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<span>History</span> </div> </a> <ul id="toc-History-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Modern_neuroethology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Modern_neuroethology"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Modern neuroethology</span> </div> </a> <ul id="toc-Modern_neuroethology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Application_to_technology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Application_to_technology"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Application to technology</span> </div> </a> <ul id="toc-Application_to_technology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Case_studies" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Case_studies"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Case studies</span> </div> </a> <button aria-controls="toc-Case_studies-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 Case studies subsection</span> </button> <ul id="toc-Case_studies-sublist" class="vector-toc-list"> <li id="toc-Jamming_avoidance_response" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Jamming_avoidance_response"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Jamming avoidance response</span> </div> </a> <ul id="toc-Jamming_avoidance_response-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Feature_analysis_in_toad_vision" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Feature_analysis_in_toad_vision"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Feature analysis in toad vision</span> </div> </a> <ul id="toc-Feature_analysis_in_toad_vision-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Computational_neuroethology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Computational_neuroethology"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Computational neuroethology</span> </div> </a> <ul id="toc-Computational_neuroethology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Model_systems" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Model_systems"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Model systems</span> </div> </a> <ul id="toc-Model_systems-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-Sources" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Sources"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Sources</span> </div> </a> <ul id="toc-Sources-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">11</span> <span>Further reading</span> </div> </a> <button aria-controls="toc-Further_reading-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 Further reading subsection</span> </button> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> <li id="toc-Textbooks" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Textbooks"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.1</span> <span>Textbooks</span> </div> </a> <ul id="toc-Textbooks-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Articles" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Articles"> <div class="vector-toc-text"> <span class="vector-toc-numb">11.2</span> <span>Articles</span> </div> </a> <ul id="toc-Articles-sublist" class="vector-toc-list"> </ul> </li> </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">12</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Neuroethology</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 22 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-22" 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">22 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%B9%D9%84%D9%85_%D8%A7%D9%84%D8%B3%D9%84%D9%88%D9%83_%D8%A7%D9%84%D8%B9%D8%B5%D8%A8%D9%8A" title="علم السلوك العصبي – Arabic" lang="ar" hreflang="ar" data-title="علم السلوك العصبي" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9D%D0%B5%D0%B9%D1%80%D0%B0%D1%8D%D1%82%D0%B0%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Нейраэталогія – Belarusian" lang="be" hreflang="be" data-title="Нейраэталогія" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%9D%D1%8D%D1%9E%D1%80%D0%B0%D1%8D%D1%82%D0%B0%D0%BB%D1%91%D0%B3%D1%96%D1%8F" title="Нэўраэталёгія – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Нэўраэталёгія" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Neuroetologia" title="Neuroetologia – Catalan" lang="ca" hreflang="ca" data-title="Neuroetologia" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Neuroetologi" title="Neuroetologi – Danish" lang="da" hreflang="da" data-title="Neuroetologi" 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/Neuroethologie" title="Neuroethologie – German" lang="de" hreflang="de" data-title="Neuroethologie" 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/Neuroetolog%C3%ADa" title="Neuroetología – Spanish" lang="es" hreflang="es" data-title="Neuroetología" 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%B1%D9%81%D8%AA%D8%A7%D8%B1%D8%B4%D9%86%D8%A7%D8%B3%DB%8C_%D8%B9%D8%B5%D8%A8%DB%8C" title="رفتارشناسی عصبی – Persian" lang="fa" hreflang="fa" data-title="رفتارشناسی عصبی" 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/Neuro%C3%A9thologie" title="Neuroéthologie – French" lang="fr" hreflang="fr" data-title="Neuroéthologie" 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%A0%EA%B2%BD%ED%96%89%EB%8F%99%ED%95%99" title="신경행동학 – Korean" lang="ko" hreflang="ko" data-title="신경행동학" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Neuroetologia" title="Neuroetologia – Italian" lang="it" hreflang="it" data-title="Neuroetologia" 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%9C%E1%83%94%E1%83%98%E1%83%A0%E1%83%9D%E1%83%94%E1%83%97%E1%83%9D%E1%83%9A%E1%83%9D%E1%83%92%E1%83%98%E1%83%90" title="ნეიროეთოლოგია – Georgian" lang="ka" hreflang="ka" data-title="ნეიროეთოლოგია" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target"><span>ქართული</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Neuroetologia" title="Neuroetologia – Polish" lang="pl" hreflang="pl" data-title="Neuroetologia" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Neuroetologia" title="Neuroetologia – Portuguese" lang="pt" hreflang="pt" data-title="Neuroetologia" 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%9D%D0%B5%D0%B9%D1%80%D0%BE%D1%8D%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F" title="Нейроэтология – Russian" lang="ru" hreflang="ru" data-title="Нейроэтология" 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/Neuroethology" title="Neuroethology – Simple English" lang="en-simple" hreflang="en-simple" data-title="Neuroethology" 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-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Neuroetologi" title="Neuroetologi – Swedish" lang="sv" hreflang="sv" data-title="Neuroetologi" 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%B6roetoloji" title="Nöroetoloji – Turkish" lang="tr" hreflang="tr" data-title="Nöroetoloji" 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%B5%D1%82%D0%BE%D0%BB%D0%BE%D0%B3%D1%96%D1%8F" title="Нейроетологія – Ukrainian" lang="uk" hreflang="uk" data-title="Нейроетологія" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%B9%D8%B5%D8%A8%DB%8C_%D8%B3%D9%84%D9%88%DA%A9%DB%8C%D8%A7%D8%AA" title="عصبی سلوکیات – Urdu" lang="ur" hreflang="ur" data-title="عصبی سلوکیات" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Th%E1%BA%A7n_kinh_h%E1%BB%8Dc_%C4%91%E1%BB%99ng_v%E1%BA%ADt" title="Thần kinh học động vật – Vietnamese" lang="vi" hreflang="vi" data-title="Thần kinh học động vật" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%A5%9E%E7%BB%8F%E8%A1%8C%E4%B8%BA%E5%AD%A6" title="神经行为学 – Chinese" lang="zh" hreflang="zh" data-title="神经行为学" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q1336265#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div 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searchaux" style="display:none">Study of animal behavior and its underlying mechanistic control by the nervous system</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-Citation_style plainlinks metadata ambox ambox-style ambox-citation_style" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>has an unclear <a href="/wiki/Wikipedia:Citing_sources#Citation_style" title="Wikipedia:Citing sources">citation style</a></b>. The reason given is: <b>numerous sources not properly notated.</b><span class="hide-when-compact"> The references used may be made clearer with a different or consistent style of <a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources">citation</a> and <a href="/wiki/Help:Footnotes" title="Help:Footnotes">footnoting</a>.</span> <span class="date-container"><i>(<span class="date">March 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> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Neuroethology_of_bat_echolocation.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/eb/Neuroethology_of_bat_echolocation.png/220px-Neuroethology_of_bat_echolocation.png" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/e/eb/Neuroethology_of_bat_echolocation.png 1.5x" data-file-width="311" data-file-height="311" /></a><figcaption><i><a href="/wiki/Animal_Echolocation" class="mw-redirect" title="Animal Echolocation">Echolocation</a></i> in bats is one model system in neuroethology.</figcaption></figure> <p><b>Neuroethology</b> is the evolutionary and comparative approach to the study of <a href="/wiki/Animal" title="Animal">animal</a> behavior and its underlying mechanistic control by the nervous system.<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><sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> It is an interdisciplinary science that combines both <a href="/wiki/Neuroscience" title="Neuroscience">neuroscience</a> (study of the nervous system) and <a href="/wiki/Ethology" title="Ethology">ethology</a> (study of animal behavior in natural conditions). A central theme of neuroethology, which differentiates it from other branches of neuroscience, is its focus on behaviors that have been favored by <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> (e.g., finding mates, navigation, locomotion, and predator avoidance) rather than on behaviors that are specific to a particular disease state or laboratory experiment. </p><p>Neuroethologists hope to uncover general principles of the nervous system from the study of animals with exaggerated or specialized behaviors. They endeavor to understand how the nervous system translates biologically relevant stimuli into natural behavior. For example, many bats are capable of <a href="/wiki/Animal_Echolocation" class="mw-redirect" title="Animal Echolocation">echolocation</a> which is used for prey capture and navigation. The auditory system of bats is often cited as an example for how acoustic properties of sounds can be converted into a sensory map of behaviorally relevant features of sounds.<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> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Philosophy">Philosophy</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=1" title="Edit section: Philosophy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuroethology is an integrative approach to the study of animal behavior that draws upon several disciplines. Its approach stems from the <a href="/wiki/Theory" title="Theory">theory</a> that animals' nervous systems have evolved to address problems of sensing and acting in certain environmental niches and that their nervous systems are best understood in the context of the problems they have evolved to solve. In accordance with <a href="/wiki/Krogh%27s_principle" title="Krogh's principle">Krogh's principle</a>, neuroethologists often study animals that are "specialists" in the behavior the researcher wishes to study e.g. honeybees and social behavior, bat echolocation, owl sound localization, etc. </p><p>The scope of neuroethological inquiry might be summarized by <a href="/wiki/J%C3%B6rg-Peter_Ewert" title="Jörg-Peter Ewert">Jörg-Peter Ewert</a>, a pioneer of neuroethology, when he considers the types of questions central to neuroethology in his 1980 introductory text to the field: </p> <ol><li>How are stimuli detected by an organism?</li> <li>How are environmental stimuli in the external world represented in the nervous system?</li> <li>How is information about a stimulus acquired, stored and recalled by the nervous system?</li> <li>How is a behavioral pattern encoded by neural networks?</li> <li>How is behavior coordinated and controlled by the nervous system?</li> <li>How can the ontogenetic development of behavior be related to neural mechanisms?</li></ol> <p>Often central to addressing questions in neuroethology are comparative methodologies, drawing upon knowledge about related organisms' nervous systems, anatomies, life histories, behaviors and environmental niches. While it is not unusual for many types of neurobiology experiments to give rise to behavioral questions, many neuroethologists often begin their research programs by observing a species' behavior in its natural environment. Other approaches to understanding nervous systems include the systems identification approach, popular in <a href="/wiki/Engineering" title="Engineering">engineering</a>. The idea is to stimulate the system using a non-natural stimulus with certain properties. The system's response to the stimulus may be used to analyze the operation of the system. Such an approach is useful for <a href="/wiki/Linear" class="mw-redirect" title="Linear">linear</a> systems, but the nervous system is notoriously <a href="/wiki/Nonlinear" class="mw-redirect" title="Nonlinear">nonlinear</a>, and neuroethologists argue that such an approach is limited. This argument is supported by experiments in the auditory system, which show that neural responses to complex sounds, like social calls, can not be predicted by the knowledge gained from studying the responses due to pure tones (one of the non-natural stimuli favored by auditory neurophysiologists). This is because of the non-linearity of the system. </p><p>Modern neuroethology is largely influenced by the research techniques used. Neural approaches are necessarily very diverse, as is evident through the variety of questions asked, measuring techniques used, relationships explored, and model systems employed. Techniques utilized since 1984 include the use of intracellular dyes, which make maps of identified neurons possible, and the use of brain slices, which bring vertebrate brains into better observation through intracellular electrodes (Hoyle 1984). Currently, other fields toward which neuroethology may be headed include <a href="/wiki/Computational_neuroscience" title="Computational neuroscience">computational neuroscience</a>, <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a>, <a href="/wiki/Neuroendocrinology" title="Neuroendocrinology">neuroendocrinology</a> and <a href="/wiki/Epigenetics" title="Epigenetics">epigenetics</a>. The existing field of neural modeling may also expand into neuroethological terrain, due to its practical uses in <a href="/wiki/Robotics" title="Robotics">robotics</a>. In all this, neuroethologists must use the right level of simplicity to effectively guide research towards accomplishing the goals of neuroethology. </p><p>Critics of neuroethology might consider it a branch of neuroscience concerned with 'animal trivia'. Though neuroethological subjects tend not to be traditional neurobiological model systems (i.e. <i><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila</a></i>, <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i>, or <i><a href="/wiki/Danio_rerio" class="mw-redirect" title="Danio rerio">Danio rerio</a></i>), neuroethological approaches emphasizing comparative methods have uncovered many concepts central to neuroscience as a whole, such as <a href="/wiki/Lateral_inhibition" title="Lateral inhibition">lateral inhibition</a>, <a href="/wiki/Coincidence_detection_in_neurobiology" title="Coincidence detection in neurobiology">coincidence detection</a>, and <a href="/wiki/Sensory_maps" class="mw-redirect" title="Sensory maps">sensory maps</a>. The discipline of neuroethology has also discovered and explained the only vertebrate behavior for which the entire neural circuit has been described: the <a href="/wiki/Electric_fish" title="Electric fish">electric fish</a> <a href="/wiki/Jamming_avoidance_response" title="Jamming avoidance response">jamming avoidance response</a>. Beyond its conceptual contributions, neuroethology makes indirect contributions to advancing human health. By understanding simpler nervous systems, many clinicians have used concepts uncovered by neuroethology and other branches of <a href="/wiki/Neuroscience" title="Neuroscience">neuroscience</a> to develop treatments for devastating human diseases. </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=2" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuroethology owes part of its existence to the establishment of ethology as a unique discipline within <a href="/wiki/Zoology" title="Zoology">zoology</a>. Although animal behavior had been studied since the time of <a href="/wiki/Aristotle" title="Aristotle">Aristotle</a> (384–342 BC), it was not until the early twentieth century that ethology finally became distinguished from natural science (a strictly descriptive field) and ecology. The main catalysts behind this new distinction were the research and writings of <a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> and <a href="/wiki/Niko_Tinbergen" class="mw-redirect" title="Niko Tinbergen">Niko Tinbergen</a>. </p><p>Konrad Lorenz was born in Austria in 1903, and is widely known for his contribution of the theory of <a href="/wiki/Fixed_action_pattern" title="Fixed action pattern">fixed action patterns</a> (FAPs): endogenous, instinctive behaviors involving a complex sequence of movements that are triggered ("released") by a certain kind of stimulus. This sequence always proceeds to completion, even if the original stimulus is removed. It is also species-specific and performed by nearly all members. Lorenz constructed his famous "hydraulic model" to help illustrate this concept, as well as the concept of action specific energy, or drives. </p><p>Niko Tinbergen was born in the Netherlands in 1907 and worked closely with Lorenz in the development of the FAP theory; their studies focused on the egg retrieval response of nesting geese. Tinbergen performed extensive research on the releasing mechanisms of particular FAPs, and used the bill-pecking behavior of baby herring gulls as his model system. This led to the concept of the <a href="/wiki/Supernormal_stimulus" title="Supernormal stimulus">supernormal stimulus</a>. Tinbergen is also well known for his <a href="/wiki/Tinbergen%27s_four_questions" title="Tinbergen's four questions">four questions</a> that he believed ethologists should be asking about any given animal behavior; among these is that of the mechanism of the behavior, on a physiological, neural and <a href="/wiki/Molecular" class="mw-redirect" title="Molecular">molecular</a> level, and this question can be thought of in many regards as the keystone question in neuroethology. Tinbergen also emphasized the need for ethologists and neurophysiologists to work together in their studies, a unity that has become a reality in the field of neuroethology. </p><p>Unlike <a href="/wiki/Behaviorism" title="Behaviorism">behaviorism</a>, which studies animals' reactions to non-natural <a href="/wiki/Stimulus_(physiology)" title="Stimulus (physiology)">stimuli</a> in artificial, <a href="/wiki/Laboratory" title="Laboratory">laboratory</a> conditions, ethology sought to categorize and analyze the natural behaviors of animals in a <a href="/wiki/Field_research" title="Field research">field setting</a>. Similarly, neuroethology asks questions about the neural bases of <i>naturally occurring</i> behaviors, and seeks to mimic the natural context as much as possible in the laboratory. </p><p>Although the development of ethology as a distinct discipline was crucial to the advent of neuroethology, equally important was the development of a more comprehensive understanding of <a href="/wiki/Neuroscience" title="Neuroscience">neuroscience</a>. Contributors to this new understanding were the Spanish Neuroanatomist, <a href="/wiki/Ramon_y_Cajal" class="mw-redirect" title="Ramon y Cajal">Ramon y Cajal</a> (born in 1852), and physiologists <a href="/wiki/Charles_Sherrington" class="mw-redirect" title="Charles Sherrington">Charles Sherrington</a>, <a href="/wiki/Edgar_Adrian" title="Edgar Adrian">Edgar Adrian</a>, <a href="/wiki/Alan_Hodgkin" title="Alan Hodgkin">Alan Hodgkin</a>, and <a href="/wiki/Andrew_Huxley" title="Andrew Huxley">Andrew Huxley</a>. Charles Sherrington, who was born in Great Britain in 1857, is famous for his work on the nerve synapse as the site of transmission of nerve impulses, and for his work on reflexes in the spinal cord. His research also led him to hypothesize that every muscular activation is coupled to an inhibition of the opposing muscle. He was awarded a Nobel Prize for his work in 1932 along with Lord Edgar Adrian who made the first physiological recordings of neural activity from single nerve fibers. </p><p>Alan Hodgkin and Andrew Huxley (born 1914 and 1917, respectively, in Great Britain), are known for their collaborative effort to understand the production of action potentials in the giant axons of squid. The pair also proposed the existence of ion channels to facilitate action potential initiation, and were awarded the Nobel Prize in 1963 for their efforts. </p><p>As a result of this pioneering research, many scientists then sought to connect the physiological aspects of the nervous and sensory systems to specific behaviors. These scientists – <a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a>, <a href="/wiki/Erich_von_Holst" title="Erich von Holst">Erich von Holst</a>, and <a href="/wiki/Theodore_Bullock" class="mw-redirect" title="Theodore Bullock">Theodore Bullock</a> – are frequently referred to as the "fathers" of neuroethology.<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> Neuroethology did not really come into its own, though, until the 1970s and 1980s, when new, sophisticated experimental methods allowed researchers such as <a href="/wiki/Masakazu_Konishi" title="Masakazu Konishi">Masakazu Konishi</a>, <a href="/wiki/Walter_Heiligenberg" title="Walter Heiligenberg">Walter Heiligenberg</a>, <a href="/wiki/J%C3%B6rg-Peter_Ewert" title="Jörg-Peter Ewert">Jörg-Peter Ewert</a>, and others to study the neural circuits underlying verifiable behavior. </p> <div class="mw-heading mw-heading2"><h2 id="Modern_neuroethology">Modern neuroethology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=3" title="Edit section: Modern neuroethology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The International Society for Neuroethology represents the present discipline of neuroethology, which was founded on the occasion of the NATO-Advanced Study Institute "Advances in Vertebrate Neuroethology" (August 13–24, 1981) organized by J.-P. Ewert, D.J. Ingle and R.R. Capranica, held at the University of Kassel in Hofgeismar, Germany (cf. report Trends in Neurosci. 5:141-143,1982). Its first president was <a href="/wiki/Theodore_H._Bullock" class="mw-redirect" title="Theodore H. Bullock">Theodore H. Bullock</a>. The society has met every three years since its first meeting in Tokyo in 1986. </p><p>Its membership draws from many research programs around the world; many of its members are students and faculty members from medical schools and neurobiology departments from various universities. Modern advances in <a href="/wiki/Neurophysiology" title="Neurophysiology">neurophysiology</a> techniques have enabled more exacting approaches in an ever-increasing number of animal systems, as size limitations are being dramatically overcome. Survey of the most recent (2007) congress of the ISN meeting symposia topics gives some idea of the field's breadth: </p> <ul><li>Comparative aspects of spatial memory (rodents, birds, humans, bats)</li> <li>Influences of higher processing centers in active sensing (primates, owls, electric fish, rodents, frogs)</li> <li>Animal signaling plasticity over many time scales (electric fish, frogs, birds)</li> <li>Song production and learning in passerine birds</li> <li>Primate sociality</li> <li>Optimal function of sensory systems (flies, moths, frogs, fish)</li> <li>Neuronal complexity in behavior (insects, computational)</li> <li>Contributions of genes to behavior (Drosophila, honeybees, zebrafish)</li> <li>Eye and head movement (crustaceans, humans, robots)</li> <li>Hormonal actions in brain and behavior (rodents, primates, fish, frogs, and birds)</li> <li>Cognition in insects (honeybee)</li></ul> <div class="mw-heading mw-heading2"><h2 id="Application_to_technology">Application to technology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=4" title="Edit section: Application to technology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuroethology can help create advancements in <a href="/wiki/Technology" title="Technology">technology</a> through an advanced understanding of animal behavior. Model systems were generalized from the study of simple and related animals to humans. For example, the neuronal cortical space map discovered in bats, a specialized champion of hearing and navigating, elucidated the concept of a computational space map. In addition, the discovery of the space map in the barn owl led to the first neuronal example of the <a href="/wiki/Lloyd_A._Jeffress" title="Lloyd A. Jeffress">Jeffress</a> model. This understanding is translatable to understanding spatial localization in humans, a mammalian relative of the bat. Today, knowledge learned from neuroethology are being applied in new technologies. For example, Randall Beer and his colleagues used algorithms learned from insect walking behavior to create robots designed to walk on uneven surfaces (Beer et al.). Neuroethology and technology contribute to one another bidirectionally. </p><p>Neuroethologists seek to understand the neural basis of a behavior as it would occur in an animal's natural environment but the techniques for neurophysiological analysis are lab-based, and cannot be performed in the field setting. This dichotomy between field and lab studies poses a challenge for neuroethology. From the neurophysiology perspective, experiments must be designed for controls and objective rigor, which contrasts with the ethology perspective – that the experiment be applicable to the animal's natural condition, which is uncontrolled, or subject to the dynamics of the environment. An early example of this is when Walter Rudolf Hess developed focal brain stimulation technique to examine a cat's brain controls of vegetative functions in addition to other behaviors. Even though this was a breakthrough in technological abilities and technique, it was not used by many neuroethologists originally because it compromised a cat's natural state, and, therefore, in their minds, devalued the experiments' relevance to real situations. </p><p>When intellectual obstacles like this were overcome, it led to a golden age of neuroethology, by focusing on simple and robust forms of behavior, and by applying modern neurobiological methods to explore the entire chain of sensory and neural mechanisms underlying these behaviors (Zupanc 2004). New technology allows neuroethologists to attach electrodes to even very sensitive parts of an animal such as its brain while it interacts with its environment.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The founders of neuroethology ushered this understanding and incorporated technology and creative experimental design. Since then even indirect technological advancements such as battery-powered and waterproofed instruments have allowed neuroethologists to mimic natural conditions in the lab while they study behaviors objectively. In addition, the electronics required for amplifying neural signals and for transmitting them over a certain distance have enabled neuroscientists to record from behaving animals<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> performing activities in naturalistic environments. Emerging technologies can complement neuroethology, augmenting the feasibility of this valuable perspective of natural neurophysiology. </p><p>Another challenge, and perhaps part of the beauty of neuroethology, is experimental design. The value of neuroethological criteria speak to the reliability of these experiments, because these discoveries represent behavior in the environments in which they evolved. Neuroethologists foresee future advancements through using new technologies and techniques, such as computational neuroscience, neuroendocrinology, and molecular genetics that mimic natural environments.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Case_studies">Case studies</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=5" title="Edit section: Case studies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Jamming_avoidance_response">Jamming avoidance response</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=6" title="Edit section: Jamming avoidance response"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1963, Akira Watanabe and Kimihisa Takeda discovered the behavior of the <a href="/wiki/Jamming_avoidance_response" title="Jamming avoidance response">jamming avoidance response</a> in the knifefish <i>Eigenmannia</i> sp. In collaboration with T.H. Bullock and colleagues, the behavior was further developed. Finally, the work of <a href="/wiki/Walter_Heiligenberg" title="Walter Heiligenberg">W. Heiligenberg</a> expanded it into a full neuroethology study by examining the series of neural connections that led to the behavior. <i>Eigenmannia</i> is a weakly electric fish that can generate electric discharges through electrocytes in its tail. Furthermore, it has the ability to electrolocate by analyzing the perturbations in its electric field. However, when the frequency of a neighboring fish's current is very close (less than 20 Hz difference) to that of its own, the fish will avoid having their signals interfere through a behavior known as Jamming Avoidance Response. If the neighbor's frequency is higher than the fish's discharge frequency, the fish will lower its frequency, and vice versa. The sign of the frequency difference is determined by analyzing the "beat" pattern of the incoming interference which consists of the combination of the two fish's discharge patterns. </p><p>Neuroethologists performed several experiments under <i>Eigenmannia</i><span class="nowrap" style="padding-left:0.1em;">'</span>s natural conditions to study how it determined the sign of the frequency difference. They manipulated the fish's discharge by injecting it with curare which prevented its natural electric organ from discharging. Then, an electrode was placed in its mouth and another was placed at the tip of its tail. Likewise, the neighboring fish's electric field was mimicked using another set of electrodes. This experiment allowed neuroethologists to manipulate different discharge frequencies and observe the fish's behavior. From the results, they were able to conclude that the electric field frequency, rather than an internal frequency measure, was used as a reference. This experiment is significant in that not only does it reveal a crucial neural mechanism underlying the behavior but also demonstrates the value neuroethologists place on studying animals in their natural habitats. </p> <div class="mw-heading mw-heading3"><h3 id="Feature_analysis_in_toad_vision">Feature analysis in toad vision</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=7" title="Edit section: Feature analysis in toad vision"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The recognition of prey and predators in the toad was first studied in depth by <a href="https://de.wikipedia.org/wiki/J%C3%B6rg-Peter_Ewert" class="extiw" title="de:Jörg-Peter Ewert">Jörg-Peter Ewert</a> (Ewert 1974; see also 2004). He began by observing the natural prey-catching behavior of the common toad (<i>Bufo bufo</i>) and concluded that the animal followed a sequence that consisted of stalking, binocular fixation, snapping, swallowing and mouth-wiping. However, initially, the toad's actions were dependent on specific features of the sensory stimulus: whether it demonstrated worm or anti-worm configurations. It was observed that the worm configuration, which signaled prey, was initiated by movement along the object's long axis, whereas anti-worm configuration, which signaled predator, was due to movement along the short axis. (Zupanc 2004). </p><p>Ewert and coworkers adopted a variety of methods to study the predator versus prey behavior response. They conducted recording experiments where they inserted electrodes into the brain, while the toad was presented with worm or anti-worm stimuli. This technique was repeated at different levels of the visual system and also allowed <a href="/wiki/Feature_detection_(nervous_system)" title="Feature detection (nervous system)">feature detectors</a> to be identified. In focus was the discovery of prey-selective neurons in the optic tectum, whose axons could be traced towards the snapping pattern generating cells in the hypoglossal nucleus. The discharge patterns of prey-selective tectal neurons in response to prey objects – in freely moving toads – "predicted" prey-catching reactions such as snapping. Another approach, called stimulation experiment, was carried out in freely moving toads. Focal electrical stimuli were applied to different regions of the brain, and the toad's response was observed. When the thalamic-pretectal region was stimulated, the toad exhibited escape responses, but when the tectum was stimulated in an area close to prey-selective neurons, the toad engaged in prey catching behavior (Carew 2000). Furthermore, neuroanatomical experiments were carried out where the toad's thalamic-pretectal/tectal connection was lesioned and the resulting deficit noted: the prey-selective properties were abolished both in the responses of prey-selective neurons and in the prey catching behavior. These and other experiments suggest that prey selectivity results from pretecto-tectal influences. </p><p>Ewert and coworkers showed in toads that there are stimulus-response mediating pathways that translate perception (of visual sign stimuli) into action (adequate behavioral responses). In addition there are modulatory loops that initiate, modify or specify this mediation (Ewert 2004). Regarding the latter, for example, the telencephalic caudal ventral striatum is involved in a loop gating the stimulus-response mediation in a manner of directed attention. The telencephalic ventral medial pallium („primordium hippocampi"), however, is involved in loops that either modify prey-selection due to associative learning or specify prey-selection due to non-associative learning, respectively. </p> <div class="mw-heading mw-heading2"><h2 id="Computational_neuroethology">Computational neuroethology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=8" title="Edit section: Computational neuroethology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Computational neuroethology (CN<sup id="cite_ref-franklin_9-0" class="reference"><a href="#cite_note-franklin-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> or CNE<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>) is concerned with the computer modelling of the neural mechanisms underlying animal behaviors. Together with the term "artificial ethology," the term "computational neuroethology" was first published in literature by Achacoso and Yamamoto in the Spring of 1990,<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> based on their pioneering work on the <a href="/wiki/Connectome" title="Connectome">connectome</a> of C. elegans in 1989,<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> with further publications in 1992.<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> Computational neuroethology was argued for in depth later in 1990 by <a href="/wiki/Randall_Beer" title="Randall Beer">Randall Beer</a><sup id="cite_ref-beer90_15-0" class="reference"><a href="#cite_note-beer90-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> and by <a href="/wiki/Dave_Cliff_(computer_scientist)" title="Dave Cliff (computer scientist)">Dave Cliff</a><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> both of whom acknowledged the strong influence of <a href="/wiki/Michael_Arbib" class="mw-redirect" title="Michael Arbib">Michael Arbib</a>'s <i>Rana Computatrix</i> computational model of neural mechanisms for visual guidance in frogs and toads.<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> </p><p>CNE systems work within a closed-loop environment; that is, they perceive their (perhaps artificial) environment directly, rather than through human input, as is typical in <a href="/wiki/Artificial_intelligence" title="Artificial intelligence">AI</a> systems.<sup id="cite_ref-franklin_9-1" class="reference"><a href="#cite_note-franklin-9"><span class="cite-bracket">[</span>9<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> For example, Barlow et al. developed a time-dependent model for the retina of the horseshoe crab <a href="/wiki/Horseshoe_crab" title="Horseshoe crab"><i>Limulus polyphemus</i></a> on a <a href="/wiki/Connection_Machine" title="Connection Machine">Connection Machine</a> (Model CM-2).<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> Instead of feeding the model retina with idealized input signals, they exposed the simulation to digitized video sequences made underwater, and compared its response with those of real animals. </p> <div class="mw-heading mw-heading2"><h2 id="Model_systems">Model systems</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=9" title="Edit section: Model systems"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Animal_Echolocation#Bats" class="mw-redirect" title="Animal Echolocation">Bat echolocation</a> – nocturnal flight navigation and prey capture; location of objects using echo returns of its own call</li> <li><a href="/wiki/Songbird" title="Songbird">Oscine</a> <a href="/wiki/Bird_song" class="mw-redirect" title="Bird song">bird song</a> – zebra finch (<i><a href="/wiki/Taeniopygia_guttata" class="mw-redirect" title="Taeniopygia guttata">Taeniopygia guttata</a></i>), canary (<i><a href="/wiki/Serinus_canaria" class="mw-redirect" title="Serinus canaria">Serinus canaria</a></i>) and white-crowned sparrow (<i><a href="/wiki/Zonotrichia_leucophrys" class="mw-redirect" title="Zonotrichia leucophrys">Zonotrichia leucophrys</a></i>); song learning as a model for human speech development</li> <li><a href="/wiki/Electric_fish" title="Electric fish">Electric fish</a> – navigation, communication, Jamming Avoidance Response (JAR), <a href="/wiki/PID_controller" class="mw-redirect" title="PID controller">corollary discharge</a>, expectation generators, and spike timing dependent plasticity</li> <li><a href="/wiki/Barn-owl" class="mw-redirect" title="Barn-owl">Barn owl</a> auditory spatial map – nocturnal prey location and capture</li> <li><a href="/wiki/Vision_in_toads" title="Vision in toads">Toad vision</a> – discrimination of prey versus predator – <a rel="nofollow" class="external text" href="http://www.joerg-peter-ewert.de/13.html">Video</a> "Image processing in the toad's visual system: behavior, brain function, <a rel="nofollow" class="external text" href="http://www.joerg-peter-ewert.de/12.html">artificial neuronal net</a>"</li> <li><a href="/wiki/Circadian_rhythm" title="Circadian rhythm">Circadian rhythm</a> – influence of various circadian controlled behaviors by the <a href="/wiki/Suprachiasmatic_nucleus" title="Suprachiasmatic nucleus">suprachiasmatic nucleus</a></li> <li><a href="/wiki/Cricket_(insect)#Chirping" title="Cricket (insect)">Cricket song</a> – mate attraction and corollary discharge</li> <li>Fish <a href="/wiki/Mauthner_cells" class="mw-redirect" title="Mauthner cells">Mauthner cells</a> – C-start escape response and underwater directional hearing</li> <li>Fly – Microscale directional hearing in <i><a href="/wiki/Ormia" title="Ormia">Ormia</a> ochracea</i>, <a href="/wiki/Sex_Differences_in_Sensory_Systems" class="mw-redirect" title="Sex Differences in Sensory Systems">sex differences of the visual system</a> in <a href="/wiki/Bibionidae" title="Bibionidae">Bibionidae</a>, and spatial navigation in chasing behavior of <a href="/wiki/Fannia_canicularis" class="mw-redirect" title="Fannia canicularis">Fannia canicularis</a></li> <li><a href="/wiki/Noctuid_moth" class="mw-redirect" title="Noctuid moth">Noctuid moths</a> – <a href="/wiki/Ultrasound_avoidance" title="Ultrasound avoidance">ultrasound avoidance</a> response to bat calls</li> <li><i><a href="/wiki/Aplysia" title="Aplysia">Aplysia</a></i> sea hares – learning and memory in startle response</li> <li>Rat – <a href="/wiki/Spatial_memory" title="Spatial memory">spatial memory</a> and navigation</li> <li>Salmon <a href="/wiki/Homing_(biology)" title="Homing (biology)">homing</a> – olfactory imprinting and <a href="/wiki/Thyroid_hormone" class="mw-redirect" title="Thyroid hormone">thyroid hormones</a></li> <li><a href="/wiki/Crayfish" title="Crayfish">Crayfish</a> – escape and startle behaviors, aggression and formation of social hierarchies</li> <li><a href="/wiki/Cichlid" title="Cichlid">Cichlid</a> fish – aggression and attack behaviors</li> <li><a href="/wiki/Honey_bee" title="Honey bee">Honey bee</a> – learning, navigation, vision, olfaction, flight, aggression, foraging</li> <li><a href="/wiki/Monarch_(butterfly)" class="mw-redirect" title="Monarch (butterfly)">Monarch butterfly</a> – navigational mechanisms</li> <li><a rel="nofollow" class="external text" href="http://nelson.beckman.uiuc.edu/courses/neuroethol/model_systems.html">More Model Systems and Information</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=10" 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: 40em;"> <ul><li><a href="/wiki/Niko_Tinbergen" class="mw-redirect" title="Niko Tinbergen">Niko Tinbergen</a></li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Erich_von_Holst" title="Erich von Holst">Erich von Holst</a></li> <li><a href="/wiki/Theodore_H._Bullock" class="mw-redirect" title="Theodore H. Bullock">Theodore H. Bullock</a></li> <li><a href="/wiki/J%C3%B6rg-Peter_Ewert" title="Jörg-Peter Ewert">Jörg-Peter Ewert</a></li> <li><a href="/wiki/Eric_Knudsen" title="Eric Knudsen">Eric Knudsen</a></li> <li><a href="/wiki/Masakazu_Konishi" title="Masakazu Konishi">Masakazu Konishi</a></li> <li><a href="/wiki/Martin_Giurfa" title="Martin Giurfa">Martin Giurfa</a></li></ul> </div> <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=Neuroethology&action=edit&section=11" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text">Hoyle, G. (1984) The scope of Neuroethology. The Behavioral and Brain Sciences. <b>7</b>: 367–412.</span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text">Ewert, P. (1980) Neuroethology. Springer-Verlag. New York.</span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text">Camhi, J. (1984) Neuroethology. Sinauer. Sunderland Mass.</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text">Suga, N. (1989). "Principles of auditory information-processing derived from neuroethology." J Exp Biol 146: 277–86.</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBullock1999" class="citation journal cs1">Bullock, T. H. (1999). "Neuroethology has pregnant agendas". <i>Journal of Comparative Physiology A</i>. <b>185</b> (4): 291–295. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs003590050389">10.1007/s003590050389</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/10555265">10555265</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:5194095">5194095</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Comparative+Physiology+A&rft.atitle=Neuroethology+has+pregnant+agendas&rft.volume=185&rft.issue=4&rft.pages=291-295&rft.date=1999&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A5194095%23id-name%3DS2CID&rft_id=info%3Apmid%2F10555265&rft_id=info%3Adoi%2F10.1007%2Fs003590050389&rft.aulast=Bullock&rft.aufirst=T.+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEwertBorchers1974" class="citation journal cs1">Ewert, J. -P.; Borchers, H. -W. 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"Antwort von retinalen Ganglienzellen bei freibeweglichen Kröten (<i>Bufo bufo</i> L.)". <i>Journal of Comparative Physiology</i>. <b>92</b> (2): 117–130. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF00694501">10.1007/BF00694501</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:11003900">11003900</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Comparative+Physiology&rft.atitle=Antwort+von+retinalen+Ganglienzellen+bei+freibeweglichen+Kr%C3%B6ten+%28Bufo+bufo+L.%29&rft.volume=92&rft.issue=2&rft.pages=117-130&rft.date=1974&rft_id=info%3Adoi%2F10.1007%2FBF00694501&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A11003900%23id-name%3DS2CID&rft.aulast=Ewert&rft.aufirst=J.+-P.&rft.au=Borchers%2C+H.+-W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchürg-PfeifferSpreckelsenEwert1993" class="citation journal cs1">Schürg-Pfeiffer, E.; Spreckelsen, C.; Ewert, J.-P. (1993). "Temporal discharge patterns of tectal and medullary neurons chronically recorded during snapping toward prey in toads <i>Bufo bufo spinosus</i>". <i>Journal of Comparative Physiology A</i>. <b>173</b> (3): 363–376. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF00212701">10.1007/BF00212701</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:41990022">41990022</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Comparative+Physiology+A&rft.atitle=Temporal+discharge+patterns+of+tectal+and+medullary+neurons+chronically+recorded+during+snapping+toward+prey+in+toads+Bufo+bufo+spinosus&rft.volume=173&rft.issue=3&rft.pages=363-376&rft.date=1993&rft_id=info%3Adoi%2F10.1007%2FBF00212701&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A41990022%23id-name%3DS2CID&rft.aulast=Sch%C3%BCrg-Pfeiffer&rft.aufirst=E.&rft.au=Spreckelsen%2C+C.&rft.au=Ewert%2C+J.-P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text">Zupanc, Günther K.H. (2004). Behavioral Neurobiology an Integrative Approach. Oxford University Press. New York.</span> </li> <li id="cite_note-franklin-9"><span class="mw-cite-backlink">^ <a href="#cite_ref-franklin_9-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-franklin_9-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStan_Franklin1998" class="citation book cs1">Stan Franklin (1998). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=_2XbpFvR1IoC&q=%22Computational+neuroethology%22+cn&pg=PA13"><i>Artificial Minds</i></a>. MIT Press. pp. 13–14. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-262-56109-9" title="Special:BookSources/978-0-262-56109-9"><bdi>978-0-262-56109-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Artificial+Minds&rft.pages=13-14&rft.pub=MIT+Press&rft.date=1998&rft.isbn=978-0-262-56109-9&rft.au=Stan+Franklin&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D_2XbpFvR1IoC%26q%3D%2522Computational%2Bneuroethology%2522%2Bcn%26pg%3DPA13&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-10">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMargaret_A._Boden2006" class="citation book cs1">Margaret A. Boden (2006). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/mindasmachinehis0001bode"><i>Mind as machine</i></a></span>. Oxford University Press. p. 1291. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-924144-6" title="Special:BookSources/978-0-19-924144-6"><bdi>978-0-19-924144-6</bdi></a>. <q>Computational neuroethology cne.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Mind+as+machine&rft.pages=1291&rft.pub=Oxford+University+Press&rft.date=2006&rft.isbn=978-0-19-924144-6&rft.au=Margaret+A.+Boden&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fmindasmachinehis0001bode&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAchacosoYamamoto1990" class="citation journal cs1">Achacoso, Theodore B.; Yamamoto, William S. (1990). "Artificial Ethology and Computational Neuroethology: A Scientific Discipline and Its Subset by Sharpening and Extending the Definition of Artificial Intelligence". <i>Perspectives in Biology and Medicine</i>. <b>33</b> (3): 379–390. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1353%2Fpbm.1990.0020">10.1353/pbm.1990.0020</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1529-8795">1529-8795</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/2188211">2188211</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:13002088">13002088</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Perspectives+in+Biology+and+Medicine&rft.atitle=Artificial+Ethology+and+Computational+Neuroethology%3A+A+Scientific+Discipline+and+Its+Subset+by+Sharpening+and+Extending+the+Definition+of+Artificial+Intelligence&rft.volume=33&rft.issue=3&rft.pages=379-390&rft.date=1990&rft.issn=1529-8795&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A13002088%23id-name%3DS2CID&rft_id=info%3Apmid%2F2188211&rft_id=info%3Adoi%2F10.1353%2Fpbm.1990.0020&rft.aulast=Achacoso&rft.aufirst=Theodore+B.&rft.au=Yamamoto%2C+William+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAchacosoFernandezNguyenYamamoto1989" class="citation journal cs1">Achacoso, Theodore B.; Fernandez, Victor; Nguyen, Duc C.; Yamamoto, William S. (1989-11-08). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2245716">"Computer Representation of the Synaptic Connectivity of Caenorhabditis Elegans"</a>. <i>Proceedings of the Annual Symposium on Computer Application in Medical Care</i>: 330–334. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0195-4210">0195-4210</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2245716">2245716</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Annual+Symposium+on+Computer+Application+in+Medical+Care&rft.atitle=Computer+Representation+of+the+Synaptic+Connectivity+of+Caenorhabditis+Elegans&rft.pages=330-334&rft.date=1989-11-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2245716%23id-name%3DPMC&rft.issn=0195-4210&rft.aulast=Achacoso&rft.aufirst=Theodore+B.&rft.au=Fernandez%2C+Victor&rft.au=Nguyen%2C+Duc+C.&rft.au=Yamamoto%2C+William+S.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2245716&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20191015035549/https://www.crcpress.com/Ays-Neuroanatomy-of-C-Elegans-for-Computation/Achacoso-Yamamoto/p/book/9780849342349">"Ay's Neuroanatomy of C. Elegans for Computation"</a>. <i>CRC Press</i>. Archived from <a rel="nofollow" class="external text" href="https://www.crcpress.com/Ays-Neuroanatomy-of-C-Elegans-for-Computation/Achacoso-Yamamoto/p/book/9780849342349">the original</a> on 2019-10-15<span class="reference-accessdate">. Retrieved <span class="nowrap">2019-10-16</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=CRC+Press&rft.atitle=Ay%27s+Neuroanatomy+of+C.+Elegans+for+Computation&rft_id=https%3A%2F%2Fwww.crcpress.com%2FAys-Neuroanatomy-of-C-Elegans-for-Computation%2FAchacoso-Yamamoto%2Fp%2Fbook%2F9780849342349&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFYamamotoAchacoso1992" class="citation journal cs1">Yamamoto, William S.; Achacoso, Theodore B. (1992-06-01). "Scaling up the nervous system of Caenorhabditis elegans: Is one ape equal to 33 million worms?". <i>Computers and Biomedical Research</i>. <b>25</b> (3): 279–291. <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%2F0010-4809%2892%2990043-A">10.1016/0010-4809(92)90043-A</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0010-4809">0010-4809</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/1611892">1611892</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Computers+and+Biomedical+Research&rft.atitle=Scaling+up+the+nervous+system+of+Caenorhabditis+elegans%3A+Is+one+ape+equal+to+33+million+worms%3F&rft.volume=25&rft.issue=3&rft.pages=279-291&rft.date=1992-06-01&rft.issn=0010-4809&rft_id=info%3Apmid%2F1611892&rft_id=info%3Adoi%2F10.1016%2F0010-4809%2892%2990043-A&rft.aulast=Yamamoto&rft.aufirst=William+S.&rft.au=Achacoso%2C+Theodore+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-beer90-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-beer90_15-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRandall_D._Beer1990" class="citation book cs1">Randall D. Beer (1990). <i>Intelligence as Adaptive Behavior: An experiment in computational neuroethology</i>. Academic Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-12-084730-2" title="Special:BookSources/0-12-084730-2"><bdi>0-12-084730-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Intelligence+as+Adaptive+Behavior%3A+An+experiment+in+computational+neuroethology&rft.pub=Academic+Press&rft.date=1990&rft.isbn=0-12-084730-2&rft.au=Randall+D.+Beer&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text">D. Cliff (1990) Computational Neuroethology: A provisional manifesto. In J.-A. Meyer and S. W. Wilson (editors): <i>From Animals to Animats: Proceedings of the First International Conference on the Simulation of Adaptive Behaviour (SAB90)</i>. MIT Press Bradford Books, 1991, pp. 29–39. <a rel="nofollow" class="external autonumber" href="http://citeseer.ist.psu.edu/viewdoc/download;jsessionid=AC6014F67854B884C2836A5BC2B57914?doi=10.1.1.27.4422&rep=rep1&type=pdf">[1]</a></span> </li> <li id="cite_note-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-17">^</a></b></span> <span class="reference-text">M. A. Arbib (1987) Levels of Modeling of Mechanisms of Visually Guided Behavior. <i>The Behavioral and Brain Sciences</i> 10:407–465.</span> </li> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text">D. Cliff (2003) Neuroethology, Computational. In M. A. Arbib (editor): <i>The Handbook of Brain Theory and Neural Networks</i>. Second Edition. MIT Press Bradford Books. pp. 737–741.</span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBarlowPrakashSolessio1993" class="citation journal cs1">Barlow, Robert B.; Prakash, Ramkrishna; Solessio, Eduardo (1993). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Ficb%2F33.1.66">"The Neural Network of the Limulus Retina: From Computer to Behavior"</a>. <i>American Zoologist</i>. <b>33</b>: 66–78. <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/10.1093%2Ficb%2F33.1.66">10.1093/icb/33.1.66</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Zoologist&rft.atitle=The+Neural+Network+of+the+Limulus+Retina%3A+From+Computer+to+Behavior&rft.volume=33&rft.pages=66-78&rft.date=1993&rft_id=info%3Adoi%2F10.1093%2Ficb%2F33.1.66&rft.aulast=Barlow&rft.aufirst=Robert+B.&rft.au=Prakash%2C+Ramkrishna&rft.au=Solessio%2C+Eduardo&rft_id=https%3A%2F%2Fdoi.org%2F10.1093%252Ficb%252F33.1.66&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Sources">Sources</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=12" title="Edit section: Sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Beer D., Randall, Roy E. Ritzmann, Thomas McKenna (1993) Biological neural networks in invertebrate neuroethology and robotics. Boston : Academic Press.</li> <li>Camhi, J.M. (1984) Neuroethology: Nerve cells and the Natural behavior of Animals, Sinauer Associates.</li> <li>Carew, T.J. (2000) Feature analysis in Toads. Behavioral Neurobiology, Sunderland, MA: Sinauer, pp. 95–119.</li> <li>Carew, T.J. (2000) Behavioral neurobiology: The Cellular Organization of Natural Behavior, Sinauer Associates.</li> <li>Ewert, J.-P. (1974) The neural basis of visually guided behavior. Scientific American 230(3):34-42</li> <li>Ewert J.-P. (2004) Motion perception shapes the visual world of amphibians. In: Prete F.R. (Ed.) Complex Worlds from Simpler Nervous Systems. Cambridge, MA, MIT Press, pp. 117–160</li> <li>Hoyle, G. (1984) The scope of Neuroethology. Behavioural Brain Science 7:367-412. Graham Hoyle put forth a rather narrow definition of the goals and subject matter of neuroethology and links the field to the field of ethology. This is followed by commentaries from many prominent neuroethologists. It makes for fascinating reading.</li> <li>Metzner, W. (1993) The Jamming avoidance response in <i>Eigenmannia</i> is controlled by two separate motor pathways. The Journal of Neuroscience. 13(5):1862-1878</li> <li>Pfluger, H.-J. and R. Menzel (1999) Neuroethology, its roots and future. J Comp Physiol A 185:389-392.</li> <li>Zupanc, G.K.H. (2004) Behavioral Neurobiology: An Integrative Approach. Oxford University Press: Oxford, UK.</li></ul> <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=Neuroethology&action=edit&section=13" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Textbooks">Textbooks</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=14" title="Edit section: Textbooks"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>Sillar, K.T., Picton, L.P., Heitler, W.J. (2016) The Neuroethology of Predation and Escape. John Wiley & Sons Inc., New York.</li> <li>Zupanc, G.K.H. (2004) Behavioral Neurobiology an Integrative Approach. Oxford University Press, New York.</li> <li>Carew, T.J. (2000) Behavioral Neurobiology: The Cellular Organization of Natural Behavior. Sinauer, Sunderland Mass.</li> <li>Simmons, P., Young, D. (1999) Nerve Cells and Animal Behaviour. Second Edition. Cambridge University Press, New York.</li> <li>Simmons, P., Young, D. (2010) Nerve Cells and Animal Behaviour. Third Edition. Cambridge University Press, New York.</li> <li>Camhi J. (1984) Neuroethology: Nerve Cells and the Natural Behavior of Animals. Sinauer Associates, Sunderland Mass.</li> <li>Guthrie, D.M. (1980) Neuroethology: An Introduction. Wiley, New York.</li> <li>Ewert, J.-P. (1980) Neuroethology: An Introduction to the Neurophysiological Fundamentals of Behaviour. Springer-Verlag, New York.</li> <li>Ewert, J.-P. (1976) Neuroethologie: Einführung in die neurophysiologischen Grundlagen des Verhaltens. HT 181. Springer-Verlag Heidelberg, Berlin, New York.</li> <li><a href="/wiki/Eric_Kandel" title="Eric Kandel">Kandel, E.R.</a> (1976) Cellular Basis of Behavior: An Introduction to Behavioral Neurobiology. W.H. Freeman</li> <li>Roeder, K.D. (1967) Nerve Cells and Insect Behavior. Harvard University Press, Cambridge Mass.</li> <li><a href="/wiki/Peter_Marler" title="Peter Marler">Marler, P</a>., Hamilton, W.J. (1966) Mechanisms of Animal Behavior. John Wiley & Sons Inc., New York.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Articles">Articles</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Neuroethology&action=edit&section=15" title="Edit section: Articles"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeslie_Brothers_&_Brian_Ring1992" class="citation cs2">Leslie Brothers & Brian Ring (1992), "A neuroethological framework for the representation of minds", <i><a href="/wiki/Journal_of_Cognitive_Neuroscience" title="Journal of Cognitive Neuroscience">Journal of Cognitive Neuroscience</a></i>, <b>4</b> (2): 107–118, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1162%2Fjocn.1992.4.2.107">10.1162/jocn.1992.4.2.107</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/23967887">23967887</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:6607487">6607487</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Cognitive+Neuroscience&rft.atitle=A+neuroethological+framework+for+the+representation+of+minds&rft.volume=4&rft.issue=2&rft.pages=107-118&rft.date=1992&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A6607487%23id-name%3DS2CID&rft_id=info%3Apmid%2F23967887&rft_id=info%3Adoi%2F10.1162%2Fjocn.1992.4.2.107&rft.au=Leslie+Brothers+%26+Brian+Ring&rfr_id=info%3Asid%2Fen.wikipedia.org%3ANeuroethology" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="http://www.scholarpedia.org/article/Neuroethology">Günther K. H. Zupanc (2010), Neuroethology, Scholarpedia, 5(10):5306.</a></li></ul> <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=Neuroethology&action=edit&section=16" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20111222150615/http://www.neuroethology.org/">International Society for Neuroethology</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090127014505/http://nelson.beckman.uiuc.edu/neuroethology.html">Topics in Neuroethology</a></li> <li><a rel="nofollow" class="external free" href="https://web.archive.org/web/20071006201121/http://www.tamie.org/insect.png">https://web.archive.org/web/20071006201121/http://www.tamie.org/insect.png</a></li> <li><a rel="nofollow" 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style="width:1%">Concepts</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/Feed_forward_(control)" title="Feed forward (control)">Feedforward</a></li> <li><a href="/wiki/Coincidence_detection_in_neurobiology" title="Coincidence detection in neurobiology">Coincidence detector</a></li> <li><i><a href="/wiki/Umwelt" title="Umwelt">Umwelt</a></i></li> <li><a href="/wiki/Instinct" title="Instinct">Instinct</a></li> <li><a href="/wiki/Feature_detection_(nervous_system)" title="Feature detection (nervous system)">Feature detection</a></li> <li><a href="/wiki/Central_pattern_generator" title="Central pattern generator">Central pattern generator (CPG)</a></li> <li><a href="/wiki/NMDA_receptor" title="NMDA receptor">NMDA receptor</a></li> <li><a href="/wiki/Lateral_inhibition" title="Lateral inhibition">Lateral inhibition</a></li> <li><a href="/wiki/Fixed_action_pattern" title="Fixed action pattern">Fixed action pattern</a></li> <li><a href="/wiki/Krogh%27s_Principle" class="mw-redirect" title="Krogh's Principle">Krogh's Principle</a></li> <li><a href="/wiki/Hebbian_theory" title="Hebbian theory">Hebbian theory</a></li> <li><a href="/wiki/Anti-Hebbian_learning" title="Anti-Hebbian learning">Anti-Hebbian learning</a></li> <li><a href="/wiki/Sound_localization" title="Sound localization">Sound localization</a></li> <li><a href="/wiki/Ultrasound_avoidance" title="Ultrasound avoidance">Ultrasound avoidance</a> in insects</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">People</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/Theodore_Holmes_Bullock" title="Theodore Holmes Bullock">Theodore Holmes Bullock</a></li> <li><a href="/wiki/Walter_Heiligenberg" title="Walter Heiligenberg">Walter Heiligenberg</a></li> <li><a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Niko Tinbergen</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Donald_Griffin" title="Donald Griffin">Donald Griffin</a></li> <li><a href="/wiki/Donald_Kennedy" title="Donald Kennedy">Donald Kennedy</a></li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a></li> <li><a href="/wiki/Erich_von_Holst" title="Erich von Holst">Erich von Holst</a></li> <li><a href="/wiki/J%C3%B6rg-Peter_Ewert" title="Jörg-Peter Ewert">Jörg-Peter Ewert</a></li> <li><a href="/wiki/Franz_Huber" title="Franz Huber">Franz Huber</a></li> <li><a href="/wiki/Bernhard_Hassenstein" title="Bernhard Hassenstein">Bernhard Hassenstein</a></li> <li><a href="/wiki/Werner_E._Reichardt" title="Werner E. Reichardt">Werner E. Reichardt</a></li> <li><a href="/wiki/Eric_Knudsen" title="Eric Knudsen">Eric Knudsen</a></li> <li><a href="/wiki/Eric_Kandel" title="Eric Kandel">Eric Kandel</a></li> <li><a href="/wiki/Nobuo_Suga" title="Nobuo Suga">Nobuo Suga</a></li> <li><a href="/wiki/Masakazu_Konishi" title="Masakazu Konishi">Masakazu Konishi</a></li> <li><a href="/wiki/Fernando_Nottebohm" title="Fernando Nottebohm">Fernando Nottebohm</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Methods</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/Patch_clamp" title="Patch clamp">Patch clamp</a></li> <li><a href="/wiki/Slice_preparation" title="Slice preparation">Slice preparation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Systems</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/Animal_echolocation" title="Animal echolocation">Animal echolocation</a></li> <li><a href="/wiki/Waggle_dance" title="Waggle dance">Waggle dance</a></li> <li><a href="/wiki/Jamming_avoidance_response" title="Jamming avoidance response">Jamming avoidance response</a></li> <li><a href="/wiki/Vision_in_toads" title="Vision in toads">Vision in toads</a></li> <li><a href="/wiki/Frog_hearing_and_communication" title="Frog hearing and communication">Frog hearing and communication</a></li> <li><a href="/wiki/Infrared_sensing_in_snakes" title="Infrared sensing in snakes">Infrared sensing in snakes</a></li> <li><a href="/wiki/Caridoid_escape_reaction" title="Caridoid escape reaction">Caridoid escape reaction</a></li> <li><a href="/wiki/Vocal_learning" title="Vocal learning">Vocal learning</a></li> <li><a href="/wiki/Surface_wave_detection" class="mw-redirect" title="Surface wave detection">Surface wave detection</a></li> <li><a href="/wiki/Electroreception" class="mw-redirect" title="Electroreception">Electroreception</a></li> <li><a href="/wiki/Mechanoreception" class="mw-redirect" title="Mechanoreception">Mechanoreception</a> <ul><li><a href="/wiki/Lateral_line" title="Lateral line">Lateral line</a></li></ul></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Neuroethology" title="Category:Neuroethology">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Neuroethology" class="extiw" title="commons:Category:Neuroethology">Commons</a></b></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="Neuroscience" style="padding:3px"><table 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neuroscience">Cellular neuroscience</a></li> <li><a href="/wiki/Computational_neuroscience" title="Computational neuroscience">Computational neuroscience</a></li> <li><a href="/wiki/Connectomics" title="Connectomics">Connectomics</a></li> <li><a href="/wiki/Imaging_genetics" title="Imaging genetics">Imaging genetics</a></li> <li><a href="/wiki/Integrative_neuroscience" title="Integrative neuroscience">Integrative neuroscience</a></li> <li><a href="/wiki/Molecular_neuroscience" title="Molecular neuroscience">Molecular neuroscience</a></li> <li><a href="/wiki/Neural_decoding" title="Neural decoding">Neural decoding</a></li> <li><a href="/wiki/Neural_engineering" title="Neural engineering">Neural engineering</a></li> <li><a href="/wiki/Neuroanatomy" title="Neuroanatomy">Neuroanatomy</a></li> <li><a href="/wiki/Neurobiology" class="mw-redirect" title="Neurobiology">Neurobiology</a></li> <li><a href="/wiki/Neurochemistry" title="Neurochemistry">Neurochemistry</a></li> <li><a href="/wiki/Neuroendocrinology" title="Neuroendocrinology">Neuroendocrinology</a></li> <li><a href="/wiki/Neurogenetics" title="Neurogenetics">Neurogenetics</a></li> <li><a href="/wiki/Neuroinformatics" title="Neuroinformatics">Neuroinformatics</a></li> <li><a href="/wiki/Neurometrics" title="Neurometrics">Neurometrics</a></li> <li><a href="/wiki/Neuromorphology" title="Neuromorphology">Neuromorphology</a></li> <li><a href="/wiki/Neurophysics" title="Neurophysics">Neurophysics</a></li> <li><a href="/wiki/Neurophysiology" title="Neurophysiology">Neurophysiology</a></li> <li><a href="/wiki/Systems_neuroscience" title="Systems neuroscience">Systems neuroscience</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Gray739.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/130px-Gray739.png" decoding="async" width="130" height="99" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/195px-Gray739.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/260px-Gray739.png 2x" data-file-width="500" data-file-height="379" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Clinical_neuroscience" title="Clinical neuroscience">Clinical<br />neuroscience</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/Behavioral_neurology" title="Behavioral neurology">Behavioral neurology</a></li> <li><a href="/wiki/Clinical_neurophysiology" title="Clinical neurophysiology">Clinical neurophysiology</a></li> <li><a href="/wiki/Epileptology" class="mw-redirect" title="Epileptology">Epileptology</a></li> <li><a href="/wiki/Neurocardiology" title="Neurocardiology">Neurocardiology</a></li> <li><a href="/wiki/Neuroepidemiology" title="Neuroepidemiology">Neuroepidemiology</a></li> <li><a href="/wiki/Enteric_nervous_system#Function" title="Enteric nervous system">Neurogastroenterology</a></li> <li><a href="/wiki/Neuroimmunology" title="Neuroimmunology">Neuroimmunology</a></li> <li><a href="/wiki/Neurointensive_care" title="Neurointensive care">Neurointensive care</a></li> <li><a href="/wiki/Neurology" title="Neurology">Neurology</a></li> <li><a href="/wiki/Neuro-oncology" title="Neuro-oncology">Neuro-oncology</a></li> <li><a href="/wiki/Neuro-ophthalmology" title="Neuro-ophthalmology">Neuro-ophthalmology</a></li> <li><a href="/wiki/Neuropathology" title="Neuropathology">Neuropathology</a></li> <li><a href="/wiki/Neuropharmacology" title="Neuropharmacology">Neuropharmacology</a></li> <li><a href="/wiki/Neuroprosthetics" title="Neuroprosthetics">Neuroprosthetics</a></li> <li><a href="/wiki/Neuropsychiatry" title="Neuropsychiatry">Neuropsychiatry</a></li> <li><a href="/wiki/Neuroradiology" title="Neuroradiology">Neuroradiology</a></li> <li><a href="/wiki/Neurorehabilitation" title="Neurorehabilitation">Neurorehabilitation</a></li> <li><a href="/wiki/Neurosurgery" title="Neurosurgery">Neurosurgery</a></li> <li><a href="/wiki/Neurotology" title="Neurotology">Neurotology</a></li> <li><a href="/wiki/Neurovirology" title="Neurovirology">Neurovirology</a></li> <li><a href="/wiki/Nutritional_neuroscience" title="Nutritional neuroscience">Nutritional neuroscience</a></li> <li><a href="/wiki/Psychiatry" title="Psychiatry">Psychiatry</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">Cognitive<br />neuroscience</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/Affective_neuroscience" title="Affective neuroscience">Affective neuroscience</a></li> <li><a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">Behavioral neuroscience</a></li> <li><a href="/wiki/Chronobiology" title="Chronobiology">Chronobiology</a></li> <li><a href="/wiki/Molecular_cellular_cognition" title="Molecular cellular cognition">Molecular cellular cognition</a></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a></li> <li><a href="/wiki/Neurolinguistics" title="Neurolinguistics">Neurolinguistics</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Sensory_neuroscience" title="Sensory neuroscience">Sensory neuroscience</a></li> <li><a href="/wiki/Social_cognitive_neuroscience" title="Social cognitive neuroscience">Social cognitive neuroscience</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Interdisciplinary<br />fields</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/Consumer_neuroscience" title="Consumer neuroscience">Consumer neuroscience</a></li> <li><a href="/wiki/Cultural_neuroscience" title="Cultural neuroscience">Cultural neuroscience</a></li> <li><a href="/wiki/Educational_neuroscience" title="Educational neuroscience">Educational neuroscience</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Global_neurosurgery" title="Global neurosurgery">Global neurosurgery</a></li> <li><a href="/wiki/Neuroanthropology" title="Neuroanthropology">Neuroanthropology</a></li> <li><a href="/wiki/Neural_engineering" title="Neural engineering">Neural engineering</a></li> <li><a href="/wiki/Neurotechnology" title="Neurotechnology">Neurobiotics</a></li> <li><a href="/wiki/Neurocriminology" title="Neurocriminology">Neurocriminology</a></li> <li><a href="/wiki/Neuroeconomics" title="Neuroeconomics">Neuroeconomics</a></li> <li><a href="/wiki/Neuroepistemology" title="Neuroepistemology">Neuroepistemology</a></li> <li><a href="/wiki/Neuroesthetics" title="Neuroesthetics">Neuroesthetics</a></li> <li><a href="/wiki/Neuroethics" title="Neuroethics">Neuroethics</a></li> <li><a class="mw-selflink selflink">Neuroethology</a></li> <li><a href="/wiki/Neurohistory" title="Neurohistory">Neurohistory</a></li> <li><a href="/wiki/Neurolaw" title="Neurolaw">Neurolaw</a></li> <li><a href="/wiki/Neuromarketing" title="Neuromarketing">Neuromarketing</a></li> <li><a href="/wiki/Neuromorphic_engineering" class="mw-redirect" title="Neuromorphic engineering">Neuromorphic engineering</a></li> <li><a href="/wiki/Neurophenomenology" title="Neurophenomenology">Neurophenomenology</a></li> <li><a href="/wiki/Neurophilosophy" title="Neurophilosophy">Neurophilosophy</a></li> <li><a href="/wiki/Neuropolitics" title="Neuropolitics">Neuropolitics</a></li> <li><a href="/wiki/Neurorobotics" title="Neurorobotics">Neurorobotics</a></li> <li><a href="/wiki/Neuroscience_of_religion" title="Neuroscience of religion">Neurotheology</a></li> <li><a href="/wiki/Paleoneurobiology" title="Paleoneurobiology">Paleoneurobiology</a></li> <li><a href="/wiki/Social_neuroscience" title="Social neuroscience">Social neuroscience</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Concepts</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/Brain%E2%80%93computer_interface" title="Brain–computer interface">Brain–computer interface</a></li> <li><a href="/wiki/Development_of_the_nervous_system" title="Development of the nervous system">Development of the nervous system</a></li> <li><a href="/wiki/Artificial_neural_network" class="mw-redirect" title="Artificial neural network">Neural network (artificial)</a></li> <li><a href="/wiki/Neural_circuit" title="Neural circuit">Neural network (biological)</a></li> <li><a href="/wiki/Detection_theory" title="Detection theory">Detection theory</a></li> <li><a href="/wiki/Intraoperative_neurophysiological_monitoring" title="Intraoperative neurophysiological monitoring">Intraoperative neurophysiological monitoring</a></li> <li><a href="/wiki/Neurochip" title="Neurochip">Neurochip</a></li> <li><a href="/wiki/Neurodegenerative_disease" title="Neurodegenerative disease">Neurodegenerative disease</a></li> <li><a href="/wiki/Neurodevelopmental_disorder" title="Neurodevelopmental disorder">Neurodevelopmental disorder</a></li> <li><a href="/wiki/Neurodiversity" title="Neurodiversity">Neurodiversity</a></li> <li><a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a></li> <li><a href="/wiki/Neuroimaging" title="Neuroimaging">Neuroimaging</a></li> <li><a href="/wiki/Neuroimmune_system" title="Neuroimmune system">Neuroimmune system</a></li> <li><a href="/wiki/Neuromanagement" title="Neuromanagement">Neuromanagement</a></li> <li><a href="/wiki/Neuromodulation" title="Neuromodulation">Neuromodulation</a></li> <li><a href="/wiki/Neuroplasticity" title="Neuroplasticity">Neuroplasticity</a></li> <li><a href="/wiki/Neurotechnology" title="Neurotechnology">Neurotechnology</a></li> <li><a href="/wiki/Neurotoxin" title="Neurotoxin">Neurotoxin</a></li> <li><a href="/wiki/Neural_basis_of_self" title="Neural basis of self">Self-awareness</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="3"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Neuroscience" title="Category:Neuroscience">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Neuroscience" class="extiw" title="commons:Category:Neuroscience">Commons</a></b></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="Animal_cognition" style="padding:3px"><table class="nowraplinks hlist 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:Animal_cognition" title="Template:Animal cognition"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Animal_cognition" title="Template talk:Animal cognition"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Animal_cognition" title="Special:EditPage/Template:Animal cognition"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Animal_cognition" style="font-size:114%;margin:0 4em"><a href="/wiki/Animal_cognition" title="Animal cognition">Animal cognition</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cognition</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/Animal_communication" title="Animal communication">Animal communication</a></li> <li><a href="/wiki/Animal_consciousness" title="Animal consciousness">Animal consciousness</a></li> <li><a href="/wiki/Animal_language" title="Animal language">Animal language</a></li> <li><a href="/wiki/Cognitive_bias_in_animals" title="Cognitive bias in animals">Cognitive bias in animals</a></li> <li><a href="/wiki/Cognitive_ethology" title="Cognitive ethology">Cognitive ethology</a></li> <li><a href="/wiki/Comparative_cognition" title="Comparative cognition">Comparative cognition</a></li> <li><a href="/wiki/Emotion_in_animals" title="Emotion in animals">Emotion in animals</a></li> <li><a href="/wiki/Insect_cognition" title="Insect cognition">Insect</a></li> <li><a href="/wiki/Mirror_test" title="Mirror test">Mirror test</a></li> <li><a class="mw-selflink selflink">Neuroethology</a></li> <li><a href="/wiki/Observational_learning" title="Observational learning">Observational learning</a></li> <li><a href="/wiki/Primate_archaeology" title="Primate archaeology">Primate archaeology</a></li> <li><a href="/wiki/Tool_use_by_non-humans" title="Tool use by non-humans">Tool use by non-humans</a> <ul><li><a href="/wiki/Tool_use_by_sea_otters" title="Tool use by sea otters">sea otters</a></li></ul></li> <li><a href="/wiki/Vocal_learning" title="Vocal learning">Vocal learning</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Intelligence</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/Bird_intelligence" title="Bird intelligence">Bird</a> <ul><li><a href="/wiki/Pigeon_intelligence" title="Pigeon intelligence">Pigeon</a></li> <li><a href="/wiki/Talking_bird" title="Talking bird">talking</a></li></ul></li> <li><a href="/wiki/Cat_intelligence" title="Cat intelligence">Cat</a></li> <li><a href="/wiki/Cephalopod_intelligence" title="Cephalopod intelligence">Cephalopod</a></li> <li><a href="/wiki/Cetacean_intelligence" title="Cetacean intelligence">Cetacean</a></li> <li><a href="/wiki/Dog_intelligence" title="Dog intelligence">Dog</a></li> <li><a href="/wiki/Elephant_cognition" title="Elephant cognition">Elephant</a></li> <li><a href="/wiki/Fish_intelligence" title="Fish intelligence">Fish</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate</a> <ul><li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Hominid</a></li></ul></li> <li><a href="/wiki/Swarm_intelligence" title="Swarm intelligence">Swarm</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Pain</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/Pain_in_amphibians" title="Pain in amphibians">Pain in amphibians</a></li> <li><a href="/wiki/Pain_in_animals" title="Pain in animals">Pain in animals</a></li> <li><a href="/wiki/Pain_in_cephalopods" title="Pain in cephalopods">Pain in cephalopods</a></li> <li><a href="/wiki/Pain_in_crustaceans" title="Pain in crustaceans">Pain in crustaceans</a></li> <li><a href="/wiki/Pain_in_fish" title="Pain in fish">Pain in fish</a></li> <li><a href="/wiki/Pain_in_invertebrates" title="Pain in invertebrates">Pain in invertebrates</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Relation to brain</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/Brain_size" title="Brain size">Brain size</a></li> <li><a href="/wiki/Brain-to-body_mass_ratio" class="mw-redirect" title="Brain-to-body mass ratio">Brain-to-body mass ratio</a></li> <li><a href="/wiki/Encephalization_quotient" title="Encephalization quotient">Encephalization quotient</a></li> <li><a href="/wiki/Neuroscience_and_intelligence" title="Neuroscience and intelligence">Neuroscience and intelligence</a></li> <li><a href="/wiki/List_of_animals_by_number_of_neurons" title="List of animals by number of neurons">Number of neurons</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Animal_cognition" title="Category:Animal cognition">Category</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="Ethology" style="padding:3px"><table class="nowraplinks hlist 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:Ethology" title="Template:Ethology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Ethology" title="Template talk:Ethology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Ethology" title="Special:EditPage/Template:Ethology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ethology" style="font-size:114%;margin:0 4em"><a href="/wiki/Ethology" title="Ethology">Ethology</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Branches</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/Animal_cognition" title="Animal cognition">Animal cognition</a></li> <li><a href="/wiki/Animal_communication" title="Animal communication">Animal communication</a></li> <li><a href="/wiki/Animal_consciousness" title="Animal consciousness">Animal consciousness</a></li> <li><a href="/wiki/Animal_culture" title="Animal culture">Animal culture</a></li> <li><a href="/wiki/Animal_sexual_behaviour" title="Animal sexual behaviour">Animal sexual behaviour</a></li> <li><a href="/wiki/Animal_welfare_science" title="Animal welfare science">Animal welfare science</a></li> <li><a href="/wiki/Anthrozoology" title="Anthrozoology">Anthrozoology</a></li> <li><a href="/wiki/Bee_learning_and_communication" title="Bee learning and communication">Bee learning and communication</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behavioural ecology</a></li> <li><a href="/wiki/Behavioral_endocrinology" title="Behavioral endocrinology">Behavioral endocrinology</a></li> <li><a href="/wiki/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Breed" title="Breed">Breed</a></li> <li><a href="/wiki/Cognitive_ethology" title="Cognitive ethology">Cognitive ethology</a></li> <li><a href="/wiki/Comfort_behaviour_in_animals" title="Comfort behaviour in animals">Comfort behaviour</a> <ul><li><a href="/wiki/Personal_grooming" title="Personal grooming">Grooming</a></li></ul></li> <li><a href="/wiki/Comparative_psychology" title="Comparative psychology">Comparative psychology</a></li> <li><a href="/wiki/Emotion_in_animals" title="Emotion in animals">Emotion in animals</a></li> <li><a href="/wiki/Ethogram" title="Ethogram">Ethogram</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">Feeding</a></li> <li><a href="/wiki/Hover_(behaviour)" title="Hover (behaviour)">Hover</a></li> <li><a href="/wiki/Human_ethology" title="Human ethology">Human ethology</a></li> <li><a href="/wiki/Instinct" title="Instinct">Instinct</a></li> <li><a href="/wiki/Learning" title="Learning">Learning</a></li> <li><a class="mw-selflink selflink">Neuroethology</a></li> <li><a href="/wiki/Pain_in_animals" title="Pain in animals">Pain in animals</a></li> <li><a href="/wiki/Philosophical_ethology" title="Philosophical ethology">Philosophical ethology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Stereotypy_(non-human)" title="Stereotypy (non-human)">Stereotypy</a></li> <li><a href="/wiki/Structures_built_by_animals" title="Structures built by animals">Structures</a> <ul><li><a href="/wiki/Beehive" title="Beehive">Hive</a></li> <li><a href="/wiki/Honeycomb_structure" title="Honeycomb structure">Honeycomb</a></li> <li><a href="/wiki/Nest" title="Nest">Nest</a></li> <li><a href="/wiki/Nesting_instinct" title="Nesting instinct">Instinct</a></li></ul></li> <li><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarm</a></li> <li><a href="/wiki/Tool_use_by_non-humans" title="Tool use by non-humans">Tool use by non-humans</a></li> <li><a href="/wiki/Zoosemiotics" title="Zoosemiotics">Zoosemiotics</a></li> <li><a href="/wiki/Zoomusicology" title="Zoomusicology">Zoomusicology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ethologists</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/Patrick_Bateson" title="Patrick Bateson">Patrick Bateson</a></li> <li><a href="/wiki/Marc_Bekoff" title="Marc Bekoff">Marc Bekoff</a></li> <li><a href="/wiki/Donald_Broom" title="Donald Broom">Donald Broom</a></li> <li><a href="/wiki/John_B._Calhoun" title="John B. Calhoun">John B. Calhoun</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></li> <li><a href="/wiki/Marian_Dawkins" title="Marian Dawkins">Marian Dawkins</a></li> <li><a href="/wiki/Richard_Dawkins" title="Richard Dawkins">Richard Dawkins</a></li> <li><a href="/wiki/Iren%C3%A4us_Eibl-Eibesfeldt" title="Irenäus Eibl-Eibesfeldt">Irenäus Eibl-Eibesfeldt</a></li> <li><a href="/wiki/Dian_Fossey" title="Dian Fossey">Dian Fossey</a></li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a></li> <li><a href="/wiki/Jane_Goodall" title="Jane Goodall">Jane Goodall</a></li> <li><a href="/wiki/Heini_Hediger" title="Heini Hediger">Heini Hediger</a></li> <li><a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Desmond_Morris" title="Desmond Morris">Desmond Morris</a></li> <li><a href="/wiki/Thomas_Sebeok" title="Thomas Sebeok">Thomas Sebeok</a></li> <li><a href="/wiki/William_Homan_Thorpe" title="William Homan Thorpe">William Homan Thorpe</a></li> <li><a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Nikolaas Tinbergen</a></li> <li><a href="/wiki/Jakob_von_Uexk%C3%BCll" class="mw-redirect" title="Jakob von Uexküll">Jakob von Uexküll</a></li> <li><a href="/wiki/Wolfgang_Wickler" title="Wolfgang Wickler">Wolfgang Wickler</a></li> <li><a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a></li> <li><a href="/wiki/Solly_Zuckerman" class="mw-redirect" title="Solly Zuckerman">Solly Zuckerman</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Societies</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/Association_for_the_Study_of_Animal_Behaviour" title="Association for the Study of Animal Behaviour">Association for the Study of Animal Behaviour</a></li> <li><a href="/wiki/International_Society_for_Applied_Ethology" title="International Society for Applied Ethology">International Society for Applied Ethology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Journals</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/Animal_Behaviour_(journal)" title="Animal Behaviour (journal)">Animal Behaviour</a></i></li> <li><i><a href="/wiki/Animal_Cognition" title="Animal Cognition">Animal Cognition</a></i></li> <li><i><a href="/wiki/Animal_Welfare_(journal)" title="Animal Welfare (journal)">Animal Welfare</a></i></li> <li><i><a href="/wiki/Behavioral_Ecology_(journal)" title="Behavioral Ecology (journal)">Behavioral Ecology</a></i></li> <li><i><a href="/wiki/Behaviour_(journal)" title="Behaviour (journal)">Behaviour</a></i></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Ethology" title="Category:Ethology">Category</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="Zoology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="background:"><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:Zoology" title="Template:Zoology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Zoology" title="Template talk:Zoology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Zoology" title="Special:EditPage/Template:Zoology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Zoology" style="font-size:114%;margin:0 4em"><a href="/wiki/Zoology" title="Zoology">Zoology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2" style="background:"><div> <ul><li><a href="/wiki/Outline_of_zoology" title="Outline of zoology">Outline</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%">Branches</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/Anthrozoology" title="Anthrozoology">Anthrozoology</a></li> <li><a href="/wiki/Arthropodology" title="Arthropodology">Arthropodology</a> (<a href="/wiki/Arachnology" title="Arachnology">Arachnology</a> (<a href="/wiki/Acarology" title="Acarology">Acarology</a>)</li> <li><a href="/wiki/Carcinology" title="Carcinology">Carcinology</a></li> <li><a href="/wiki/Entomology" title="Entomology">Entomology</a> (<a href="/wiki/Coleopterology" title="Coleopterology">Coleopterology</a></li> <li><a href="/wiki/Lepidopterology" title="Lepidopterology">Lepidopterology</a></li> <li><a href="/wiki/Melittology" title="Melittology">Melittology</a></li> <li><a href="/wiki/Myrmecology" title="Myrmecology">Myrmecology</a></li> <li><a href="/wiki/Orthopterology" title="Orthopterology">Orthopterology</a>)</li> <li><a href="/wiki/Myriapodology" title="Myriapodology">Myriapodology</a>)</li> <li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological anthropology</a></li> <li><a href="/wiki/Bryozoology" title="Bryozoology">Bryozoology</a></li> <li><a href="/wiki/Cnidariology" title="Cnidariology">Cnidariology</a></li> <li><a href="/wiki/Ethnozoology" title="Ethnozoology">Ethnozoology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Helminthology" title="Helminthology">Helminthology</a></li> <li><a href="/wiki/Herpetology" title="Herpetology">Herpetology</a> (<a href="/wiki/Batrachology" title="Batrachology">Batrachology</a></li> <li><a href="/wiki/Ophiology" class="mw-redirect" title="Ophiology">Ophiology</a></li> <li><a href="/wiki/Testudinology" class="mw-redirect" title="Testudinology">Testudinology</a>)</li> <li><a href="/wiki/Ichthyology" title="Ichthyology">Ichthyology</a></li> <li><a href="/wiki/Malacology" title="Malacology">Malacology</a> (<a href="/wiki/Conchology" title="Conchology">Conchology</a></li> <li><a href="/wiki/Teuthology" title="Teuthology">Teuthology</a>)</li> <li><a href="/wiki/Mammalogy" title="Mammalogy">Mammalogy</a> (<a href="/wiki/Cetology" title="Cetology">Cetology</a></li> <li><a href="/wiki/Cynology" title="Cynology">Cynology</a></li> <li><a href="/wiki/Felinology" title="Felinology">Felinology</a></li> <li><a href="/wiki/Hippology" title="Hippology">Hippology</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a>)</li> <li><a href="/wiki/Nematology" title="Nematology">Nematology</a></li> <li><a class="mw-selflink selflink">Neuroethology</a></li> <li><a href="/wiki/Ornithology" title="Ornithology">Ornithology</a></li> <li><a href="/wiki/Parasitology" title="Parasitology">Parasitology</a></li> <li><a href="/wiki/Paleozoology" title="Paleozoology">Paleozoology</a></li> <li><a href="/wiki/Planktology" title="Planktology">Planktology</a></li> <li><a href="/wiki/Zooarchaeology" title="Zooarchaeology">Zooarchaeology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Animal" title="Animal">Animal</a> groups</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/Sponge" title="Sponge">Porifera</a></li> <li><a href="/wiki/Ctenophora" title="Ctenophora">Ctenophora</a></li> <li><a href="/wiki/Placozoa" title="Placozoa">Placozoa</a></li> <li><a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a></li> <li><a href="/wiki/Xenacoelomorpha" title="Xenacoelomorpha">Xenacoelomorpha</a></li> <li><a href="/wiki/Ambulacraria" title="Ambulacraria">Ambulacraria</a></li> <li><a href="/wiki/Chordate" title="Chordate">Chordata</a></li> <li><a href="/wiki/Aschelminthes" class="mw-redirect" title="Aschelminthes">Aschelminthes</a></li> <li><a href="/wiki/Arthropod" title="Arthropod">Arthropoda</a></li> <li><a href="/wiki/Flatworm" title="Flatworm">Platyhelminthes</a></li> <li><a href="/wiki/Mollusca" title="Mollusca">Mollusca</a></li> <li><a href="/wiki/Annelid" title="Annelid">Annelida</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Anatomy" title="Anatomy">Animal anatomy</a><br /><a href="/wiki/Morphology_(biology)" title="Morphology (biology)">Animal morphology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Histology" title="Histology">Histology</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/Tissue_(biology)#Animal_tissue" title="Tissue (biology)">Tissues</a> <ul><li><a href="/wiki/Epithelium" title="Epithelium">Epithelial tissue</a></li> <li><a href="/wiki/Muscle_tissue" class="mw-redirect" title="Muscle tissue">Muscular tissue</a></li> <li><a href="/wiki/Nervous_tissue" title="Nervous tissue">Nervous tissue</a></li> <li><a href="/wiki/Connective_tissue" title="Connective tissue">Connective tissues</a></li> <li><a href="/wiki/Mineralized_tissues" title="Mineralized tissues">Mineralized tissues</a></li></ul></li> <li><a href="/wiki/Molecular_anatomy" title="Molecular anatomy">Molecular anatomy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Comparative_anatomy" title="Comparative anatomy">Anatomy <br /> and morphology</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/Mammal#Anatomy" title="Mammal">Mammalian anatomy and morphology</a> <ul><li><a href="/wiki/Human_anatomy" class="mw-redirect" title="Human anatomy">Human</a></li> <li><a href="/wiki/Dog_anatomy" title="Dog anatomy">Dog</a></li> <li><a href="/wiki/Elephant#Anatomy_and_morphology" title="Elephant">Elephant</a></li> <li><a href="/wiki/Cat_anatomy" title="Cat anatomy">Cat</a></li></ul></li> <li><a href="/wiki/Bird_anatomy" title="Bird anatomy">Bird anatomy</a></li> <li><a href="/wiki/Fish_anatomy" title="Fish anatomy">Fish anatomy</a> <ul><li><a href="/wiki/Shark_anatomy" title="Shark anatomy">Shark anatomy</a></li></ul></li> <li><a href="/wiki/Spider_anatomy" title="Spider anatomy">Spider anatomy</a></li> <li><a href="/wiki/Insect_morphology" title="Insect morphology">Insect morphology</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/Physiology#Animals" title="Physiology">Animal physiology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General physiology</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/Respiration_(physiology)" title="Respiration (physiology)">Respiration</a> <ul><li><a href="/wiki/Respiratory_system" title="Respiratory system">Respiratory system</a></li> <li><a href="/wiki/Breathing" title="Breathing">Breathing</a></li> <li><a href="/wiki/Gas_exchange" title="Gas exchange">Gas exchange</a></li> <li><a href="/wiki/Respiratory_pigment" title="Respiratory pigment">Respiratory pigment</a></li> <li><a href="/wiki/Cellular_respiration" title="Cellular respiration">Cellular respiration</a></li></ul></li> <li><a href="/wiki/Vascular_system" class="mw-redirect" title="Vascular system">Vascular system</a> <ul><li><a href="/wiki/Blood" title="Blood">Blood</a></li> <li><a href="/wiki/Lymph" title="Lymph">Lymph</a></li> <li><a href="/wiki/Blood_vessel" title="Blood vessel">Blood vessels</a> <ul><li><a href="/wiki/Arteries" class="mw-redirect" title="Arteries">Arteries</a></li> <li><a href="/wiki/Vein" title="Vein">Veins</a></li> <li><a href="/wiki/Capillaries" class="mw-redirect" title="Capillaries">Capillaries</a></li></ul></li> <li><a href="/wiki/Heart" title="Heart">Heart</a></li> <li><a href="/wiki/Water_vascular_system" title="Water vascular system">Water vascular system</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">By species</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/Insect_physiology" title="Insect physiology">Insect physiology</a></li> <li><a href="/wiki/Fish_physiology" title="Fish physiology">Fish physiology</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%"><a href="/wiki/List_of_zoologists" class="mw-redirect" title="List of zoologists">Zoologists</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/Karl_Ernst_von_Baer" title="Karl Ernst von Baer">Karl Ernst von Baer</a></li> <li><a href="/wiki/Georges_Cuvier" title="Georges Cuvier">Georges Cuvier</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></li> <li><a href="/wiki/Jean-Henri_Fabre" title="Jean-Henri Fabre">Jean-Henri Fabre</a></li> <li><a href="/wiki/William_Kirby_(entomologist)" title="William Kirby (entomologist)">William Kirby</a></li> <li><a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a></li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a></li> <li><a href="/wiki/Thomas_Say" title="Thomas Say">Thomas Say</a></li> <li><a href="/wiki/Jakob_von_Uexk%C3%BCll" class="mw-redirect" title="Jakob von Uexküll">Jakob von Uexküll</a></li> <li><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="background:;width:1%">History</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/History_of_zoology_through_1859" title="History of zoology through 1859">Pre-Darwin</a></li> <li><a href="/wiki/History_of_zoology_(1859%E2%80%93present)" title="History of zoology (1859–present)">Post-Darwin</a></li> <li><a href="/wiki/Timeline_of_zoology" title="Timeline of zoology">Timeline of zoology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="background:"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" 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