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Eyespot (mimicry) - Wikipedia

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<div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Evolution</span> </div> </a> <ul id="toc-Evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Artificial_eyespots" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Artificial_eyespots"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Artificial eyespots</span> </div> </a> <ul id="toc-Artificial_eyespots-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">5</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"> 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Available in 19 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-19" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">19 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%B9%D9%8A%D9%8A%D9%86%D8%A9_(%D8%AA%D9%84%D9%88%D9%86)" title="عيينة (تلون) – Arabic" lang="ar" hreflang="ar" data-title="عيينة (تلون)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9B%D1%8A%D0%B6%D0%BB%D0%B8%D0%B2%D0%BE_%D0%BE%D0%BA%D0%BE" title="Лъжливо око – Bulgarian" lang="bg" hreflang="bg" data-title="Лъжливо око" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Ocel_(mimetisme)" title="Ocel (mimetisme) – Catalan" lang="ca" hreflang="ca" data-title="Ocel (mimetisme)" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Augenfleck_(Abwehr)" title="Augenfleck (Abwehr) – German" lang="de" hreflang="de" data-title="Augenfleck (Abwehr)" 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/Ocelo_(mimetismo)" title="Ocelo (mimetismo) – Spanish" lang="es" hreflang="es" data-title="Ocelo (mimetismo)" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Okulmakulo" title="Okulmakulo – Esperanto" lang="eo" hreflang="eo" data-title="Okulmakulo" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%86%D8%B4%D9%85_%D8%AF%D8%B1%D9%88%D8%BA%DB%8C%D9%86" title="چشم دروغین – Persian" lang="fa" hreflang="fa" data-title="چشم دروغین" 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/Ocelle_(tache)" title="Ocelle (tache) – French" lang="fr" hreflang="fr" data-title="Ocelle (tache)" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Ocelo_(mimetismo)" title="Ocelo (mimetismo) – Galician" lang="gl" hreflang="gl" data-title="Ocelo (mimetismo)" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Ocello_(mimetismo)" title="Ocello (mimetismo) – Italian" lang="it" hreflang="it" data-title="Ocello (mimetismo)" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Bintik_mata" title="Bintik mata – Malay" lang="ms" hreflang="ms" data-title="Bintik mata" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Oogvlek" title="Oogvlek – Dutch" lang="nl" hreflang="nl" data-title="Oogvlek" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%9C%BC%E7%8A%B6%E7%B4%8B" title="眼状紋 – Japanese" lang="ja" hreflang="ja" data-title="眼状紋" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Ocelo_(mimetismo)" title="Ocelo (mimetismo) – Portuguese" lang="pt" hreflang="pt" data-title="Ocelo (mimetismo)" 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%93%D0%BB%D0%B0%D0%B7%D1%87%D0%B0%D1%82%D0%BE%D0%B5_%D0%BF%D1%8F%D1%82%D0%BD%D0%BE" title="Глазчатое пятно – Russian" lang="ru" hreflang="ru" data-title="Глазчатое пятно" 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/Eye-spot_(mimicry)" title="Eye-spot (mimicry) – Simple English" lang="en-simple" hreflang="en-simple" data-title="Eye-spot (mimicry)" 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-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Silm%C3%A4laikku" title="Silmälaikku – Finnish" lang="fi" hreflang="fi" data-title="Silmälaikku" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%9C%BC%E6%96%91" title="眼斑 – Cantonese" lang="yue" hreflang="yue" data-title="眼斑" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E7%9C%BC%E7%8B%80%E6%96%91%E9%BB%9E" title="眼狀斑點 – Chinese" lang="zh" 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data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> <div id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><span typeof="mw:File"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. 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Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><p class="mw-empty-elt"> </p> <div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Eye-like marking used for mimicry or distraction GT </div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Mycalesis_patnia.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Mycalesis_patnia.jpg/300px-Mycalesis_patnia.jpg" decoding="async" width="300" height="265" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Mycalesis_patnia.jpg/450px-Mycalesis_patnia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/6/66/Mycalesis_patnia.jpg 2x" data-file-width="469" data-file-height="415" /></a><figcaption>Many butterflies, such as this <a href="/wiki/Mycalesis_patnia" title="Mycalesis patnia">gladeye bushbrown</a> (<i>Mycalesis patnia</i>), have eyespots on their wings.</figcaption></figure> <p>An <b>eyespot</b> (sometimes <b>ocellus</b>) is an <a href="/wiki/Eye" title="Eye">eye</a>-like marking. They are found in butterflies, reptiles, cats, birds and fish. </p><p>Eyespots could be explained in at least three different ways. They may be a form of <a href="/wiki/Mimicry" title="Mimicry">mimicry</a> in which a spot on the body of an animal resembles an eye of a different animal, to deceive potential <a href="/wiki/Predator" class="mw-redirect" title="Predator">predator</a> or prey species. They may be a form of <a href="/wiki/Automimicry" title="Automimicry">self-mimicry</a>, to draw a predator's attention away from the prey's most vulnerable body parts. Or they may serve to make the prey appear inedible or dangerous. Eyespot markings may play a role in intraspecies communication or courtship; a well-known example is the eyespots on a <a href="/wiki/Peacock" class="mw-redirect" title="Peacock">peacock</a>'s display feathers. </p><p>The pattern-forming biological process (<a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenesis</a>) of eyespots in a wide variety of animals is controlled by a small number of <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">genes active in embryonic development</a>, including the genes called <a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a>, <a href="/wiki/Distal-less" class="mw-redirect" title="Distal-less">Distal-less</a>, <a href="/wiki/Hedgehog_(gene)" class="mw-redirect" title="Hedgehog (gene)">Hedgehog</a>, <a href="/wiki/Antennapedia" title="Antennapedia">Antennapedia</a>, and the <a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a>. </p><p>Artificial eyespots have been shown to reduce predation of cattle by lions. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Zoological_distribution">Zoological distribution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=1" title="Edit section: Zoological distribution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="In_butterflies_and_moths">In butterflies and moths</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=2" title="Edit section: In butterflies and moths"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Antheraea_polyphemus_-_Polyphemus_Moth.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Antheraea_polyphemus_-_Polyphemus_Moth.jpg/220px-Antheraea_polyphemus_-_Polyphemus_Moth.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Antheraea_polyphemus_-_Polyphemus_Moth.jpg/330px-Antheraea_polyphemus_-_Polyphemus_Moth.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Antheraea_polyphemus_-_Polyphemus_Moth.jpg/440px-Antheraea_polyphemus_-_Polyphemus_Moth.jpg 2x" data-file-width="1400" data-file-height="1050" /></a><figcaption><a href="/wiki/Antheraea_polyphemus" title="Antheraea polyphemus">Polyphemus moth</a></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Automeris_ioPCCA20040704-2975AB1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1d/Automeris_ioPCCA20040704-2975AB1.jpg/220px-Automeris_ioPCCA20040704-2975AB1.jpg" decoding="async" width="220" height="121" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1d/Automeris_ioPCCA20040704-2975AB1.jpg/330px-Automeris_ioPCCA20040704-2975AB1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1d/Automeris_ioPCCA20040704-2975AB1.jpg/440px-Automeris_ioPCCA20040704-2975AB1.jpg 2x" data-file-width="1024" data-file-height="564" /></a><figcaption><i><a href="/wiki/Automeris_io" title="Automeris io">Io moth</a></i></figcaption></figure> <p>The eye-like markings in some <a href="/wiki/Lepidoptera" title="Lepidoptera">butterflies and moths</a> and certain other insects, as well as birds like the <a href="/wiki/Sunbittern" title="Sunbittern">sunbittern</a>, serve functions in addition to <a href="/wiki/Mimic" class="mw-redirect" title="Mimic">mimicry</a>;<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup> indeed, it is unclear whether they actually mimic eyes.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> There is evidence that eyespots in butterflies are <a href="/wiki/Antipredator_adaptation" class="mw-redirect" title="Antipredator adaptation">antipredator adaptations</a>, either in <a href="/wiki/Deimatic_display" class="mw-redirect" title="Deimatic display">deimatic displays</a> to intimidate predators, or to deflect attacks away from vital body parts.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Prudic_Jeon_Cao_Monteiro_pp._73–75_4-0" class="reference"><a href="#cite_note-Prudic_Jeon_Cao_Monteiro_pp._73–75-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> In species such as <i><a href="/wiki/Grayling_(butterfly)" title="Grayling (butterfly)">Hipparchia semele</a></i>, the conspicuous eyespots are hidden at rest to decrease detectability, and only exposed when they believe potential predators are nearby.<sup id="cite_ref-Stevens2005_5-0" class="reference"><a href="#cite_note-Stevens2005-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> Butterfly eyespots can mimic dead leaves for <a href="/wiki/Camouflage" title="Camouflage">camouflage</a> from predators, as seen in <i><a href="/wiki/Bicyclus_anynana" title="Bicyclus anynana">Bicyclus anynana</a></i>; this is a response to a seasonal fall in temperature, causing a shift in selection towards smaller, less conspicuous eyespots among those individuals developing at that time.<sup id="cite_ref-Bhardwaj_Jolander_2020_6-0" class="reference"><a href="#cite_note-Bhardwaj_Jolander_2020-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> Butterfly eyespots may play a role in mate recognition and <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> Sexual selection drives the diversification of eyespots in different species of butterflies, as mates select for characteristics like size and brightness.<sup id="cite_ref-Breuker_Brakefield_2002_8-0" class="reference"><a href="#cite_note-Breuker_Brakefield_2002-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>Some species of <a href="/wiki/Caterpillar" title="Caterpillar">caterpillar</a>, such as those of hawkmoths (<a href="/wiki/Sphingidae" title="Sphingidae">Sphingidae</a>), have eyespots on their anterior abdominal segments. When alarmed, they retract the head and the thoracic segments into the body, leaving the apparently threatening large eyes at the front of the visible part of the body.<sup id="cite_ref-Stevens2005_5-1" class="reference"><a href="#cite_note-Stevens2005-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> </p><p>Butterflies such as the blues (<a href="/wiki/Lycaenidae" title="Lycaenidae">Lycaenidae</a>) have filamentous "tails" at the ends of their wings and nearby patterns of markings, which combine to create a "false head". This <a href="/wiki/Automimicry" title="Automimicry">automimicry</a> misdirects predators such as birds and jumping spiders (<a href="/wiki/Salticidae" class="mw-redirect" title="Salticidae">Salticidae</a>). Spectacular examples occur in the <a href="/wiki/Theclinae" title="Theclinae">hairstreak</a> butterflies; they commonly perch upside down with the false head raised and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this deflects attacks from the insect's head.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_reptiles_and_mammals">In reptiles and mammals</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=3" title="Edit section: In reptiles and mammals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some reptiles, such as the <a href="/wiki/Sand_lizard" title="Sand lizard">sand lizard</a> of Europe, have eyespots; in the sand lizard's case, there is a row of spots along the back, and a row on each side.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> </p><p>Many species of <a href="/wiki/Felidae" title="Felidae">cat</a>, including <a href="/wiki/Geoffroy%27s_cat" title="Geoffroy&#39;s cat">Geoffroy's cats</a>, <a href="/wiki/Jungle_cat" title="Jungle cat">jungle cats</a>, <a href="/wiki/Pampas_cat" title="Pampas cat">pampas cats</a>, and <a href="/wiki/Serval" title="Serval">servals</a>, have white markings, whether spots or bars, on the backs of their ears; it is possible that these signal "follow me" to the young of the species. There may be an <a href="/wiki/Evolutionary_trade-off" class="mw-redirect" title="Evolutionary trade-off">evolutionary trade-off</a> in this case between nocturnal camouflage and intraspecific signalling.<sup id="cite_ref-Sunquist_Sunquist_2017_13-0" class="reference"><a href="#cite_note-Sunquist_Sunquist_2017-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Graipel_Bogoni_2019_14-0" class="reference"><a href="#cite_note-Graipel_Bogoni_2019-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_birds">In birds</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=4" title="Edit section: In birds"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Fan_of_Colours.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/Fan_of_Colours.jpg/220px-Fan_of_Colours.jpg" decoding="async" width="220" height="157" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/85/Fan_of_Colours.jpg/330px-Fan_of_Colours.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/85/Fan_of_Colours.jpg/440px-Fan_of_Colours.jpg 2x" data-file-width="4582" data-file-height="3264" /></a><figcaption><a href="/wiki/Indian_peafowl" title="Indian peafowl">Indian peafowl</a> display</figcaption></figure> <p>Male birds of some species, such as the <a href="/wiki/Peacock" class="mw-redirect" title="Peacock">peacock</a>, have conspicuous eyespots in their plumage, used to signal their quality to <a href="/wiki/Sexual_selection" title="Sexual selection">sexually selecting</a> females. The number of eyespots in a peacock's train predicts his mating success; when a peacock's train is experimentally pruned, females lose interest.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> Several species of <a href="/wiki/Pygmy_owl" title="Pygmy owl">pygmy owl</a> bear false eyes on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_fish">In fish</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=5" title="Edit section: In fish"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></div> <p>Some fish have eyespots. The <a href="/wiki/Foureye_butterflyfish" title="Foureye butterflyfish">foureye butterflyfish</a> gets its name from a large and conspicuous eyespot on each side of the body near the tail. A black vertical bar on the head runs through the true eye, making it hard to see.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> This may deceive <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> in two ways: into attacking the tail rather than the more vulnerable head, and about the fish's likely direction of travel. The foureye butterflyfish eyespot is thus an example of <a href="/wiki/Self-mimicry" class="mw-redirect" title="Self-mimicry">self-mimicry</a>.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> For the same reason, many juvenile fish display eyespots that disappear during their adult phase.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> Some species of fish, like the <a href="/wiki/Synchiropus_picturatus" class="mw-redirect" title="Synchiropus picturatus">spotted mandarin fish</a> and <a href="/wiki/Raja_montagui" class="mw-redirect" title="Raja montagui">spotted ray</a>, maintain their eyespots throughout their adult lives. These eyespots can take a form very similar to those seen in most butterflies, with a focus surrounded by concentric rings of other pigmentation.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> </p> <ul class="gallery mw-gallery-nolines center"> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Deilephila_elpenor_11.JPG" class="mw-file-description" title="Larva of elephant hawkmoth (Deilephila elpenor) displaying eyespots when alarmed"><img alt="Larva of elephant hawkmoth (Deilephila elpenor) displaying eyespots when alarmed" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Deilephila_elpenor_11.JPG/180px-Deilephila_elpenor_11.JPG" decoding="async" width="180" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Deilephila_elpenor_11.JPG/270px-Deilephila_elpenor_11.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Deilephila_elpenor_11.JPG/360px-Deilephila_elpenor_11.JPG 2x" data-file-width="1024" data-file-height="768" /></a></span></div> <div class="gallerytext">Larva of <a href="/wiki/Deilephila_elpenor" title="Deilephila elpenor">elephant hawkmoth</a> (<i>Deilephila elpenor</i>) displaying eyespots when alarmed</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Notodontidae_-_Cerura_vinula.jpg" class="mw-file-description" title="Larva of puss moth (Cerura vinula) in a defensive posture, with black eyespots visible just above its head"><img alt="Larva of puss moth (Cerura vinula) in a defensive posture, with black eyespots visible just above its head" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Notodontidae_-_Cerura_vinula.jpg/180px-Notodontidae_-_Cerura_vinula.jpg" decoding="async" width="180" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Notodontidae_-_Cerura_vinula.jpg/270px-Notodontidae_-_Cerura_vinula.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/66/Notodontidae_-_Cerura_vinula.jpg/360px-Notodontidae_-_Cerura_vinula.jpg 2x" data-file-width="1023" data-file-height="767" /></a></span></div> <div class="gallerytext">Larva of <a href="/wiki/Cerura_vinula" title="Cerura vinula">puss moth</a> (<i>Cerura vinula</i>) in a defensive posture, with black eyespots visible just above its head</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Gray_Hairstreak_(One_more_time...)_(6222138633).jpg" class="mw-file-description" title="Automimicry: gray hairstreak (Strymon melinus) has false head at rear"><img alt="Automimicry: gray hairstreak (Strymon melinus) has false head at rear" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg/180px-Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg" decoding="async" width="180" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg/270px-Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/95/Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg/360px-Gray_Hairstreak_%28One_more_time...%29_%286222138633%29.jpg 2x" data-file-width="1024" data-file-height="1024" /></a></span></div> <div class="gallerytext"><a href="/wiki/Automimicry" title="Automimicry">Automimicry</a>: <a href="/wiki/Strymon_melinus" class="mw-redirect" title="Strymon melinus">gray hairstreak</a> (<i>Strymon melinus</i>) has false head at rear</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Zauneidechse_1469_(detail).jpg" class="mw-file-description" title="Sand lizard (Lacerta agilis) with rows of eyespots"><img alt="Sand lizard (Lacerta agilis) with rows of eyespots" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/Zauneidechse_1469_%28detail%29.jpg/180px-Zauneidechse_1469_%28detail%29.jpg" decoding="async" width="180" height="129" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/Zauneidechse_1469_%28detail%29.jpg/270px-Zauneidechse_1469_%28detail%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/da/Zauneidechse_1469_%28detail%29.jpg/360px-Zauneidechse_1469_%28detail%29.jpg 2x" data-file-width="2523" data-file-height="1804" /></a></span></div> <div class="gallerytext"><a href="/wiki/Sand_lizard" title="Sand lizard">Sand lizard</a> (<i>Lacerta agilis</i>) with rows of eyespots</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Glaucidium_californicum_Verdi_Sierra_Pines_2_(detail).jpg" class="mw-file-description" title="Pygmy owl (Glaucidium californicum) with eyespots behind head"><img alt="Pygmy owl (Glaucidium californicum) with eyespots behind head" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg/161px-Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg" decoding="async" width="161" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg/242px-Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/be/Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg/322px-Glaucidium_californicum_Verdi_Sierra_Pines_2_%28detail%29.jpg 2x" data-file-width="400" data-file-height="447" /></a></span></div> <div class="gallerytext"> <a href="/wiki/Glaucidium_californicum" class="mw-redirect" title="Glaucidium californicum">Pygmy owl</a> (<i>Glaucidium californicum</i>) with eyespots behind head</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Serval_from_back.jpg" class="mw-file-description" title="Serval (Leptailurus serval) from back, ocelli clearly visible on the ears"><img alt="Serval (Leptailurus serval) from back, ocelli clearly visible on the ears" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Serval_from_back.jpg/120px-Serval_from_back.jpg" decoding="async" width="120" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Serval_from_back.jpg/180px-Serval_from_back.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/be/Serval_from_back.jpg/240px-Serval_from_back.jpg 2x" data-file-width="2592" data-file-height="3888" /></a></span></div> <div class="gallerytext"><a href="/wiki/Serval" title="Serval">Serval</a> (<i>Leptailurus serval</i>) from back, ocelli clearly visible on the ears</div> </li> <li class="gallerybox" style="width: 185px"> <div class="thumb" style="width: 180px;"><span typeof="mw:File"><a href="/wiki/File:Chaetodon_capistratus_14zz.jpg" class="mw-file-description" title="Foureye butterflyfish (Chaetodon capistratus) has false eyes near tail and disruptive eye mask concealing eye"><img alt="Foureye butterflyfish (Chaetodon capistratus) has false eyes near tail and disruptive eye mask concealing eye" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Chaetodon_capistratus_14zz.jpg/180px-Chaetodon_capistratus_14zz.jpg" decoding="async" width="180" height="137" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Chaetodon_capistratus_14zz.jpg/270px-Chaetodon_capistratus_14zz.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Chaetodon_capistratus_14zz.jpg/360px-Chaetodon_capistratus_14zz.jpg 2x" data-file-width="673" data-file-height="511" /></a></span></div> <div class="gallerytext"><a href="/wiki/Foureye_butterflyfish" title="Foureye butterflyfish">Foureye butterflyfish</a> (<i>Chaetodon capistratus</i>) has false eyes near tail and <a href="/wiki/Disruptive_eye_mask" title="Disruptive eye mask">disruptive eye mask</a> concealing eye</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="Morphogenesis">Morphogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=6" title="Edit section: Morphogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Butterflywiki.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Butterflywiki.jpg/260px-Butterflywiki.jpg" decoding="async" width="260" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Butterflywiki.jpg/390px-Butterflywiki.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Butterflywiki.jpg/520px-Butterflywiki.jpg 2x" data-file-width="573" data-file-height="386" /></a><figcaption>Plan of a typical butterfly, showing the <a href="/wiki/Morphogenesis" title="Morphogenesis">morphogenetic</a> foci on the wings that create eyespots</figcaption></figure> <p>Butterfly eyespots are formed during <a href="/wiki/Embryogenesis" class="mw-redirect" title="Embryogenesis">embryogenesis</a> as a result of a <a href="/wiki/Morphogenetic" class="mw-redirect" title="Morphogenetic">morphogenetic</a> signalling centre or organizer, called the focus. This induces neighbouring cells to produce specific pigments which pattern the eyespot.<sup id="cite_ref-Breakfield_22-0" class="reference"><a href="#cite_note-Breakfield-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Keys_23-0" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-French_24-0" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> </p><p>Early experiments on eyespot morphogenesis used <a href="/wiki/Cautery" class="mw-redirect" title="Cautery">cautery</a> on the butterfly wing eyespot foci to demonstrate that a long range signaling mechanism or morphogen gradient controlled the formation of eyespots in both space and time.<sup id="cite_ref-French_24-1" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> The findings cannot be explained by a simple source/<a href="/wiki/Diffusion" title="Diffusion">diffusion</a> model,<sup id="cite_ref-French_24-2" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> but could be explained by either a source/threshold model, in which the focus creates the morphogen, or by the sink model, in which the focus generates a gradient by removing a morphogen which was created elsewhere.<sup id="cite_ref-French_24-3" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> Several genes involved in eyespot formation have been identified that can fit into these models, but only two of them have been functionally tested. These genes are the <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factor</a> Distalless (Dll) and the <a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">ligand</a> (a signalling substance that binds a <a href="/wiki/Cell_surface_receptor" title="Cell surface receptor">cell surface receptor</a>) Hedgehog (Hh).<sup id="cite_ref-Monteiro_25-0" class="reference"><a href="#cite_note-Monteiro-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p><p>Butterfly eyespot morphology appears to be the result of the <a href="/wiki/Evolution" title="Evolution">evolution</a> of an altered version of the <a href="/wiki/Regulatory_circuit" class="mw-redirect" title="Regulatory circuit">regulatory circuit</a> which patterns the wings of other insects. This rogue regulatory circuit is able to pattern both the anterior and posterior eyespots independent of the usual anterior/posterior wing compartmentalization restrictions seen in the fruit fly <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i>.<sup id="cite_ref-Keys_23-1" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-French_24-4" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> The altered regulatory circuit redeploys early developmental signaling sources, like the canonical <a href="/wiki/Hedgehog_(gene)" class="mw-redirect" title="Hedgehog (gene)">hedgehog</a> (Hh) pathway, <a href="/wiki/Distal-less" class="mw-redirect" title="Distal-less">Distal-less</a> (Dll), and <a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">engrailed</a> (En), breaking the anterior/posterior compartmentalization restrictions through increased localized levels of Hh signaling.<sup id="cite_ref-Breakfield_22-1" class="reference"><a href="#cite_note-Breakfield-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Keys_23-2" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-French_24-5" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> In turn, this raises expression of its receptor <a href="/wiki/Patched" title="Patched">Patched</a> (Ptc) and transcription factor.<sup id="cite_ref-French_24-6" class="reference"><a href="#cite_note-French-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> Normally, in <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i>, engrailed acts in the posterior compartment to restrict Ptc and <i><a href="/wiki/Cubitus_interruptus" class="mw-redirect" title="Cubitus interruptus">Cubitus interruptus</a></i> (Ci) expression to the anterior compartment by <a href="/wiki/Repressor_(genetics)" class="mw-redirect" title="Repressor (genetics)">repressing transcription</a> of Ci, thereby preventing Ptc expression.<sup id="cite_ref-Keys_23-3" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> From the perspective of <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">evolutionary developmental biology</a>, understanding the redeployment and <a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">plasticity</a> of existing regulatory mechanisms in butterfly eyespot locus development has given more insight into a fundamental mechanism for the evolution of novel structures.<sup id="cite_ref-Breakfield_22-2" class="reference"><a href="#cite_note-Breakfield-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Keys_23-4" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Distal-less">Distal-less</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=7" title="Edit section: Distal-less"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Distal-less" class="mw-redirect" title="Distal-less">Distal-less</a></div> <p>The Distal-less gene is present in almost all eyespot organizers, making it an ideal candidate to carry out major functions of eyespot formation. During the wing imaginal disc development Dll, has two expression domains separated by a temporal component. First Dll is expressed in a group of cells in the center of what will become the focus and eventually the eyespot. This expression starts during the middle of the fifth <a href="/wiki/Instar" title="Instar">instar</a> <a href="/wiki/Larva" title="Larva">larva</a> and lasts until the <a href="/wiki/Pupa" title="Pupa">pupal</a> stage. The second domain starts around 20 hours after pupation around the original central cluster of cells, in an area in which a black ring of the eyespot will be formed. Functional experiments using <a href="/wiki/Transgenic" class="mw-redirect" title="Transgenic">transgenic</a> <i><a href="/wiki/Bicyclus_anynana" title="Bicyclus anynana">Bicyclus anynana</a></i> (the squinting bush brown butterfly) have shown that overexpression or down-regulation of Dll in the first expression domain correlates with bigger and smaller eyespots respectively. However, if this is done on the second domain then the overall size of the eyespots remains the same, but the width of the black ring raises with a higher amount of Dll. This suggests that Dll might be responsible for the differentiation of the focus in the first expression domain and might be involved in establishing the ring colour patterns in the second domain. These experiments together with the wide distribution of Dll across eyespot forming butterflies suggest that this transcription factor is a central regulator for the correct patterning of the eyespots.<sup id="cite_ref-Monteiro_25-1" class="reference"><a href="#cite_note-Monteiro-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Hedgehog">Hedgehog</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=8" title="Edit section: Hedgehog"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Hedgehog_(gene)" class="mw-redirect" title="Hedgehog (gene)">Hedgehog (gene)</a></div> <p>The Hedgehog (Hh) gene is the other element that has been functionally tested in the formation of eyespots. Investigating genes involved in wing development and morphogenetic activity has led to the discovery that Hh has a primary role in the morphogenetic signaling center of the foci.<sup id="cite_ref-Keys_23-5" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> In a manner that is similar to the development of <i><a href="/wiki/Drosophila" title="Drosophila">Drosophila</a></i> fruit flies, Hh is expressed in all cells in the posterior compartment of the developing butterfly wing during the mid fifth instar of butterfly wing development. However, in butterflies, Hh expression is significantly higher in those cells that flank the potential foci.<sup id="cite_ref-Keys_23-6" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> Higher transcription levels of Hh, along with other known associates of the Hh pathway, namely patched (Ptc) the Hh receptor, and cubitus interruptus (Ci), the Hh transcription factor is seen throughout the mid to late fifth instar as well, which further implies a role for Hh signaling in eyespot development and patterning.<sup id="cite_ref-Keys_23-7" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> </p><p>Furthermore, cells that are flanked by the cells expressing the highest level of Hh signaling are fated to become the foci, indicating that focus cell fate determination relies on high concentrations of Hh in surrounding cells.<sup id="cite_ref-Keys_23-8" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> However, this observation has not been totally confirmed as a rule for multiple butterfly species.<sup id="cite_ref-Monteiro_25-2" class="reference"><a href="#cite_note-Monteiro-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> Studies tried to extrapolate the result of Hh pathway involvement by looking for the expression of Ci in <i>Bicyclus anynana</i>.<sup id="cite_ref-Keys_23-9" class="reference"><a href="#cite_note-Keys-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> Here they observed that both seem to be expressed in eyespots, suggesting a relation with the Hh signaling pathway. However, other studies did not find evidence of Hh expression in <i>B. anynana</i>.<sup id="cite_ref-Monteiro_25-3" class="reference"><a href="#cite_note-Monteiro-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Notch">Notch</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=9" title="Edit section: Notch"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></div> <p>The Notch (N) gene expression precedes an upregulation of Dll in the cells that will become the center of the focus. This makes N the earliest developmental signal, so far studied, that is related with the establishment of the eyespots. Loss of N completely disrupts Dll expression, and eventually eyespot formation, in several butterfly species. A variety of other wing patterns are determined by N and Dll patterns of expression in early development of the wing imaginal disc, suggesting that a single mechanism patterns multiple coloration structures of the wing.<sup id="cite_ref-Reed_26-0" class="reference"><a href="#cite_note-Reed-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=10" title="Edit section: Evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Butterfly eyespots are formed by an interplay of at least 3 genes, namely Distal-less (Dll), spalt (sal), and Antennapedia (Antp), hence their evolution has been shaped by differential expression of these genes in different butterfly taxa, as shown in <i><a href="/wiki/Bicyclus_anynana" title="Bicyclus anynana">Bicyclus anynana</a></i>.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Artificial_eyespots">Artificial eyespots</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=11" title="Edit section: Artificial eyespots"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Eyespot_experiment_on_cattle_in_Botswana.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/84/Eyespot_experiment_on_cattle_in_Botswana.jpg/660px-Eyespot_experiment_on_cattle_in_Botswana.jpg" decoding="async" width="660" height="138" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/8/84/Eyespot_experiment_on_cattle_in_Botswana.jpg 1.5x" data-file-width="962" data-file-height="201" /></a><figcaption>Eyespot experiment on cattle in Botswana. Both the eyespots (left) and the cross markings (centre) protected the cattle from predation by lions, compared to the unmarked controls (right).<sup id="cite_ref-Radford_McNutt_Rogers_Maslen_2020_28-0" class="reference"><a href="#cite_note-Radford_McNutt_Rogers_Maslen_2020-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure><div style="clear:both;" class=""></div> <p>Eyespots painted on the rumps of cows have been shown to reduce cattle predation in Africa. The study authors, Cameron Radford and colleagues, note that in the <a href="/wiki/Sundarbans" title="Sundarbans">Sundarbans</a>, forest users wear face masks with eye markings on the backs of their heads in the hope of reducing tiger attacks. In the study on 2061 cattle in 14 herds over 4 years, 683 were given eye markings, 543 were painted with crosses, and 835 were unpainted. None of the eyed cattle were predated, but 4 cross-marked and 15 unmarked cattle were killed, one by a leopard, the rest by lions. Both the eyespots and the cross markings provided <a href="/wiki/Statistically_significant" class="mw-redirect" title="Statistically significant">statistically significant</a> protection. The cattle were always in mixed groups of marked and unmarked animals; it is not known whether marking all animals in a herd would provide effective protection.<sup id="cite_ref-Radford_McNutt_Rogers_Maslen_2020_28-1" class="reference"><a href="#cite_note-Radford_McNutt_Rogers_Maslen_2020-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Eyespot_(mimicry)&amp;action=edit&amp;section=12" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Eyespot" class="extiw" title="commons:Category:Eyespot">Eyespot</a></span>.</div></div> </div> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Nazar_(amulet)" title="Nazar (amulet)">Nazar (amulet)</a></li> <li><a href="/wiki/Patterns_in_nature" title="Patterns in nature">Patterns in nature</a></li></ul> <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=Eyespot_(mimicry)&amp;action=edit&amp;section=13" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 28em;"> <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"><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="CITEREFKodandaramaiah2011" class="citation journal cs1">Kodandaramaiah, U. 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title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Communications+Biology&amp;rft.atitle=Artificial+eyespots+on+cattle+reduce+predation+by+large+carnivores&amp;rft.volume=3&amp;rft.issue=1&amp;rft.pages=article+430&amp;rft.date=2020-08-07&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7414152%23id-name%3DPMC&amp;rft.issn=2399-3642&amp;rft_id=info%3Apmid%2F32770111&amp;rft_id=info%3Adoi%2F10.1038%2Fs42003-020-01156-0&amp;rft.aulast=Radford&amp;rft.aufirst=Cameron&amp;rft.au=McNutt%2C+John+Weldon&amp;rft.au=Rogers%2C+Tracey&amp;rft.au=Maslen%2C+Ben&amp;rft.au=Jordan%2C+Neil&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC7414152&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AEyespot+%28mimicry%29" class="Z3988"></span></span> </li> </ol></div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist 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href="/wiki/Aggressive_mimicry" title="Aggressive mimicry">Aggressive/Wicklerian</a></li> <li><a href="/wiki/Ant_mimicry" title="Ant mimicry">Ant/Myrmecomorphy</a></li> <li><a href="/wiki/Distraction_display" title="Distraction display">Aristotelian/Distraction display</a></li> <li><a href="/wiki/Automimicry" title="Automimicry">Automimicry</a></li> <li><a href="/wiki/Batesian_mimicry" title="Batesian mimicry">Batesian</a> <ul><li><a href="/wiki/Locomotor_mimicry" title="Locomotor mimicry">Locomotor</a></li></ul></li> <li><a href="/wiki/Brood_parasitism" title="Brood parasitism">Brood</a></li> <li><a href="/wiki/Chemical_mimicry" title="Chemical mimicry">Chemical</a></li> <li><a href="/wiki/Emsleyan_mimicry" title="Emsleyan mimicry">Emsleyan/Mertensian</a></li> <li><a class="mw-selflink selflink">Eyespot</a></li> <li><a href="/wiki/M%C3%BCllerian_mimicry" title="Müllerian mimicry">Müllerian</a></li> <li><a href="/wiki/Sexual_mimicry" title="Sexual mimicry">Sexual</a></li> <li><a href="/wiki/Mimicry_in_vertebrates" title="Mimicry in vertebrates">In vertebrates</a></li> <li><a href="/wiki/Wasmannian_mimicry" class="mw-redirect" title="Wasmannian mimicry">Wasmannian</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="3" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Batesplate_ArM.jpg" class="mw-file-description" title="Batesian mimicry in butterflies"><img alt="Batesian mimicry in butterflies" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/70px-Batesplate_ArM.jpg" decoding="async" width="70" height="60" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/105px-Batesplate_ArM.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/95/Batesplate_ArM.jpg/140px-Batesplate_ArM.jpg 2x" data-file-width="1200" data-file-height="1024" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mimicry_in_plants" title="Mimicry in plants">In plants</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mimicry_in_plants#Bakerian" title="Mimicry in plants">Bakerian</a></li> <li><a href="/wiki/Mimicry_in_plants#Dodsonian" title="Mimicry in plants">Dodsonian</a></li> <li><a href="/wiki/Gilbertian_mimicry" title="Gilbertian mimicry">Gilbertian</a></li> <li><a href="/wiki/Pouyannian_mimicry" title="Pouyannian mimicry">Pouyannian</a></li> <li><a href="/wiki/Vavilovian_mimicry" title="Vavilovian mimicry">Vavilovian</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptation</a></li> <li><a href="/wiki/Animal_communication" title="Animal communication">Animal communication</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a>/<a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Co-evolution</a></li> <li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Coloration_evidence_for_natural_selection" title="Coloration evidence for natural selection">Coloration evidence for natural selection</a></li> <li><i><a href="/wiki/Dazzled_and_Deceived" title="Dazzled and Deceived">Dazzled and Deceived</a></i></li> <li><a href="/wiki/Deception_in_animals" title="Deception in animals">Deception in animals</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Mimicry#Evolution" title="Mimicry">Mimicry#Evolution</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Frequency-dependent_selection" title="Frequency-dependent selection">Frequency-dependent selection</a></li> <li><a href="/wiki/Phagomimicry" title="Phagomimicry">Phagomimicry</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li>&#160;<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:Mimicry" title="Category:Mimicry">Category mimicry</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="Vision_in_animals" 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="3"><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:Vision_in_animals" title="Template:Vision in animals"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Vision_in_animals" title="Template talk:Vision in animals"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Vision_in_animals" title="Special:EditPage/Template:Vision in animals"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Vision_in_animals" style="font-size:114%;margin:0 4em">Vision in animals</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Visual_system" title="Visual system">Vision</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/Bird_vision" title="Bird vision">Birds</a></li> <li><a href="/wiki/Chameleon_vision" title="Chameleon vision">Chameleons</a></li> <li><a href="/wiki/Dinosaur_vision" title="Dinosaur vision">Dinosaurs</a></li> <li><a href="/wiki/Vision_in_fish" title="Vision in fish">Fish</a></li> <li><a href="/wiki/Vision_in_toads" title="Vision in toads">Toads</a></li> <li><a href="/wiki/Mammalian_vision" title="Mammalian vision">Mammals</a> <ul><li><a href="/wiki/Equine_vision" title="Equine vision">horses</a></li> <li><a href="/wiki/Dog_anatomy#Vision" title="Dog anatomy">dogs</a></li> <li><a href="/wiki/Cat_senses#Sight" title="Cat senses">cats</a></li></ul></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:Astronotus_ocellatus_-_closeup_(aka).jpg" class="mw-file-description" title="Fish surveying its world"><img alt="Fish surveying its world" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/140px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg" decoding="async" width="140" height="93" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/210px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Astronotus_ocellatus_-_closeup_%28aka%29.jpg/280px-Astronotus_ocellatus_-_closeup_%28aka%29.jpg 2x" data-file-width="3008" data-file-height="2000" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eye" title="Eye">Eyes</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/Arthropod_eye" title="Arthropod eye">Arthropod eye</a></li> <li><a href="/wiki/Compound_eye" title="Compound eye">Compound eye</a></li> <li><a href="/wiki/Eagle_eye" title="Eagle eye">Eagle eye</a></li> <li><a href="/wiki/Tapetum_lucidum" title="Tapetum lucidum">Eye shine</a></li> <li><a href="/wiki/Simple_eye_in_invertebrates" title="Simple eye in invertebrates">Simple eye in invertebrates</a></li> <li><a href="/wiki/Mammalian_eye" title="Mammalian eye">Mammalian eye</a> <ul><li><a href="/wiki/Human_eye" title="Human eye">human</a></li></ul></li> <li><a href="/wiki/Mollusc_eye" title="Mollusc eye">Mollusc eye</a> <ul><li><a href="/wiki/Cephalopod_eye" title="Cephalopod eye">cephalopod</a></li> <li><a href="/wiki/Gastropod_eye" class="mw-redirect" title="Gastropod eye">gastropod</a></li></ul></li> <li><a href="/wiki/Holochroal_eye" title="Holochroal eye">Holochroal eye</a></li> <li><a href="/wiki/Parietal_eye" title="Parietal eye">Parietal eye</a></li> <li><a href="/wiki/Schizochroal_eye" class="mw-redirect" title="Schizochroal eye">Schizochroal eye</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</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/Evolution_of_the_eye" title="Evolution of the eye">Evolution of the eye</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Evolution of color vision</a></li> <li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Evolution of color vision in primates</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Coloration</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/Albinism" title="Albinism">Albinism</a></li> <li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Chromatophore" title="Chromatophore">Chromatophore</a></li> <li><a href="/wiki/Counter-illumination" title="Counter-illumination">Counter-illumination</a></li> <li><a href="/wiki/Countershading" title="Countershading">Countershading</a></li> <li><a href="/wiki/Crypsis" title="Crypsis">Crypsis</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Disruptive_coloration" title="Disruptive coloration">Disruptive coloration</a> <ul><li><a href="/wiki/Coincident_disruptive_coloration" title="Coincident disruptive coloration">coincident</a></li></ul></li> <li><a class="mw-selflink selflink">Eyespot (mimicry)</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Structural_coloration" title="Structural coloration">Structural coloration</a></li> <li><a href="/wiki/Underwater_camouflage" title="Underwater camouflage">Underwater camouflage</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</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/Sense" title="Sense">Animal senses</a></li> <li><a href="/wiki/Blindness_in_animals" title="Blindness in animals">Blindness in animals</a></li> <li><a href="/wiki/Eyespot_apparatus" title="Eyespot apparatus">Eyespot apparatus</a></li> <li><a href="/wiki/Feature_detection_(nervous_system)" title="Feature detection (nervous system)">Feature detection</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/Monocular_deprivation" title="Monocular deprivation">Monocular deprivation</a></li> <li><a href="/wiki/Ommatidium" title="Ommatidium">Ommatidium</a></li> <li><a href="/wiki/Palpebral_(bone)" title="Palpebral (bone)">Palpebral (bone)</a></li> <li><a href="/wiki/Pseudopupil" title="Pseudopupil">Pseudopupil</a></li> <li><a href="/wiki/Rhopalium" title="Rhopalium">Rhopalium</a></li> <li><a href="/wiki/Underwater_vision" title="Underwater vision">Underwater vision</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</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="The_development_of_phenotype" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Genarch" title="Template:Genarch"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genarch" title="Template talk:Genarch"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genarch" title="Special:EditPage/Template:Genarch"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="The_development_of_phenotype" style="font-size:114%;margin:0 4em">The <a href="/wiki/Morphogenesis" title="Morphogenesis">development</a> of <a href="/wiki/Phenotype" title="Phenotype">phenotype</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key concepts</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">Genotype–phenotype distinction</a></li> <li><a href="/wiki/Reaction_norm" title="Reaction norm">Reaction norm</a></li> <li><a href="/wiki/Gene%E2%80%93environment_interaction" title="Gene–environment interaction">Gene–environment interaction</a></li> <li><a href="/wiki/Gene%E2%80%93environment_correlation" title="Gene–environment correlation">Gene–environment correlation</a></li> <li><a href="/wiki/Operon" title="Operon">Operon</a></li> <li><a href="/wiki/Heritability" title="Heritability">Heritability</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative genetics</a></li> <li><a href="/wiki/Heterochrony" title="Heterochrony">Heterochrony</a> <ul><li><a href="/wiki/Neoteny" title="Neoteny">Neoteny</a></li></ul></li> <li><a href="/wiki/Heterotopy" title="Heterotopy">Heterotopy</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_architecture" title="Genetic architecture">Genetic architecture</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Dominance_(genetics)" title="Dominance (genetics)">Dominance</a></li> <li><a href="/wiki/Epistasis" title="Epistasis">Epistasis</a></li> <li><a href="/wiki/Fitness_landscape" title="Fitness landscape">Fitness landscape</a>/<a href="/wiki/Evolutionary_landscape" title="Evolutionary landscape">evolutionary landscape</a></li> <li><a href="/wiki/Pleiotropy" title="Pleiotropy">Pleiotropy</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Plasticity</a></li> <li><a href="/wiki/Polygenic_inheritance" class="mw-redirect" title="Polygenic inheritance">Polygenic inheritance</a></li> <li><a href="/wiki/Transgressive_segregation" title="Transgressive segregation">Transgressive segregation</a></li> <li><a href="/wiki/Sequence_space_(evolution)" title="Sequence space (evolution)">Sequence space</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-genetic influences</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Maternal_effect" title="Maternal effect">Maternal effect</a></li> <li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">Genomic imprinting</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Polyphenism" title="Polyphenism">Polyphenism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Developmental architecture</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Developmental_biology" title="Developmental biology">Developmental biology</a></li> <li><a href="/wiki/Morphogenesis" title="Morphogenesis">Morphogenesis</a> <ul><li><a class="mw-selflink selflink">Eyespot</a></li> <li><a href="/wiki/Pattern_formation" title="Pattern formation">Pattern formation</a></li></ul></li> <li><a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">Segmentation</a></li> <li><a href="/wiki/Metamerism_(biology)" title="Metamerism (biology)">Metamerism</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution of genetic systems</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolvability" title="Evolvability">Evolvability</a></li> <li><a href="/wiki/Robustness_(evolution)" title="Robustness (evolution)">Robustness</a></li> <li><a href="/wiki/Neutral_network_(evolution)" title="Neutral network (evolution)">Neutral networks</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li> <li><a href="/wiki/Evolutionary_tinkering" title="Evolutionary tinkering">Tinkering</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Control of development</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">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/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation of gene expression</a></li> <li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Evo-devo_gene_toolkit" title="Evo-devo gene toolkit">Evo-devo gene toolkit</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Homeobox" title="Homeobox">Homeobox</a></li> <li><a href="/wiki/Hedgehog_signaling_pathway" title="Hedgehog signaling pathway">Hedgehog signaling pathway</a></li> <li><a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">Notch signaling pathway</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Elements</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/Homeotic_gene" title="Homeotic gene">Homeotic gene</a></li> <li><a href="/wiki/Hox_gene" title="Hox gene">Hox gene</a></li> <li><a href="/wiki/Pax_genes" title="Pax genes">Pax genes</a> <ul><li><a href="/wiki/PAX6" title="PAX6">eyeless gene</a></li></ul></li> <li><a href="/wiki/DLX_gene_family" title="DLX gene family">Distal-less</a></li> <li><a href="/wiki/Engrailed_(gene)" title="Engrailed (gene)">Engrailed</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li> <li><a href="/wiki/Ligand_(biochemistry)" title="Ligand (biochemistry)">Ligand</a></li> <li><a href="/wiki/Morphogen" title="Morphogen">Morphogen</a></li> <li><a href="/wiki/Receptor_(biochemistry)" title="Receptor (biochemistry)">Cell surface receptor</a></li> <li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential figures</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/C._H._Waddington" title="C. H. Waddington">C. H. Waddington</a></li> <li><a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a></li> <li><a href="/wiki/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a> + <a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a> <ul><li><a href="/wiki/Lac_operon" title="Lac operon">Lac operon</a></li></ul></li> <li><a href="/wiki/Eric_F._Wieschaus" title="Eric F. Wieschaus">Eric F. Wieschaus</a></li> <li><a href="/wiki/Christiane_N%C3%BCsslein-Volhard" title="Christiane Nüsslein-Volhard">Christiane Nüsslein-Volhard</a></li> <li><a href="/wiki/William_McGinnis" title="William McGinnis">William McGinnis</a></li> <li><a href="/wiki/Michael_Levine_(biologist)" title="Michael Levine (biologist)">Mike Levine</a></li> <li><a href="/wiki/Sean_B._Carroll" title="Sean B. Carroll">Sean B. Carroll</a> <ul><li><i><a href="/wiki/Endless_Forms_Most_Beautiful_(book)" title="Endless Forms Most Beautiful (book)">Endless Forms Most Beautiful</a></i></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Debates</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Morphogenetic_field" title="Morphogenetic field">Morphogenetic field</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index of evolutionary biology articles</a></div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐rqr4k Cached time: 20241123161544 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 0.599 seconds Real time usage: 0.772 seconds Preprocessor visited node count: 2659/1000000 Post‐expand include size: 118260/2097152 bytes Template argument size: 1214/2097152 bytes Highest expansion depth: 13/100 Expensive parser function count: 6/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 155197/5000000 bytes Lua time usage: 0.354/10.000 seconds Lua memory usage: 6114716/52428800 bytes Number of Wikibase entities loaded: 0/400 --> <!-- Transclusion expansion time report (%,ms,calls,template) 100.00% 658.609 1 -total 41.86% 275.684 1 Template:Reflist 30.95% 203.853 23 Template:Cite_journal 15.61% 102.836 4 Template:Navbox 14.86% 97.893 1 Template:Mimicry 11.43% 75.256 4 Template:Main_other 11.13% 73.327 1 Template:Short_description 8.61% 56.688 1 Template:Commons_category 8.13% 53.542 1 Template:Sister_project 7.71% 50.767 1 Template:Side_box --> <!-- Saved in parser cache with key enwiki:pcache:idhash:11615271-0!canonical and timestamp 20241123161544 and revision id 1253646521. 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