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cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Short-term plasticity subsection</span> </button> <ul id="toc-Short-term_plasticity-sublist" class="vector-toc-list"> <li id="toc-Synaptic_enhancement" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Synaptic_enhancement"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Synaptic enhancement</span> </div> </a> <ul id="toc-Synaptic_enhancement-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Synaptic_depression" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Synaptic_depression"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Synaptic depression</span> </div> </a> <ul id="toc-Synaptic_depression-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Long-term_plasticity" class="vector-toc-list-item vector-toc-level-1 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href="#Long-term_potentiation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Long-term potentiation</span> </div> </a> <ul id="toc-Long-term_potentiation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Synaptic_strength" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Synaptic_strength"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Synaptic strength</span> </div> </a> <ul id="toc-Synaptic_strength-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Computational_use_of_plasticity" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Computational_use_of_plasticity"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Computational use of plasticity</span> </div> </a> <ul id="toc-Computational_use_of_plasticity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>External links</span> </div> </a> <button aria-controls="toc-External_links-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 External links subsection</span> </button> <ul id="toc-External_links-sublist" class="vector-toc-list"> <li id="toc-Videos,_podcasts" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Videos,_podcasts"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.1</span> <span>Videos, podcasts</span> </div> </a> <ul id="toc-Videos,_podcasts-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main 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class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Synaptic plasticity</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 19 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-19" 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">19 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/%D9%84%D8%AF%D9%88%D9%86%D8%A9_%D9%85%D8%B4%D8%A8%D9%83%D9%8A%D8%A9" title="لدونة مشبكية – Arabic" lang="ar" hreflang="ar" data-title="لدونة مشبكية" 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/Sinapsna_plasti%C4%8Dnost" title="Sinapsna plastičnost – Bosnian" lang="bs" hreflang="bs" data-title="Sinapsna plastičnost" 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/Plasticitat_sin%C3%A0ptica" title="Plasticitat sinàptica – Catalan" lang="ca" hreflang="ca" data-title="Plasticitat sinàptica" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de badge-Q70894304 mw-list-item" title=""><a href="https://de.wikipedia.org/wiki/Synaptische_Plastizit%C3%A4t" title="Synaptische Plastizität – German" lang="de" hreflang="de" data-title="Synaptische Plastizität" 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/Neuroplasticidad" title="Neuroplasticidad – Spanish" lang="es" hreflang="es" data-title="Neuroplasticidad" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A7%D9%86%D8%B9%D8%B7%D8%A7%D9%81%E2%80%8C%D9%BE%D8%B0%DB%8C%D8%B1%DB%8C_%D8%B3%DB%8C%D9%86%D8%A7%D9%BE%D8%B3%DB%8C" title="انعطاف‌پذیری سیناپسی – Persian" lang="fa" hreflang="fa" data-title="انعطاف‌پذیری سیناپسی" 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/Plasticit%C3%A9_synaptique" title="Plasticité synaptique – French" lang="fr" hreflang="fr" data-title="Plasticité synaptique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%8B%9C%EB%83%85%EC%8A%A4_%EA%B0%80%EC%86%8C%EC%84%B1" title="시냅스 가소성 – Korean" lang="ko" hreflang="ko" data-title="시냅스 가소성" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8D%D5%AB%D5%B6%D5%A1%D5%BA%D5%BD%D5%A1%D5%B5%D5%AB%D5%B6_%D5%B3%D5%AF%D5%B8%D6%82%D5%B6%D5%B8%D6%82%D5%A9%D5%B5%D5%B8%D6%82%D5%B6" title="Սինապսային ճկունություն – Armenian" lang="hy" hreflang="hy" data-title="Սինապսային ճկունություն" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Plasticit%C3%A0_sinaptica" title="Plasticità sinaptica – Italian" lang="it" hreflang="it" data-title="Plasticità sinaptica" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%A1%E1%83%98%E1%83%9C%E1%83%90%E1%83%A4%E1%83%A1%E1%83%A3%E1%83%A0%E1%83%98_%E1%83%9E%E1%83%9A%E1%83%90%E1%83%A1%E1%83%A2%E1%83%98%E1%83%A3%E1%83%A0%E1%83%9D%E1%83%91%E1%83%90" title="სინაფსური პლასტიურობა – Georgian" lang="ka" hreflang="ka" data-title="სინაფსური პლასტიურობა" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Plasticitas_synaptica" title="Plasticitas synaptica – Latin" lang="la" hreflang="la" data-title="Plasticitas synaptica" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target"><span>Latina</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Synaptische_plasticiteit" title="Synaptische plasticiteit – Dutch" lang="nl" hreflang="nl" data-title="Synaptische plasticiteit" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Plasticidade_sin%C3%A1ptica" title="Plasticidade sináptica – Portuguese" lang="pt" hreflang="pt" data-title="Plasticidade sináptica" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A1%D0%B8%D0%BD%D0%B0%D0%BF%D1%82%D0%B8%D1%87%D0%B5%D1%81%D0%BA%D0%B0%D1%8F_%D0%BF%D0%BB%D0%B0%D1%81%D1%82%D0%B8%D1%87%D0%BD%D0%BE%D1%81%D1%82%D1%8C" title="Синаптическая пластичность – Russian" lang="ru" hreflang="ru" data-title="Синаптическая пластичность" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Synaptisk_plasticitet" title="Synaptisk plasticitet – Swedish" lang="sv" hreflang="sv" data-title="Synaptisk plasticitet" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%A1%D0%B8%D0%BD%D0%B0%D0%BF%D1%82%D0%B8%D1%87%D0%BD%D0%B0_%D0%BF%D0%BB%D0%B0%D1%81%D1%82%D0%B8%D1%87%D0%BD%D1%96%D1%81%D1%82%D1%8C" title="Синаптична пластичність – Ukrainian" lang="uk" hreflang="uk" data-title="Синаптична пластичність" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%AA%81%E8%A7%B8%E5%8F%AF%E5%A1%91%E6%80%A7" title="突觸可塑性 – Cantonese" lang="yue" hreflang="yue" data-title="突觸可塑性" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%AA%81%E8%A7%A6%E5%8F%AF%E5%A1%91%E6%80%A7" title="突触可塑性 – Chinese" lang="zh" hreflang="zh" data-title="突触可塑性" data-language-autonym="中文" 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id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Ability of a synapse to strengthen or weaken over time according to its activity</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about synaptic plasticity. For the role of synapse formation and stabilization in plasticity, see <a href="/wiki/Synaptic_stabilization" title="Synaptic stabilization">Synaptic stabilization</a>. For the general concept of brain plasticity, see <a href="/wiki/Neuroplasticity" title="Neuroplasticity">neuroplasticity</a> and <a href="/wiki/Synaptic_stabilization" title="Synaptic stabilization">Synaptic stabilization</a>.</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Synaptic_Plasticity_Rule.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Synaptic_Plasticity_Rule.png/220px-Synaptic_Plasticity_Rule.png" decoding="async" width="220" height="226" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Synaptic_Plasticity_Rule.png/330px-Synaptic_Plasticity_Rule.png 1.5x, //upload.wikimedia.org/wikipedia/commons/4/41/Synaptic_Plasticity_Rule.png 2x" data-file-width="360" data-file-height="369" /></a><figcaption>Synaptic plasticity rule for gradient estimation by dynamic perturbation of conductances</figcaption></figure> <p>In <a href="/wiki/Neuroscience" title="Neuroscience">neuroscience</a>, <b>synaptic plasticity</b> is the ability of <a href="/wiki/Synapses" class="mw-redirect" title="Synapses">synapses</a> to <a href="/wiki/Chemical_synapse#Synaptic_strength" title="Chemical synapse">strengthen or weaken</a> over time, in response to increases or decreases in their activity.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Since <a href="/wiki/Memory" title="Memory">memories</a> are postulated to be represented by vastly interconnected <a href="/wiki/Neural_circuit" title="Neural circuit">neural circuits</a> in the <a href="/wiki/Brain" title="Brain">brain</a>, synaptic plasticity is one of the important neurochemical foundations of <a href="/wiki/Learning" title="Learning">learning</a> and <a href="/wiki/Memory" title="Memory">memory</a> (<i>see <a href="/wiki/Hebbian_theory" title="Hebbian theory">Hebbian theory</a></i>). </p><p>Plastic change often results from the alteration of the number of <a href="/wiki/Neurotransmitter_receptor" title="Neurotransmitter receptor">neurotransmitter receptors</a> located on a synapse.<sup id="cite_ref-NewT_2-0" class="reference"><a href="#cite_note-NewT-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> There are several underlying mechanisms that cooperate to achieve synaptic plasticity, including changes in the quantity of <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitters</a> released into a synapse and changes in how effectively cells respond to those neurotransmitters.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Synaptic plasticity in both <a href="/wiki/Excitatory_synapse" title="Excitatory synapse">excitatory</a> and <a href="/wiki/Inhibitory_synapse" class="mw-redirect" title="Inhibitory synapse">inhibitory</a> synapses has been found to be dependent upon <a href="/wiki/Postsynaptic" class="mw-redirect" title="Postsynaptic">postsynaptic</a> <a href="/wiki/Calcium" title="Calcium">calcium</a> release.<sup id="cite_ref-NewT_2-1" class="reference"><a href="#cite_note-NewT-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Historical_discoveries">Historical discoveries</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=1" title="Edit section: Historical discoveries"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1973, <a href="/wiki/Terje_L%C3%B8mo" title="Terje Lømo">Terje Lømo</a> and <a href="/wiki/Tim_Bliss" class="mw-redirect" title="Tim Bliss">Tim Bliss</a> first described the now widely studied phenomenon of <a href="/wiki/Long-term_potentiation" title="Long-term potentiation">long-term potentiation</a> (LTP) in a publication in the <i>Journal of Physiology</i>. The experiment described was conducted on the synapse between the <a href="/wiki/Perforant_path" title="Perforant path">perforant path</a> and <a href="/wiki/Dentate_gyrus" title="Dentate gyrus">dentate gyrus</a> in the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampi</a> of anaesthetised rabbits. They were able to show a burst of tetanic (100 Hz) stimulus on perforant path fibres led to a dramatic and long-lasting augmentation in the post-synaptic response of cells onto which these fibres synapse in the dentate gyrus. In the same year, the pair published very similar data recorded from awake rabbits. This discovery was of particular interest due to the proposed role of the hippocampus in certain forms of memory. </p> <div class="mw-heading mw-heading2"><h2 id="Biochemical_mechanisms">Biochemical mechanisms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=2" title="Edit section: Biochemical mechanisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Two molecular mechanisms for synaptic plasticity involve the <a href="/wiki/NMDA" class="mw-redirect" title="NMDA">NMDA</a> and <a href="/wiki/AMPA" title="AMPA">AMPA</a> glutamate receptors. Opening of NMDA channels (which relates to the level of cellular <a href="/wiki/Depolarization" title="Depolarization">depolarization</a>) leads to a rise in post-synaptic Ca<sup>2+</sup> concentration and this has been linked to long-term potentiation, LTP (as well as to protein <a href="/wiki/Kinase" title="Kinase">kinase</a> activation); strong depolarization of the post-synaptic cell completely displaces the <a href="/wiki/Magnesium" title="Magnesium">magnesium</a> ions that block NMDA ion channels and allows calcium ions to enter a cell – probably causing LTP, while weaker depolarization only partially displaces the Mg<sup>2+</sup> ions, resulting in less Ca<sup>2+</sup> entering the post-synaptic neuron and lower intracellular Ca<sup>2+</sup> concentrations (which activate protein phosphatases and induce <a href="/wiki/Long-term_depression" title="Long-term depression">long-term depression</a>, LTD).<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>These activated protein kinases serve to phosphorylate post-synaptic excitatory receptors (e.g. <a href="/wiki/AMPA_receptor" title="AMPA receptor">AMPA receptors</a>), improving cation conduction, and thereby potentiating the synapse. Also, these signals recruit additional receptors into the post-synaptic membrane, stimulating the production of a modified receptor type, thereby facilitating an influx of calcium. This in turn increases post-synaptic excitation by a given pre-synaptic stimulus. This process can be reversed via the activity of protein phosphatases, which act to dephosphorylate these cation channels.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The second mechanism depends on a <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">second messenger</a> cascade regulating <a href="/wiki/Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">gene transcription</a> and changes in the levels of key proteins such as <a href="/wiki/CaMKII" class="mw-redirect" title="CaMKII">CaMKII</a> and PKAII. Activation of the second messenger pathway leads to increased levels of CaMKII and PKAII within the <a href="/wiki/Dendritic_spine" title="Dendritic spine">dendritic spine</a>. These protein kinases have been linked to growth in dendritic spine volume and LTP processes such as the addition of AMPA receptors to the <a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> and phosphorylation of ion channels for enhanced permeability.<sup id="cite_ref-Haining09_6-0" class="reference"><a href="#cite_note-Haining09-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Localization or compartmentalization of activated proteins occurs in the presence of their given stimulus which creates local effects in the dendritic spine. Calcium influx from NMDA receptors is necessary for the activation of CaMKII. This activation is localized to spines with focal stimulation and is inactivated before spreading to adjacent spines or the shaft, indicating an important mechanism of LTP in that particular changes in protein activation can be localized or compartmentalized to enhance the responsivity of single dendritic spines. Individual dendritic spines are capable of forming unique responses to presynaptic cells.<sup id="cite_ref-Seok-Jin09_7-0" class="reference"><a href="#cite_note-Seok-Jin09-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> This second mechanism can be triggered by <a href="/wiki/Protein_phosphorylation" title="Protein phosphorylation">protein phosphorylation</a> but takes longer and lasts longer, providing the mechanism for long-lasting memory storage. The duration of the LTP can be regulated by breakdown of these <a href="/wiki/Second_messenger" class="mw-redirect" title="Second messenger">second messengers</a>. <a href="/wiki/Phosphodiesterase" title="Phosphodiesterase">Phosphodiesterase</a>, for example, breaks down the secondary messenger <a href="/wiki/Cyclic_adenosine_monophosphate" title="Cyclic adenosine monophosphate">cAMP</a>, which has been implicated in increased AMPA receptor synthesis in the post-synaptic neuron <sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2011)">citation needed</span></a></i>]</sup>. </p><p>Long-lasting changes in the efficacy of synaptic connections (<a href="/wiki/Long-term_potentiation" title="Long-term potentiation">long-term potentiation</a>, or LTP) between two neurons can involve the making and breaking of synaptic contacts. Genes such as activin ß-A, which encodes a subunit of <a href="/wiki/Activin_A" class="mw-redirect" title="Activin A">activin A</a>, are up-regulated during early stage LTP. The activin molecule modulates the actin dynamics in dendritic spines through the <a href="/wiki/Mitogen-activated_protein_kinase" title="Mitogen-activated protein kinase">MAP-kinase pathway</a>. By changing the <a href="/wiki/F-actin" class="mw-redirect" title="F-actin">F-actin</a> <a href="/wiki/Cytoskeletal" class="mw-redirect" title="Cytoskeletal">cytoskeletal</a> structure of dendritic spines, spine necks are lengthened producing increased electrical isolation.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The end result is long-term maintenance of LTP.<sup id="cite_ref-Synapse_9-0" class="reference"><a href="#cite_note-Synapse-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>The number of <a href="/wiki/Ion_channel" title="Ion channel">ion channels</a> on the post-synaptic membrane affects the strength of the synapse.<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> Research suggests that the density of receptors on post-synaptic membranes changes, affecting the neuron's excitability in response to stimuli. In a dynamic process that is maintained in equilibrium, <a href="/wiki/NMDA_receptor" title="NMDA receptor">N-methyl D-aspartate receptor (NMDA receptor)</a> and AMPA receptors are added to the membrane by <a href="/wiki/Exocytosis" title="Exocytosis">exocytosis</a> and removed by <a href="/wiki/Endocytosis" title="Endocytosis">endocytosis</a>.<sup id="cite_ref-Shi99_11-0" class="reference"><a href="#cite_note-Shi99-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Song02_12-0" class="reference"><a href="#cite_note-Song02-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-PO05_13-0" class="reference"><a href="#cite_note-PO05-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> These processes, and by extension the number of receptors on the membrane, can be altered by synaptic activity.<sup id="cite_ref-Shi99_11-1" class="reference"><a href="#cite_note-Shi99-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-PO05_13-1" class="reference"><a href="#cite_note-PO05-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Experiments have shown that AMPA receptors are delivered to the synapse through vesicular <a href="/wiki/Membrane_fusion" class="mw-redirect" title="Membrane fusion">membrane fusion</a> with the postsynaptic membrane via the protein kinase CaMKII, which is activated by the influx of calcium through NMDA receptors. CaMKII also improves AMPA ionic conductance through phosphorylation.<sup id="cite_ref-Bear_2007_14-0" class="reference"><a href="#cite_note-Bear_2007-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> When there is high-frequency NMDA receptor activation, there is an increase in the expression of a protein <a href="/wiki/PSD-95" class="mw-redirect" title="PSD-95">PSD-95</a> that increases synaptic capacity for AMPA receptors.<sup id="cite_ref-stabilization_plasticity_15-0" class="reference"><a href="#cite_note-stabilization_plasticity-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> This is what leads to a long-term increase in AMPA receptors and thus synaptic strength and plasticity. </p><p>If the strength of a synapse is only reinforced by stimulation or weakened by its lack, a <a href="/wiki/Positive_feedback_loop" class="mw-redirect" title="Positive feedback loop">positive feedback loop</a> will develop, causing some cells never to fire and some to fire too much. But two regulatory forms of plasticity, called scaling and <a href="/wiki/Metaplasticity" title="Metaplasticity">metaplasticity</a>, also exist to provide <a href="/wiki/Negative_feedback" title="Negative feedback">negative feedback</a>.<sup id="cite_ref-PO05_13-2" class="reference"><a href="#cite_note-PO05-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Synaptic scaling is a primary mechanism by which a neuron is able to stabilize firing rates up or down.<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> </p><p><a href="/wiki/Synaptic_scaling" title="Synaptic scaling">Synaptic scaling</a> serves to maintain the strengths of synapses relative to each other, lowering amplitudes of small <a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">excitatory postsynaptic potentials</a> in response to continual excitation and raising them after prolonged blockage or inhibition.<sup id="cite_ref-PO05_13-3" class="reference"><a href="#cite_note-PO05-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This effect occurs gradually over hours or days, by changing the numbers of <a href="/wiki/NMDA_receptor" title="NMDA receptor">NMDA receptors</a> at the synapse (Pérez-Otaño and Ehlers, 2005). <a href="/wiki/Metaplasticity" title="Metaplasticity">Metaplasticity</a> varies the threshold level at which plasticity occurs, allowing integrated responses to synaptic activity spaced over time and preventing saturated states of LTP and LTD. Since LTP and LTD (<a href="/wiki/Long-term_depression" title="Long-term depression">long-term depression</a>) rely on the influx of <a href="/wiki/Calcium_in_biology" title="Calcium in biology">Ca<sup>2+</sup></a> through NMDA channels, metaplasticity may be due to changes in NMDA receptors, altered calcium buffering, altered states of kinases or phosphatases and a priming of protein synthesis machinery.<sup id="cite_ref-Abraham97_17-0" class="reference"><a href="#cite_note-Abraham97-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Synaptic scaling is a primary mechanism by which a neuron to be selective to its varying inputs.<sup id="cite_ref-Abbot2000_18-0" class="reference"><a href="#cite_note-Abbot2000-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The neuronal circuitry affected by LTP/LTD and modified by scaling and metaplasticity leads to reverberatory neural circuit development and regulation in a Hebbian manner which is manifested as memory, whereas the changes in neural circuitry, which begin at the level of the synapse, are an integral part in the ability of an organism to learn.<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> </p><p>There is also a specificity element of biochemical interactions to create synaptic plasticity, namely the importance of location. Processes occur at microdomains – such as <a href="/wiki/Exocytosis" title="Exocytosis">exocytosis</a> of AMPA receptors is spatially regulated by the <a href="/wiki/T-SNARE" class="mw-redirect" title="T-SNARE">t-SNARE</a> <a href="/wiki/STX4" title="STX4">STX4</a>.<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> Specificity is also an important aspect of CAMKII signaling involving nanodomain calcium.<sup id="cite_ref-Seok-Jin09_7-1" class="reference"><a href="#cite_note-Seok-Jin09-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The spatial gradient of PKA between dendritic spines and shafts is also important for the strength and regulation of synaptic plasticity.<sup id="cite_ref-Haining09_6-1" class="reference"><a href="#cite_note-Haining09-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> It is important to remember that the biochemical mechanisms altering synaptic plasticity occur at the level of individual synapses of a neuron. Since the biochemical mechanisms are confined to these "microdomains," the resulting synaptic plasticity affects only the specific synapse at which it took place. </p> <div class="mw-heading mw-heading2"><h2 id="Theoretical_mechanisms">Theoretical mechanisms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=3" title="Edit section: Theoretical mechanisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A bidirectional model, describing both LTP and LTD, of synaptic plasticity has proved necessary for a number of different learning mechanisms in <a href="/wiki/Computational_neuroscience" title="Computational neuroscience">computational neuroscience</a>, <a href="/wiki/Neural_networks_(biology)" class="mw-redirect" title="Neural networks (biology)">neural networks</a>, and <a href="/wiki/Biophysics" title="Biophysics">biophysics</a>. Three major hypotheses for the molecular nature of this plasticity have been well-studied, and none are required to be the exclusive mechanism: </p> <ol><li>Change in the probability of glutamate release.</li> <li>Insertion or removal of post-synaptic AMPA receptors.</li> <li><a href="/wiki/Phosphorylation" title="Phosphorylation">Phosphorylation</a> and de-phosphorylation inducing a change in AMPA receptor conductance.</li></ol> <p>Of these, the latter two hypotheses have been recently mathematically examined to have identical calcium-dependent dynamics which provides strong theoretical evidence for a calcium-based model of plasticity, which in a linear model where the total number of receptors are conserved looks like </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\frac {dW_{i}(t)}{dt}}={\frac {1}{\tau ([Ca^{2+}]_{i})}}\left(\Omega ([Ca^{2+}]_{i})-W_{i}\right),}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi>d</mi> <msub> <mi>W</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> </mrow> <mrow> <mi>d</mi> <mi>t</mi> </mrow> </mfrac> </mrow> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mn>1</mn> <mrow> <mi>τ</mi> <mo stretchy="false">(</mo> <mo stretchy="false">[</mo> <mi>C</mi> <msup> <mi>a</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mo>+</mo> </mrow> </msup> <msub> <mo stretchy="false">]</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">)</mo> </mrow> </mfrac> </mrow> <mrow> <mo>(</mo> <mrow> <mi mathvariant="normal">Ω</mi> <mo stretchy="false">(</mo> <mo stretchy="false">[</mo> <mi>C</mi> <msup> <mi>a</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mo>+</mo> </mrow> </msup> <msub> <mo stretchy="false">]</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">)</mo> <mo>−</mo> <msub> <mi>W</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> </mrow> <mo>)</mo> </mrow> <mo>,</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\frac {dW_{i}(t)}{dt}}={\frac {1}{\tau ([Ca^{2+}]_{i})}}\left(\Omega ([Ca^{2+}]_{i})-W_{i}\right),}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/9e2b8401c9b7f1a704d4900dc33b6edc0edd24a6" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.671ex; width:42.357ex; height:6.509ex;" alt="{\displaystyle {\frac {dW_{i}(t)}{dt}}={\frac {1}{\tau ([Ca^{2+}]_{i})}}\left(\Omega ([Ca^{2+}]_{i})-W_{i}\right),}"></span></dd></dl> <p>where </p> <ul><li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle W_{i}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>W</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle W_{i}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/7301a4cfd04d4f5db4549fdf23746a0d2ce9f387" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:2.993ex; height:2.509ex;" alt="{\displaystyle W_{i}}"></span> is the <a href="/wiki/Synaptic_weight" title="Synaptic weight">synaptic weight</a> of the <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle i}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>i</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle i}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/add78d8608ad86e54951b8c8bd6c8d8416533d20" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:0.802ex; height:2.176ex;" alt="{\displaystyle i}"></span>th input axon,</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle [Ca^{2+}]}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mo stretchy="false">[</mo> <mi>C</mi> <msup> <mi>a</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mo>+</mo> </mrow> </msup> <mo stretchy="false">]</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle [Ca^{2+}]}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/2486b37d36b435e1328d6650185cb2652e8e0760" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:6.623ex; height:3.176ex;" alt="{\displaystyle [Ca^{2+}]}"></span> is the concentration of calcium,</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \tau }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>τ</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \tau }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/38a7dcde9730ef0853809fefc18d88771f95206c" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.202ex; height:1.676ex;" alt="{\displaystyle \tau }"></span> is a time constant dependent on the insertion and removal rates of neurotransmitter receptors, which is dependent on <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle [Ca^{2+}]}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mo stretchy="false">[</mo> <mi>C</mi> <msup> <mi>a</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mo>+</mo> </mrow> </msup> <mo stretchy="false">]</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle [Ca^{2+}]}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/2486b37d36b435e1328d6650185cb2652e8e0760" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:6.623ex; height:3.176ex;" alt="{\displaystyle [Ca^{2+}]}"></span>, and</li> <li><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Omega =\beta A_{m}^{\rm {fp}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi mathvariant="normal">Ω</mi> <mo>=</mo> <mi>β</mi> <msubsup> <mi>A</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>m</mi> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mi mathvariant="normal">f</mi> <mi mathvariant="normal">p</mi> </mrow> </mrow> </msubsup> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Omega =\beta A_{m}^{\rm {fp}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/c4d228b0d316224d4d688045bcf50fd87f5c85fd" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:9.61ex; height:3.176ex;" alt="{\displaystyle \Omega =\beta A_{m}^{\rm {fp}}}"></span> is also a function of the concentration of calcium that depends linearly on the number of receptors on the membrane of the neuron at some fixed point.</li></ul> <p>Both <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Omega }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi mathvariant="normal">Ω</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Omega }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/24b0d5ca6f381068d756f6337c08e0af9d1eeb6f" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.678ex; height:2.176ex;" alt="{\displaystyle \Omega }"></span> and <span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \tau }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>τ</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \tau }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/38a7dcde9730ef0853809fefc18d88771f95206c" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:1.202ex; height:1.676ex;" alt="{\displaystyle \tau }"></span> are found experimentally and agree on results from both hypotheses. The model makes important simplifications that make it unsuited for actual experimental predictions, but provides a significant basis for the hypothesis of a calcium-based synaptic plasticity dependence.<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="Short-term_plasticity">Short-term plasticity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=4" title="Edit section: Short-term plasticity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Short-term synaptic plasticity acts on a timescale of tens of milliseconds to a few minutes unlike long-term plasticity, which lasts from minutes to hours. Short-term plasticity can either strengthen or weaken a synapse. </p> <div class="mw-heading mw-heading3"><h3 id="Synaptic_enhancement">Synaptic enhancement</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=5" title="Edit section: Synaptic enhancement"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Short-term synaptic enhancement results from an increased probability of synaptic terminals releasing transmitters in response to pre-synaptic action potentials. Synapses will strengthen for a short time because of an increase in the amount of packaged transmitter released in response to each action potential.<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> Depending on the time scales over which it acts synaptic enhancement is classified as <a href="/wiki/Neural_facilitation" title="Neural facilitation">neural facilitation</a>, <a href="/wiki/Synaptic_augmentation" title="Synaptic augmentation">synaptic augmentation</a> or <a href="/wiki/Post-tetanic_potentiation" title="Post-tetanic potentiation">post-tetanic potentiation</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Synaptic_depression">Synaptic depression</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=6" title="Edit section: Synaptic depression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Synaptic_fatigue" title="Synaptic fatigue">Synaptic fatigue</a> or depression is usually attributed to the depletion of the readily releasable vesicles. Depression can also arise from post-synaptic processes and from feedback activation of presynaptic receptors.<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> <a href="/wiki/Heterosynaptic_plasticity" title="Heterosynaptic plasticity">heterosynaptic</a> depression is thought to be linked to the release of <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP) from <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a>.<sup id="cite_ref-Glia_24-0" class="reference"><a href="#cite_note-Glia-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Long-term_plasticity">Long-term plasticity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=7" title="Edit section: Long-term plasticity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Long-term_depression" title="Long-term depression">Long-term depression</a> (LTD) and <a href="/wiki/Long-term_potentiation" title="Long-term potentiation">long-term potentiation</a> (LTP) are two forms of long-term plasticity, lasting minutes or more, that occur at excitatory synapses.<sup id="cite_ref-NewT_2-2" class="reference"><a href="#cite_note-NewT-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> NMDA-dependent LTD and LTP have been extensively researched, and are found to require the binding of <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>, and <a href="/wiki/Glycine" title="Glycine">glycine</a> or <a href="/wiki/D-serine" class="mw-redirect" title="D-serine">D-serine</a> for activation of NMDA receptors.<sup id="cite_ref-Glia_24-1" class="reference"><a href="#cite_note-Glia-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> The turning point for the synaptic modification of a synapse has been found to be modifiable itself, depending on the history of the synapse.<sup id="cite_ref-pmid7619513_25-0" class="reference"><a href="#cite_note-pmid7619513-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Recently, a number of attempts have been made to offer a comprehensive model that could account for most forms of synaptic plasticity.<sup id="cite_ref-pmid21348800_26-0" class="reference"><a href="#cite_note-pmid21348800-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Long-term_depression">Long-term depression</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=8" title="Edit section: Long-term depression"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Brief activation of an excitatory pathway can produce what is known as long-term depression (LTD) of synaptic transmission in many areas of the brain. LTD is induced by a minimum level of postsynaptic depolarization and simultaneous increase in the intracellular calcium concentration at the postsynaptic neuron. LTD can be initiated at inactive synapses if the calcium concentration is raised to the minimum required level by heterosynaptic activation, or if the extracellular concentration is raised. These alternative conditions capable of causing LTD differ from the Hebb rule, and instead depend on synaptic activity modifications. <a href="/wiki/D-serine" class="mw-redirect" title="D-serine">D-serine</a> release by <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a> has been found to lead to a significant reduction of LTD in the hippocampus.<sup id="cite_ref-Glia_24-2" class="reference"><a href="#cite_note-Glia-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Activity-dependent LTD was investigated in 2011 for the electrical synapses (modification of Gap Junctions efficacy through their activity).<sup id="cite_ref-pmid22021860_27-0" class="reference"><a href="#cite_note-pmid22021860-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> In the brain, cerebellum is one of the structures where LTD is a form of neuroplasticity.<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> <div class="mw-heading mw-heading3"><h3 id="Long-term_potentiation">Long-term potentiation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=9" title="Edit section: Long-term potentiation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Long-term potentiation, commonly referred to as LTP, is an increase in synaptic response following potentiating pulses of electrical stimuli that sustains at a level above the baseline response for hours or longer. LTP involves interactions between postsynaptic neurons and the specific presynaptic inputs that form a synaptic association, and is specific to the stimulated pathway of synaptic transmission. The long-term stabilization of synaptic changes is determined by a parallel increase of pre- and postsynaptic structures such as <a href="/wiki/Bouton_(synapse)" class="mw-redirect" title="Bouton (synapse)">axonal bouton</a>, <a href="/wiki/Dendritic_spine" title="Dendritic spine">dendritic spine</a> and <a href="/wiki/Postsynaptic_density" title="Postsynaptic density">postsynaptic density</a>.<sup id="cite_ref-stabilization_plasticity_15-1" class="reference"><a href="#cite_note-stabilization_plasticity-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> On the molecular level, an increase of the postsynaptic scaffolding proteins <a href="/wiki/PSD-95" class="mw-redirect" title="PSD-95">PSD-95</a> and <a href="/wiki/HOMER1" title="HOMER1">Homer1c</a> has been shown to correlate with the stabilization of synaptic enlargement.<sup id="cite_ref-stabilization_plasticity_15-2" class="reference"><a href="#cite_note-stabilization_plasticity-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Modification of astrocyte coverage at the synapses in the hippocampus has been found to result from the <a href="/wiki/LTP_induction" title="LTP induction">induction of LTP</a>, which has been found to be linked to the release of <a href="/wiki/D-serine" class="mw-redirect" title="D-serine">D-serine</a>, <a href="/wiki/Nitric_oxide" title="Nitric oxide">nitric oxide</a>, and the <a href="/wiki/Chemokine" title="Chemokine">chemokine</a>, <a href="/wiki/S100B" title="S100B">s100B</a> by <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a>.<sup id="cite_ref-Glia_24-3" class="reference"><a href="#cite_note-Glia-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> LTP is also a model for studying the synaptic basis of Hebbian plasticity. Induction conditions resemble those described for the initiation of long-term depression (LTD), but a stronger depolarization and a greater increase of calcium are necessary to achieve LTP.<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> Experiments performed by stimulating an array of individual dendritic spines, have shown that synaptic cooperativity by as few as two adjacent dendritic spines prevents LTD, allowing only LTP.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Synaptic_strength">Synaptic strength</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=10" title="Edit section: Synaptic strength"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The modification of <a href="/wiki/Synaptic_strength" class="mw-redirect" title="Synaptic strength">synaptic strength</a> is referred to as functional plasticity. Changes in synaptic strength involve distinct mechanisms of particular types of <a href="/wiki/Glial_cell" class="mw-redirect" title="Glial cell">glial cells</a>, the most researched type being <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a>.<sup id="cite_ref-Glia_24-4" class="reference"><a href="#cite_note-Glia-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Computational_use_of_plasticity">Computational use of plasticity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=11" title="Edit section: Computational use of plasticity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Every kind of synaptic plasticity has different computational uses.<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> Short-term facilitation has been demonstrated to serve as both working memory and mapping input for readout, short-term depression for removing auto-correlation. Long-term potentiation is used for spatial memory storage while long-term depression for both encoding space features, selective weakening of synapses and clearing old memory traces respectively. Forward <a href="/wiki/Spike-timing-dependent_plasticity" title="Spike-timing-dependent plasticity">spike-timing-dependent plasticity</a> is used for long range temporal correlation, temporal coding and spatiotemporal coding. The reversed <a href="/wiki/Spike-timing-dependent_plasticity" title="Spike-timing-dependent plasticity">spike-timing-dependent plasticity</a> acts as sensory filtering. </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=12" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 30em;"> <ul><li><a href="/wiki/Homosynaptic_plasticity" title="Homosynaptic plasticity">Homosynaptic plasticity</a></li> <li><a href="/wiki/Homeostatic_plasticity" title="Homeostatic plasticity">Homeostatic plasticity</a></li> <li><a href="/wiki/Inhibitory_postsynaptic_potential" title="Inhibitory postsynaptic potential">Inhibitory postsynaptic potential</a></li> <li><a href="/wiki/Activity-dependent_plasticity" title="Activity-dependent plasticity">Activity-dependent plasticity</a></li> <li><a href="/wiki/Neural_backpropagation" title="Neural backpropagation">Neural backpropagation</a></li> <li><a href="/wiki/Neuroplasticity" title="Neuroplasticity">Neuroplasticity</a></li> <li><a href="/wiki/Postsynaptic_potential" title="Postsynaptic potential">Postsynaptic potential</a></li> <li><a href="/wiki/Non-synaptic_plasticity" class="mw-redirect" title="Non-synaptic plasticity">Non-synaptic plasticity</a></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span 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"Atomic scale nanoelectronics for quantum neuromorphic devices: comparing different materials". <i>International Journal of Nanotechnology</i>. <b>13</b> (7): 509–523. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://arxiv.org/abs/1606.01884">1606.01884</a></span>. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2016IJNT...13..509P">2016IJNT...13..509P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1504%2FIJNT.2016.078543">10.1504/IJNT.2016.078543</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:18697109">18697109</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+Nanotechnology&rft.atitle=Atomic+scale+nanoelectronics+for+quantum+neuromorphic+devices%3A+comparing+different+materials&rft.volume=13&rft.issue=7&rft.pages=509-523&rft.date=2016&rft_id=info%3Aarxiv%2F1606.01884&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A18697109%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1504%2FIJNT.2016.078543&rft_id=info%3Abibcode%2F2016IJNT...13..509P&rft.aulast=Prati&rft.aufirst=E&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASynaptic+plasticity" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=14" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFThornton2003" class="citation journal cs1">Thornton JK (2003). <a rel="nofollow" class="external text" href="http://www.maps.org/news-letters/v13n1/13124tho.html">"New LSD Research: Gene Expression within the Mammalian Brain"</a>. <i>MAPS</i>. <b>13</b> (1)<span class="reference-accessdate">. Retrieved <span class="nowrap">2007-06-08</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=MAPS&rft.atitle=New+LSD+Research%3A+Gene+Expression+within+the+Mammalian+Brain&rft.volume=13&rft.issue=1&rft.date=2003&rft.aulast=Thornton&rft.aufirst=JK&rft_id=http%3A%2F%2Fwww.maps.org%2Fnews-letters%2Fv13n1%2F13124tho.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASynaptic+plasticity" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChapouthier2004" class="citation journal cs1"><a href="/wiki/Georges_Chapouthier" title="Georges Chapouthier">Chapouthier G</a> (2004). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2567045">"From the search for a molecular code of memory to the role of neurotransmitters: a historical perspective"</a>. <i>Neural Plasticity</i>. <b>11</b> (3–4): 151–8. <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/10.1155%2FNP.2004.151">10.1155/NP.2004.151</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/PMC2567045">2567045</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/15656266">15656266</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neural+Plasticity&rft.atitle=From+the+search+for+a+molecular+code+of+memory+to+the+role+of+neurotransmitters%3A+a+historical+perspective&rft.volume=11&rft.issue=3%E2%80%934&rft.pages=151-8&rft.date=2004&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2567045%23id-name%3DPMC&rft_id=info%3Apmid%2F15656266&rft_id=info%3Adoi%2F10.1155%2FNP.2004.151&rft.aulast=Chapouthier&rft.aufirst=G&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2567045&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASynaptic+plasticity" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHawkinsKandelBailey2006" class="citation journal cs1">Hawkins RD, Kandel ER, Bailey CH (June 2006). "Molecular mechanisms of memory storage in Aplysia". <i>The Biological Bulletin</i>. <b>210</b> (3): 174–91. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F4134556">10.2307/4134556</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/4134556">4134556</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/16801493">16801493</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:16448344">16448344</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Biological+Bulletin&rft.atitle=Molecular+mechanisms+of+memory+storage+in+Aplysia&rft.volume=210&rft.issue=3&rft.pages=174-91&rft.date=2006-06&rft_id=info%3Apmid%2F16801493&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A16448344%23id-name%3DS2CID&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F4134556%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.2307%2F4134556&rft.aulast=Hawkins&rft.aufirst=RD&rft.au=Kandel%2C+ER&rft.au=Bailey%2C+CH&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASynaptic+plasticity" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeDoux2002" class="citation book cs1"><a href="/wiki/Joseph_E._LeDoux" title="Joseph E. LeDoux">LeDoux J</a> (2002). <i>Synaptic Self: How Our Brains Become Who We Are</i>. New York: Penguin Books. pp. 1–324.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Synaptic+Self%3A+How+Our+Brains+Become+Who+We+Are.&rft.place=New+York&rft.pages=1-324&rft.pub=Penguin+Books&rft.date=2002&rft.aulast=LeDoux&rft.aufirst=J&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASynaptic+plasticity" class="Z3988"></span></li></ul> </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=Synaptic_plasticity&action=edit&section=15" 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://icwww.epfl.ch/~gerstner//SPNM/node71.html">Overview</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170502074457/http://icwww.epfl.ch/~gerstner/SPNM/node71.html">Archived</a> 2017-05-02 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090210041801/http://cnr.iop.kcl.ac.uk/default.aspx?pageid=169">Finnerty lab, MRC Centre for Neurodegeneration Research, London</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100119002009/http://www.bris.ac.uk/synaptic/public/plasticity.htm">Brain Basics Synaptic Plasticity Synaptic transmission is plastic</a></li> <li><a rel="nofollow" class="external text" href="http://nba.uth.tmc.edu/neuroscience/s1/chapter07.html">Synaptic Plasticity</a>, <i>Neuroscience Online</i> (electronic neuroscience textbook by UT Houston Medical School)</li></ul> <div class="mw-heading mw-heading3"><h3 id="Videos,_podcasts"><span id="Videos.2C_podcasts"></span>Videos, podcasts</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Synaptic_plasticity&action=edit&section=16" title="Edit section: Videos, podcasts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://videocast.nih.gov/Summary.asp?file=13746">Synaptic plasticity: Multiple mechanisms and functions</a> - a lecture by Robert Malenka, M.D., Ph.D., <a href="/wiki/Stanford_University" title="Stanford University">Stanford University</a>. Video podcast, runtime: 01:05:17.</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 .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 dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · ";font-weight:bold}.mw-parser-output .hlist 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system">nervous system</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Primarily CNS</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/Arousal" title="Arousal">Arousal</a> <ul><li><a href="/wiki/Wakefulness" title="Wakefulness">Wakefulness</a></li></ul></li> <li><a href="/wiki/Intracranial_pressure" title="Intracranial pressure">Intracranial pressure</a></li> <li><a href="/wiki/Lateralization_of_brain_function" title="Lateralization of brain function">Lateralization of brain function</a></li> <li><a href="/wiki/Sleep" title="Sleep">Sleep</a></li> <li><a href="/wiki/Memory" title="Memory">Memory</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Primarily PNS</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/Reflex" title="Reflex">Reflex</a></li> <li><a href="/wiki/Sensory_nervous_system" title="Sensory nervous system">Sensation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Both</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/Evoked_potential" title="Evoked potential">Evoked potential</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/Bereitschaftspotential" title="Bereitschaftspotential">Bereitschaftspotential</a></li> <li><a href="/wiki/P300_(neuroscience)" title="P300 (neuroscience)">P300</a></li> <li><a href="/wiki/Evoked_potential" title="Evoked potential">Auditory evoked potential</a></li> <li><a href="/wiki/Somatosensory_evoked_potentials" class="mw-redirect" title="Somatosensory evoked potentials">Somatosensory evoked potentials</a></li> <li><a href="/wiki/Visual_evoked_potential" class="mw-redirect" title="Visual evoked potential">Visual evoked potential</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other short term</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/Neurotransmission" title="Neurotransmission">Neurotransmission</a></li> <li><a href="/wiki/Chronaxie" title="Chronaxie">Chronaxie</a></li> <li><a href="/wiki/Membrane_potential" title="Membrane potential">Membrane potential</a></li> <li><a href="/wiki/Action_potential" title="Action potential">Action potential</a></li> <li><a href="/wiki/Postsynaptic_potential" title="Postsynaptic potential">Postsynaptic potential</a> <ul><li><a href="/wiki/Excitatory_postsynaptic_potential" title="Excitatory postsynaptic potential">Excitatory</a></li> <li><a href="/wiki/Inhibitory_postsynaptic_potential" title="Inhibitory postsynaptic potential">Inhibitory</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Long term</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/Axoplasmic_transport" class="mw-redirect" title="Axoplasmic transport">Axoplasmic transport</a></li> <li><a href="/wiki/Neuroregeneration" title="Neuroregeneration">Neuroregeneration</a>/Nerve regeneration</li> <li><a href="/wiki/Neuroplasticity" title="Neuroplasticity">Neuroplasticity</a>/<a class="mw-selflink selflink">Synaptic plasticity</a> <ul><li><a href="/wiki/Long-term_potentiation" title="Long-term potentiation">Long-term potentiation</a></li> <li><a href="/wiki/Long-term_depression" title="Long-term depression">Long-term depression</a></li></ul></li></ul> 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