 2024</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Peptide signals play a role in information processing that is different from that of conventional neurotransmitters, and many appear to be particularly associated with specific behaviours. For example, <a href="/wiki/Oxytocin" title="Oxytocin">oxytocin</a> and <a href="/wiki/Vasopressin" title="Vasopressin">vasopressin</a> have striking and specific effects on social behaviours, including maternal behaviour and pair bonding. <a href="/wiki/Crustacean_Cardioactive_Peptide" class="mw-redirect" title="Crustacean Cardioactive Peptide">CCAP</a> has several functions including regulating heart rate, <a href="/wiki/Allatostatin" title="Allatostatin">allatostatin</a> and <a href="/wiki/Proctolin" title="Proctolin">proctolin</a> regulate food intake and growth, <a href="/wiki/Bursicon" title="Bursicon">bursicon</a> controls tanning of the cuticle and <a href="/wiki/Corazonin" title="Corazonin">corazonin</a> has a role in cuticle pigmentation and moulting. </p> <div class="mw-heading mw-heading2"><h2 id="Synthesis">Synthesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=2" title="Edit section: Synthesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuropeptides are synthesized from inactive <a href="/wiki/Protein_precursor" title="Protein precursor">precursor proteins</a> called prepropeptides.<sup id="cite_ref-:3_1-0" class="reference"><a href="#cite_note-:3-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Prepropeptides contain sequences for a family of distinct peptides and often contain duplicated copies of the same peptides, depending on the organism.<sup id="cite_ref-Evolution_of_neuropeptide_signallin_2-0" class="reference"><a href="#cite_note-Evolution_of_neuropeptide_signallin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In addition to the precursor peptide sequences, prepropeptides also contain a signal peptide, spacer peptides, and cleavage sites.<sup id="cite_ref-:6_3-0" class="reference"><a href="#cite_note-:6-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The signal peptide sequence guides the protein to the secretory pathway, starting at the <a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">endoplasmic reticulum</a>. The signal peptide sequence is removed in the endoplasmic reticulum, yielding a propeptide. The propeptide travels to the <a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a> where it is proteolytically cleaved and processed into multiple peptides. Peptides are packaged into dense core vesicles, where further cleaving and processing, such as C-terminal amidation, can occur. Dense core vesicles are transported throughout the neuron and can release peptides at the synaptic cleft, cell body, and along the axon.<sup id="cite_ref-:3_1-1" class="reference"><a href="#cite_note-:3-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:4_4-0" class="reference"><a href="#cite_note-:4-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_5-0" class="reference"><a href="#cite_note-:5-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_6-0" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mechanism">Mechanism</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=3" title="Edit section: Mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuropeptides are released by dense core vesicles after <a href="/wiki/Depolarization" title="Depolarization">depolarization</a> of the cell. Compared to classical <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a> signaling, neuropeptide signaling is more sensitive. Neuropeptide receptor affinity is in the nanomolar to micromolar range while neurotransmitter affinity is in the micromolar to millimolar range. Additionally, dense core vesicles contain a small amount of neuropeptide (3 - 10mM) compared to synaptic vesicles containing neurotransmitters (e.g. 100mM for acetylcholine).<sup id="cite_ref-:8_7-0" class="reference"><a href="#cite_note-:8-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Evidence shows that neuropeptides are released after high-frequency firing or bursts, distinguishing dense core vesicle from synaptic vesicle release.<sup id="cite_ref-:4_4-1" class="reference"><a href="#cite_note-:4-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Neuropeptides utilize volume transmission and are not reuptaken quickly, allowing diffusion across broad areas (nm to mm) to reach targets. Almost all neuropeptides bind to <a href="/wiki/G_protein-coupled_receptor" title="G protein-coupled receptor">G protein-coupled receptors</a> (GPCRs), inducing second messenger cascades to modulate neural activity on long time-scales.<sup id="cite_ref-:3_1-2" class="reference"><a href="#cite_note-:3-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:4_4-2" class="reference"><a href="#cite_note-:4-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:5_5-1" class="reference"><a href="#cite_note-:5-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Expression of neuropeptides in the nervous system is diverse. Neuropeptides are often co-released with other neuropeptides and neurotransmitters, yielding a diversity of effects depending on the combination of release.<sup id="cite_ref-:5_5-2" class="reference"><a href="#cite_note-:5-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:0_8-0" class="reference"><a href="#cite_note-:0-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> For example, <a href="/wiki/Vasoactive_intestinal_peptide" title="Vasoactive intestinal peptide">vasoactive intestinal peptide</a> is typically co-released with acetylcholine.<sup id="cite_ref-:9_9-0" class="reference"><a href="#cite_note-:9-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Neuropeptide release can also be specific. In <i>Drosophila</i> larvae, for example, eclosion hormone is expressed in just two neurons.<sup id="cite_ref-:1_6-1" class="reference"><a href="#cite_note-:1-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Receptor_targets">Receptor targets</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=4" title="Edit section: Receptor targets"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most neuropeptides act on G-protein coupled receptors (GPCRs). Neuropeptide-GPCRs fall into two families: rhodopsin-like and the secretin class.<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>  Most peptides activate a single GPCR, while some activate multiple GPCRs (e.g. AstA, AstC, DTK).<sup id="cite_ref-:0_8-1" class="reference"><a href="#cite_note-:0-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Peptide-GPCR binding relationships are highly conserved across animals. Aside from conserved structural relationships, some peptide-GPCR functions are also conserved across the animal kingdom. For example, neuropeptide F/neuropeptide Y signaling is structurally and functionally conserved between insects and mammals.<sup id="cite_ref-:0_8-2" class="reference"><a href="#cite_note-:0-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>Although peptides mostly target metabotropic receptors, there is some evidence that neuropeptides bind to other receptor targets. Peptide-gated ion channels (FMRFamide-gated sodium channels) have been found in snails and Hydra.<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> Other examples of non-GPCR targets include: insulin-like peptides and tyrosine-kinase receptors in <i>Drosophila</i> and atrial natriuretic peptide and eclosion hormone with membrane-bound guanylyl cyclase receptors in mammals and insects.<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> <div class="mw-heading mw-heading2"><h2 id="Actions">Actions</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=5" title="Edit section: Actions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Due to their modulatory and diffusive nature, neuropeptides can act on multiple time and spatial scales. Below are some examples of neuropeptide actions: </p> <div class="mw-heading mw-heading3"><h3 id="Co-release">Co-release</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=6" title="Edit section: Co-release"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuropeptides are often co-released with other neurotransmitters and neuropeptides to modulate synaptic activity. <a href="/wiki/Synaptic_vesicle" title="Synaptic vesicle">Synaptic vesicles</a> and dense core vesicles can have differential activation properties for release, resulting in context-dependent co-release combinations.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><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><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> For example, insect <a href="/wiki/Motor_neuron" title="Motor neuron">motor neurons</a> are <a href="/wiki/Glutamatergic" title="Glutamatergic">glutamatergic</a> and some contain dense core vesicles with <a href="/wiki/Proctolin" title="Proctolin">proctolin</a>. At low frequency activation, only glutamate is released, yielding fast and rapid excitation of the muscle. At high frequency activation however, dense core vesicles release proctolin, inducing prolonged contractions.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Thus, neuropeptide release can be fine-tuned to modulate synaptic activity in certain contexts. </p><p>Some regions of the nervous system are specialized to release distinctive sets of peptides. For example, the hypothalamus and the pituitary gland release peptides (e.g. TRH, GnRH, CRH, SST) that act as hormones<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> In one subpoplation of the <a href="/wiki/Arcuate_nucleus" class="mw-redirect" title="Arcuate nucleus">arcuate nucleus</a> of the <a href="/wiki/Hypothalamus" title="Hypothalamus">hypothalamus</a>, three <a href="/wiki/Anorectic" title="Anorectic">anorectic</a> peptides are co-expressed: <a href="/wiki/%CE%91-melanocyte-stimulating_hormone" class="mw-redirect" title="Α-melanocyte-stimulating hormone">α-melanocyte-stimulating hormone</a> (α-MSH), <a href="/wiki/Galanin-like_peptide" title="Galanin-like peptide">galanin-like peptide</a>, and <a href="/wiki/Cocaine-and-amphetamine-regulated_transcript" class="mw-redirect" title="Cocaine-and-amphetamine-regulated transcript">cocaine-and-amphetamine-regulated transcript</a> (CART), and in another subpopulation two <a href="/wiki/Orexigenic" class="mw-redirect" title="Orexigenic">orexigenic</a> peptides are co-expressed, <a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">neuropeptide Y</a> and <a href="/wiki/Agouti-related_peptide" title="Agouti-related peptide">agouti-related peptide</a> (AGRP).<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> These peptides are all released in different combinations to signal hunger and satiation cues.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>The following is a list of neuroactive peptides co-released with other neurotransmitters. Transmitter names are shown in bold. </p><p><b><a href="/wiki/Norepinephrine" title="Norepinephrine">Norepinephrine</a></b> (noradrenaline). In neurons of the A2 cell group in the <a href="/wiki/Nucleus_of_the_solitary_tract" class="mw-redirect" title="Nucleus of the solitary tract">nucleus of the solitary tract</a>), norepinephrine co-exists with: </p> <ul><li><a href="/wiki/Galanin" title="Galanin">Galanin</a></li> <li><a href="/wiki/Enkephalin" title="Enkephalin">Enkephalin</a></li> <li><a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">Neuropeptide Y</a></li></ul> <p><b><a href="/wiki/Gamma-Aminobutyric_acid" class="mw-redirect" title="Gamma-Aminobutyric acid">GABA</a></b> </p> <ul><li><a href="/wiki/Somatostatin" title="Somatostatin">Somatostatin</a> (in the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>)</li> <li><a href="/wiki/Cholecystokinin" title="Cholecystokinin">Cholecystokinin</a></li> <li><a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">Neuropeptide Y</a> (in the <a href="/wiki/Arcuate_nucleus" class="mw-redirect" title="Arcuate nucleus">arcuate nucleus</a>)</li></ul> <p><b><a href="/wiki/Acetylcholine" title="Acetylcholine">Acetylcholine</a></b> </p> <ul><li><a href="/wiki/Vasoactive_intestinal_peptide" title="Vasoactive intestinal peptide">VIP</a></li> <li><a href="/wiki/Substance_P" title="Substance P">Substance P</a></li></ul> <p><b><a href="/wiki/Dopamine" title="Dopamine">Dopamine</a></b> </p> <ul><li><a href="/wiki/Cholecystokinin" title="Cholecystokinin">Cholecystokinin</a></li> <li><a href="/wiki/Neurotensin" title="Neurotensin">Neurotensin</a></li> <li><a href="/wiki/Glucagon-like_peptide-1" title="Glucagon-like peptide-1">Glucagon-like peptide-1</a> (in the <a href="/wiki/Nucleus_accumbens" title="Nucleus accumbens">nucleus accumbens</a>)</li></ul> <p><b><a href="/wiki/Epinephrine" class="mw-redirect" title="Epinephrine">Epinephrine</a></b> (adrenaline) </p> <ul><li><a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">Neuropeptide Y</a></li> <li><a href="/wiki/Neurotensin" title="Neurotensin">Neurotensin</a></li></ul> <p><b><a href="/wiki/Serotonin" title="Serotonin">Serotonin</a></b> (5-HT) </p> <ul><li><a href="/wiki/Substance_P" title="Substance P">Substance P</a></li> <li><a href="/wiki/Thyrotropin-releasing_hormone" title="Thyrotropin-releasing hormone">TRH</a></li> <li><a href="/wiki/Enkephalin" title="Enkephalin">Enkephalin</a></li></ul> <p>Some neurons make several different peptides. For instance, <a href="/wiki/Vasopressin" title="Vasopressin">vasopressin</a> co-exists with <a href="/wiki/Dynorphin" title="Dynorphin">dynorphin</a> and <a href="/wiki/Galanin" title="Galanin">galanin</a> in magnocellular neurons of the <a href="/wiki/Supraoptic_nucleus" title="Supraoptic nucleus">supraoptic nucleus</a> and <a href="/wiki/Paraventricular_nucleus" class="mw-redirect" title="Paraventricular nucleus">paraventricular nucleus</a>, and with <a href="/wiki/Corticotropin-releasing_hormone" title="Corticotropin-releasing hormone">CRF</a> (in parvocellular neurons of the <a href="/wiki/Paraventricular_nucleus" class="mw-redirect" title="Paraventricular nucleus">paraventricular nucleus</a>) </p><p><a href="/wiki/Oxytocin" title="Oxytocin">Oxytocin</a> in the <a href="/wiki/Supraoptic_nucleus" title="Supraoptic nucleus">supraoptic nucleus</a> co-exists with <a href="/wiki/Enkephalin" title="Enkephalin">enkephalin</a>, <a href="/wiki/Dynorphin" title="Dynorphin">dynorphin</a>, <a href="/wiki/Cocaine-and_amphetamine_regulated_transcript" class="mw-redirect" title="Cocaine-and amphetamine regulated transcript">cocaine-and amphetamine regulated transcript</a> (CART) and <a href="/wiki/Cholecystokinin" title="Cholecystokinin">cholecystokinin</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Evolution_of_Neuropeptide_Signaling">Evolution of Neuropeptide Signaling</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=7" title="Edit section: Evolution of Neuropeptide Signaling"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Peptide" title="Peptide">Peptides</a> are ancient signaling systems that are found in almost all animals on Earth.<sup id="cite_ref-:2_21-0" class="reference"><a href="#cite_note-:2-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><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> Genome sequencing reveals evidence of neuropeptide genes in <a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a>, <a href="/wiki/Ctenophora" title="Ctenophora">Ctenophora</a>, and <a href="/wiki/Placozoa" title="Placozoa">Placozoa</a>, some of oldest living animals with nervous systems or neural-like tissues.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Evolution_of_neuropeptide_signallin_2-1" class="reference"><a href="#cite_note-Evolution_of_neuropeptide_signallin-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Recent studies also show genomic evidence of neuropeptide processing machinery in metazoans and <a href="/wiki/Choanoflagellate" title="Choanoflagellate">choanoflagellates</a>, suggesting that neuropeptide signaling may predate the development of nervous tissues.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Additionally, <a href="/wiki/Ctenophora" title="Ctenophora">Ctenophore</a> and <a href="/wiki/Placozoa" title="Placozoa">Placozoa</a> neural signaling is entirely peptidergic and lacks the major amine <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitters</a> such as acetylcholine, dopamine, and serotonin.<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-:2_21-1" class="reference"><a href="#cite_note-:2-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> This also suggests that neuropeptide signaling developed before amine neurotransmitters. </p> <div class="mw-heading mw-heading2"><h2 id="Research_history">Research history</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuropeptide&action=edit&section=8" title="Edit section: Research history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the early 1900s, chemical messengers were crudely extracted from whole animal brains and tissues and studied for their physiological effects. In 1931, von Euler and Gaddum, used a similar method to try and isolate acetylcholine but instead discovered a peptide substance that induced physiological changes including muscle contractions and depressed blood pressure. These effects were not abolished using atropine, ruling out the substance as acetylcholine. <sup id="cite_ref-:7_28-0" class="reference"><a href="#cite_note-:7-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:9_9-1" class="reference"><a href="#cite_note-:9-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>In insects, <a href="/wiki/Proctolin" title="Proctolin">proctolin</a> was the first neuropeptide to be isolated and sequenced.<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><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> In 1975, Alvin Starratt and Brian Brown extracted the peptide from hindgut muscles of the cockroach and found that its application enhanced muscle contractions. While Starratt and Brown initially thought of proctolin as an excitatory neurotransmitter, proctolin was later confirmed as a neuromodulatory peptide.<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><p><a href="/wiki/David_de_Wied" title="David de Wied">David de Wied</a> first used the term "neuropeptide" in the 1970s to delineate peptides derived from the nervous system.<sup id="cite_ref-:6_3-1" class="reference"><a href="#cite_note-:6-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:8_7-1" class="reference"><a href="#cite_note-:8-7"><span class="cite-bracket">[</span>7<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=Neuropeptide&action=edit&section=9" 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-:3-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-:3_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:3_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:3_1-2"><sup><i><b>c</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="CITEREFMainsEipper1999" class="citation book cs1">Mains RE, Eipper BA (1999). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/books/NBK28247/">"The Neuropeptides"</a>. <i>Basic Neurochemistry</i> (6th ed.). 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class="Z3988"></span></span> </li> <li id="cite_note-24"><span class="mw-cite-backlink"><b><a href="#cite_ref-24">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTakahashiTakeda2015" class="citation journal cs1">Takahashi T, Takeda N (January 2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4346854">"Insight into the molecular and functional diversity of cnidarian neuropeptides"</a>. <i>International Journal of Molecular Sciences</i>. <b>16</b> (2): 2610–2625. <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%2Fijms16022610">10.3390/ijms16022610</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely 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"Proctolin". <i>Handbook of Hormones</i>. <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%2FB978-0-12-801028-0.00067-2">10.1016/B978-0-12-801028-0.00067-2</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780128010280" title="Special:BookSources/9780128010280"><bdi>9780128010280</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Proctolin&rft.btitle=Handbook+of+Hormones&rft.date=2016&rft_id=info%3Adoi%2F10.1016%2FB978-0-12-801028-0.00067-2&rft.isbn=9780128010280&rft.aulast=Tanaka&rft.aufirst=Y&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuropeptide" 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 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typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <a href="https://commons.wikimedia.org/wiki/Category:Neuropeptides" class="extiw" title="commons:Category:Neuropeptides"><span style="font-style:italic; font-weight:bold;">Neuropeptides</span></a>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/40px-Wiktionary-logo-en-v2.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/60px-Wiktionary-logo-en-v2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Wiktionary-logo-en-v2.svg/80px-Wiktionary-logo-en-v2.svg.png 2x" data-file-width="512" data-file-height="512" /></span></span></div> <div class="side-box-text plainlist">Look up <i><b><a href="https://en.wiktionary.org/wiki/neuropeptide" class="extiw" title="wiktionary:neuropeptide">neuropeptide</a></b></i> in Wiktionary, the free dictionary.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://www.journals.elsevier.com/neuropeptides"><i>Neuropeptides</i> Journal</a></li> <li><a rel="nofollow" class="external text" href="http://isyslab.info/NeuroPep/">Neuropeptides reference website (a comprehensive neuropeptide database)</a></li> <li><a rel="nofollow" class="external text" href="http://www.morganclaypool.com/toc/npe/1/1"><i>Neuropeptides</i> eBook series</a></li> <li><a rel="nofollow" class="external text" href="http://www.wormbook.org/chapters/www_neuropeptides/neuropeptides.html">Neuropeptide chapter in the C. elegans Wormbook</a> excellent, and very accessible, discussion of neuropeptide biology in C. elegans</li></ul> <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 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ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Signal_transduction" title="Template:Signal transduction"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Signal_transduction" title="Template talk:Signal transduction"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Signal_transduction" title="Special:EditPage/Template:Signal transduction"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Cell_signaling_/_Signal_transduction" style="font-size:114%;margin:0 4em"><a href="/wiki/Cell_signaling" title="Cell signaling">Cell signaling</a> / <a href="/wiki/Signal_transduction" title="Signal transduction">Signal transduction</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Signaling pathways</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/G_protein-coupled_receptor" title="G protein-coupled receptor">GPCR</a></li> <li class="mw-empty-elt"></li> <li><a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">Wnt</a></li> <li><a href="/wiki/Receptor_tyrosine_kinase" title="Receptor tyrosine kinase">RTK</a> <ul><li><a href="/wiki/TGF_beta_signaling_pathway" title="TGF beta signaling pathway">TGF beta</a></li> <li><a href="/wiki/MAPK/ERK_pathway" title="MAPK/ERK pathway">MAPK/ERK</a></li></ul></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch</a></li> <li><a href="/wiki/JAK-STAT_signaling_pathway" title="JAK-STAT signaling pathway">JAK-STAT</a></li> <li><a href="/wiki/Akt/PKB_signaling_pathway" title="Akt/PKB signaling pathway">Akt/PKB</a></li> <li><a href="/wiki/Apoptosis" title="Apoptosis">Fas apoptosis</a></li> <li><a href="/wiki/Hippo_signaling_pathway" title="Hippo signaling pathway">Hippo</a></li> <li><a href="/wiki/PI3K/AKT/mTOR_pathway" title="PI3K/AKT/mTOR pathway">PI3K/AKT/mTOR pathway</a></li> <li><a href="/wiki/Integrin#Signal_transduction" title="Integrin">Integrin receptors</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Agents</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/Ligand_(biochemistry)" title="Ligand (biochemistry)">Receptor ligands</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/Hormone" title="Hormone">Hormones</a></li> <li><a href="/wiki/Neurotransmitter" title="Neurotransmitter">Neurotransmitters</a>/<a class="mw-selflink selflink">Neuropeptides</a>/<a href="/wiki/Neurohormone" title="Neurohormone">Neurohormones</a></li> <li><a href="/wiki/Cytokine" title="Cytokine">Cytokines</a></li> <li><a href="/wiki/Growth_factor" title="Growth factor">Growth factors</a></li> <li><a href="/wiki/Cell_signaling" title="Cell signaling">Signaling molecules</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Receptors</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/Cell_surface_receptor" title="Cell surface receptor">Cell surface</a></li> <li><a href="/wiki/Intracellular_receptor" title="Intracellular receptor">Intracellular</a></li> <li><a href="/wiki/Co-receptor" title="Co-receptor">Co-receptor</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Second_messenger_system" title="Second messenger system">Second messenger</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><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/CAMP-dependent_pathway" title="CAMP-dependent pathway">cAMP-dependent pathway</a></li> <li><a href="/wiki/Calcium_signaling" title="Calcium signaling">Ca<sup>2+</sup> signaling</a></li> <li><a href="/wiki/Lipid_signaling" title="Lipid signaling">Lipid signaling</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Assistants:</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/Signal_transducing_adaptor_protein" title="Signal transducing adaptor protein">Signal transducing adaptor protein</a></li> <li><a href="/wiki/Scaffold_protein" title="Scaffold protein">Scaffold protein</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/Transcription_factor" title="Transcription factor">Transcription factors</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_transcription_factor" title="General transcription factor">General</a></li> <li><a href="/wiki/Transcription_preinitiation_complex" title="Transcription preinitiation complex">Transcription preinitiation complex</a></li> <li><a href="/wiki/Transcription_factor_II_D" title="Transcription factor II D">TFIID</a></li> <li><a href="/wiki/Transcription_factor_II_H" title="Transcription factor II H">TFIIH</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By distance</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/Juxtacrine_signalling" title="Juxtacrine signalling">Juxtacrine</a></li> <li><a href="/wiki/Autocrine_signalling" class="mw-redirect" title="Autocrine signalling">Autocrine</a> / <a href="/wiki/Paracrine_signalling" class="mw-redirect" title="Paracrine signalling">Paracrine</a></li> <li><a href="/wiki/Endocrine_system" title="Endocrine system">Endocrine</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other concepts</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/Intracrine" title="Intracrine">Intracrine action</a></li> <li>Neurocrine signaling <ul><li><a href="/wiki/Synaptic_transmission" class="mw-redirect" title="Synaptic transmission">Synaptic transmission</a></li> <li><a href="/wiki/Chemical_synapse" title="Chemical synapse">Chemical synapse</a></li></ul></li> <li><a href="/wiki/Neuroendocrine_cell" title="Neuroendocrine cell">Neuroendocrine signaling</a></li> <li><a href="/wiki/Exocrine_gland" title="Exocrine gland">Exocrine signalling</a> <ul><li><a href="/wiki/Pheromone" title="Pheromone">Pheromones</a></li></ul></li> <li><a href="/wiki/Mechanotransduction" title="Mechanotransduction">Mechanotransduction</a></li> <li><a href="/wiki/Visual_phototransduction" title="Visual phototransduction">Phototransduction</a></li> <li><a href="/wiki/Ion_channel" title="Ion channel">Ion channel gating</a></li> <li><a href="/wiki/Gap_junction" title="Gap junction">Gap junction</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="Peptides:_neuropeptides" 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:Neuropeptides" title="Template:Neuropeptides"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Neuropeptides" title="Template talk:Neuropeptides"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Neuropeptides" title="Special:EditPage/Template:Neuropeptides"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Peptides:_neuropeptides" style="font-size:114%;margin:0 4em"><a href="/wiki/Peptide" title="Peptide">Peptides</a>: <a class="mw-selflink selflink">neuropeptides</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Hormones</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"><i>see <a href="/wiki/Template:Hormones" title="Template:Hormones">hormones</a></i></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Opioid_peptide" title="Opioid peptide">Opioid peptides</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:12em"><a href="/wiki/Dynorphin" title="Dynorphin">Dynorphins</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Dynorphin_A" title="Dynorphin A">Dynorphin A</a> <ul><li><a href="/wiki/Dynorphin_A1%E2%80%938" class="mw-redirect" title="Dynorphin A1–8">Dynorphin A<sub>1–8</sub></a></li></ul></li> <li><a href="/wiki/Dynorphin_B" title="Dynorphin B">Dynorphin B</a></li> <li><a href="/wiki/Big_dynorphin" title="Big dynorphin">Big dynorphin</a></li> <li><a href="/wiki/Leumorphin" title="Leumorphin">Leumorphin</a></li> <li><a href="/wiki/Alpha-Neoendorphin" class="mw-redirect" title="Alpha-Neoendorphin">α-Neoendorphin</a></li> <li><a href="/wiki/Beta-Neoendorphin" class="mw-redirect" title="Beta-Neoendorphin">β-Neoendorphin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em"><a href="/wiki/Endomorphin" title="Endomorphin">Endomorphins</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Endomorphin-1" title="Endomorphin-1">Endomorphin-1</a></li> <li><a href="/wiki/Endomorphin-2" title="Endomorphin-2">Endomorphin-2</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em"><a href="/wiki/Endorphins" title="Endorphins">Endorphins</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Alpha-Endorphin" class="mw-redirect" title="Alpha-Endorphin">α-Endorphin</a></li> <li><a href="/wiki/Beta-Endorphin" class="mw-redirect" title="Beta-Endorphin">β-Endorphin</a></li> <li><a href="/wiki/Gamma-Endorphin" class="mw-redirect" title="Gamma-Endorphin">γ-Endorphin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em"><a href="/wiki/Enkephalin" title="Enkephalin">Enkephalins</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Met-enkephalin" title="Met-enkephalin">Met-enkephalin</a></li> <li><a href="/wiki/Leu-enkephalin" title="Leu-enkephalin">Leu-enkephalin</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em">Others</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adrenorphin" title="Adrenorphin">Adrenorphin</a></li> <li><a href="/wiki/Amidorphin" title="Amidorphin">Amidorphin</a></li> <li><a href="/wiki/Hemorphin" title="Hemorphin">Hemorphin</a> <ul><li><a href="/wiki/Hemorphin-4" title="Hemorphin-4">Hemorphin-4</a></li></ul></li> <li><a href="/wiki/Nociceptin" title="Nociceptin">Nociceptin</a></li> <li><a href="/wiki/Opiorphin" title="Opiorphin">Opiorphin</a></li> <li><a href="/wiki/Spinorphin" title="Spinorphin">Spinorphin</a></li> <li><a href="/wiki/Valorphin" title="Valorphin">Valorphin</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other<br /> neuropeptides</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:12em"><a href="/wiki/Kinin" title="Kinin">Kinins</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <dl><dd><a href="/wiki/Bradykinin" title="Bradykinin">Bradykinins</a></dd> <dd><a href="/wiki/Tachykinin_peptides" title="Tachykinin peptides">Tachykinins</a>: mammal <ul><li><a href="/wiki/Substance_P" title="Substance P">Substance P</a></li> <li><a href="/wiki/Neurokinin_A" title="Neurokinin A">Neurokinin A</a></li> <li><a href="/wiki/Neurokinin_B" title="Neurokinin B">Neurokinin B</a></li></ul></dd> <dd>amphibian <ul><li><a href="/wiki/Kassinin" title="Kassinin">Kassinin</a></li> <li><a href="/wiki/Physalaemin" title="Physalaemin">Physalaemin</a></li></ul></dd></dl> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em">Neuromedins</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Neuromedin_B" title="Neuromedin B">B</a></li> <li><a href="/wiki/Neuromedin_N" title="Neuromedin N">N</a></li> <li><a href="/wiki/Neuromedin_S" title="Neuromedin S">S</a></li> <li><a href="/wiki/Neuromedin_U" title="Neuromedin U">U</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em"><a href="/wiki/Orexin" title="Orexin">Orexins</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Orexin-A" title="Orexin-A">A</a></li> <li><a href="/wiki/Orexin_B" class="mw-redirect" title="Orexin B">B</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:12em">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Angiotensin" title="Angiotensin">Angiotensin</a></li> <li><a href="/wiki/Bombesin" title="Bombesin">Bombesin</a></li> <li><a href="/wiki/Calcitonin_gene-related_peptide" title="Calcitonin gene-related peptide">Calcitonin gene-related peptide</a></li> <li><a href="/wiki/Carnosine" title="Carnosine">Carnosine</a></li> <li><a href="/wiki/Cocaine_and_amphetamine_regulated_transcript" title="Cocaine and amphetamine regulated transcript">Cocaine- and amphetamine-regulated transcript</a></li> <li><a href="/wiki/Delta-sleep-inducing_peptide" title="Delta-sleep-inducing peptide">Delta-sleep-inducing peptide</a></li> <li><a href="/wiki/FMRFamide" title="FMRFamide">FMRFamide</a></li> <li><a href="/wiki/Galanin" title="Galanin">Galanin</a></li> <li><a href="/wiki/Galanin-like_peptide" title="Galanin-like peptide">Galanin-like peptide</a></li> <li><a href="/wiki/Gastrin-releasing_peptide" title="Gastrin-releasing peptide">Gastrin-releasing peptide</a></li> <li><a href="/wiki/Ghrelin" title="Ghrelin">Ghrelin</a></li> <li><a href="/w/index.php?title=Neuropeptide_AF&action=edit&redlink=1" class="new" title="Neuropeptide AF (page does not exist)">Neuropeptide AF</a></li> <li><a href="/wiki/Neuropeptide_FF" title="Neuropeptide FF">Neuropeptide FF</a></li> <li><a href="/w/index.php?title=Neuropeptide_SF&action=edit&redlink=1" class="new" title="Neuropeptide SF (page does not exist)">Neuropeptide SF</a></li> <li><a href="/w/index.php?title=Neuropeptide_VF&action=edit&redlink=1" class="new" title="Neuropeptide VF (page does not exist)">Neuropeptide VF</a></li> <li><a href="/wiki/Neuropeptide_S" title="Neuropeptide S">Neuropeptide S</a></li> <li><a href="/wiki/Neuropeptide_Y" title="Neuropeptide Y">Neuropeptide Y</a></li> <li><a href="/wiki/Neurophysins" title="Neurophysins">Neurophysins</a></li> <li><a href="/wiki/Neurotensin" title="Neurotensin">Neurotensin</a></li> <li><a href="/wiki/Pancreatic_polypeptide" title="Pancreatic polypeptide">Pancreatic polypeptide</a></li> <li><a href="/wiki/Pituitary_adenylate_cyclase-activating_peptide" title="Pituitary adenylate cyclase-activating peptide">Pituitary adenylate cyclase-activating peptide</a></li> <li><a href="/wiki/RVD-Hp%CE%B1" title="RVD-Hpα">RVD-Hpα</a></li> <li><a href="/wiki/VGF" title="VGF">VGF</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"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" 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