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class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Rate of evolution</span> </div> </a> <ul id="toc-Rate_of_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Origins_of_the_eye" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Origins_of_the_eye"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Origins of the eye</span> </div> </a> <ul id="toc-Origins_of_the_eye-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stages_of_evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Stages_of_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Stages of evolution</span> </div> </a> <button aria-controls="toc-Stages_of_evolution-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 Stages of evolution subsection</span> </button> <ul id="toc-Stages_of_evolution-sublist" class="vector-toc-list"> <li id="toc-Early_eyes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Early_eyes"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Early eyes</span> </div> </a> <ul id="toc-Early_eyes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Lens_formation_and_diversification" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Lens_formation_and_diversification"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Lens formation and diversification</span> </div> </a> <ul id="toc-Lens_formation_and_diversification-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Crystallins" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Crystallins"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Crystallins</span> </div> </a> <ul id="toc-Crystallins-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Aqueous_humor,_iris,_and_cornea" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Aqueous_humor,_iris,_and_cornea"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Aqueous humor, iris, and cornea</span> </div> </a> <ul id="toc-Aqueous_humor,_iris,_and_cornea-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_developments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Other_developments"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Other developments</span> </div> </a> <ul id="toc-Other_developments-sublist" class="vector-toc-list"> <li id="toc-Color_vision" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Color_vision"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5.1</span> <span>Color vision</span> </div> </a> <ul id="toc-Color_vision-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Polarization_vision" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Polarization_vision"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5.2</span> <span>Polarization vision</span> </div> </a> <ul id="toc-Polarization_vision-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Focusing_mechanism" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Focusing_mechanism"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5.3</span> <span>Focusing mechanism</span> </div> </a> <ul id="toc-Focusing_mechanism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Placement" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Placement"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5.4</span> <span>Placement</span> </div> </a> <ul id="toc-Placement-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </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-Explanatory_footnotes" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Explanatory_footnotes"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Explanatory footnotes</span> </div> </a> <ul id="toc-Explanatory_footnotes-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">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</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 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</div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Evolution of the eye</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 15 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-15" 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">15 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%AA%D8%B7%D9%88%D8%B1_%D8%A7%D9%84%D8%B9%D9%8A%D9%86" title="تطور العين – Arabic" lang="ar" hreflang="ar" data-title="تطور العين" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%9A%E0%A7%8B%E0%A6%96%E0%A7%87%E0%A6%B0_%E0%A6%AC%E0%A6%BF%E0%A6%AC%E0%A6%B0%E0%A7%8D%E0%A6%A4%E0%A6%A8" title="চোখের বিবর্তন – Bangla" lang="bn" hreflang="bn" data-title="চোখের বিবর্তন" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Augenevolution" title="Augenevolution – German" lang="de" hreflang="de" data-title="Augenevolution" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%95%CE%BE%CE%AD%CE%BB%CE%B9%CE%BE%CE%B7_%CF%84%CE%BF%CF%85_%CE%BC%CE%B1%CF%84%CE%B9%CE%BF%CF%8D" title="Εξέλιξη του ματιού – Greek" lang="el" hreflang="el" data-title="Εξέλιξη του ματιού" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Evoluci%C3%B3n_del_ojo" title="Evolución del ojo – Spanish" lang="es" hreflang="es" data-title="Evolución del ojo" 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/%D9%81%D8%B1%DA%AF%D8%B4%D8%AA_%DA%86%D8%B4%D9%85" title="فرگشت چشم – Persian" lang="fa" hreflang="fa" data-title="فرگشت چشم" 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/Histoire_%C3%A9volutive_de_l%27%C5%93il" title="Histoire évolutive de l'œil – French" lang="fr" hreflang="fr" data-title="Histoire évolutive de l'œil" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%90%D7%91%D7%95%D7%9C%D7%95%D7%A6%D7%99%D7%94_%D7%A9%D7%9C_%D7%94%D7%A2%D7%99%D7%9F" title="אבולוציה של העין – Hebrew" lang="he" hreflang="he" data-title="אבולוציה של העין" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D8%AA%D8%B7%D9%88%D8%B1_%D8%A7%D9%84%D8%B9%D9%8A%D9%86" title="تطور العين – Egyptian Arabic" lang="arz" hreflang="arz" data-title="تطور العين" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target"><span>مصرى</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%9C%BC%E3%81%AE%E9%80%B2%E5%8C%96" title="眼の進化 – Japanese" lang="ja" hreflang="ja" data-title="眼の進化" 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-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Evolution_of_the_eye" title="Evolution of the eye – Simple English" lang="en-simple" hreflang="en-simple" data-title="Evolution of the eye" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%BE%DB%95%D8%B1%DB%95%D8%B3%DB%95%D9%86%D8%AF%D9%86%DB%8C_%DA%86%D8%A7%D9%88" title="پەرەسەندنی چاو – Central Kurdish" lang="ckb" hreflang="ckb" data-title="پەرەسەندنی چاو" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target"><span>کوردی</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Silm%C3%A4n_evoluutio" title="Silmän evoluutio – Finnish" lang="fi" hreflang="fi" data-title="Silmän evoluutio" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%A7%E0%B8%B4%E0%B8%A7%E0%B8%B1%E0%B8%92%E0%B8%99%E0%B8%B2%E0%B8%81%E0%B8%B2%E0%B8%A3%E0%B8%82%E0%B8%AD%E0%B8%87%E0%B8%95%E0%B8%B2" title="วิวัฒนาการของตา – Thai" lang="th" hreflang="th" data-title="วิวัฒนาการของตา" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target"><span>ไทย</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/G%C3%B6z%C3%BCn_evrimi" title="Gözün evrimi – Turkish" lang="tr" hreflang="tr" data-title="Gözün evrimi" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</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/Q2732276#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-associated-pages" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-nstab-main" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Evolution_of_the_eye" title="View the content page [c]" 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<div class="mw-indicators"> </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"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Origins and diversification of the organs of sight through geologic time</div> <p class="mw-empty-elt"> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Diagram_of_eye_evolution.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Diagram_of_eye_evolution.svg/350px-Diagram_of_eye_evolution.svg.png" decoding="async" width="350" height="488" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Diagram_of_eye_evolution.svg/525px-Diagram_of_eye_evolution.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Diagram_of_eye_evolution.svg/700px-Diagram_of_eye_evolution.svg.png 2x" data-file-width="584" data-file-height="815" /></a><figcaption>Major stages in the evolution of the eye in <a href="/wiki/Vertebrates" class="mw-redirect" title="Vertebrates">vertebrates</a></figcaption></figure> <p>Many scientists have found the <b>evolution of the eye</b> attractive to study because the <a href="/wiki/Eye" title="Eye">eye</a> distinctively exemplifies an <a href="/wiki/Convergent_evolution" title="Convergent evolution">analogous</a> <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organ</a> found in many <a href="/wiki/Taxon" title="Taxon">animal forms</a>. Simple light detection is found in bacteria, single-celled organisms, plants and animals. Complex, image-forming eyes have evolved independently several times.<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> </p><p>Diverse eyes are known from the <a href="/wiki/Burgess_shale" class="mw-redirect" title="Burgess shale">Burgess shale</a> of the <a href="/wiki/Middle_Cambrian" class="mw-redirect" title="Middle Cambrian">Middle Cambrian</a>, and from the slightly older <a href="/wiki/Emu_Bay_Shale" title="Emu Bay Shale">Emu Bay Shale</a>.<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> Eyes vary in their <a href="/wiki/Visual_acuity" title="Visual acuity">visual acuity</a>, the range of wavelengths they can detect, their sensitivity in no light, their ability to detect motion or to resolve objects, and whether they can <a href="/wiki/Colour_vision" class="mw-redirect" title="Colour vision">discriminate colours</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History_of_research">History of research</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=1" title="Edit section: History of research"><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:Eye_iris.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Eye_iris.jpg/220px-Eye_iris.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Eye_iris.jpg/330px-Eye_iris.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/65/Eye_iris.jpg/440px-Eye_iris.jpg 2x" data-file-width="2788" data-file-height="1864" /></a><figcaption>The human eye, showing the <a href="/wiki/Iris_(anatomy)" title="Iris (anatomy)">iris</a> and <a href="/wiki/Pupil" title="Pupil">pupil</a></figcaption></figure> <p>In 1802, philosopher <a href="/wiki/William_Paley" title="William Paley">William Paley</a> called it a miracle of "<a href="/wiki/Design" title="Design">design</a>." In 1859, <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> himself wrote in his <i><a href="/wiki/Origin_of_Species" class="mw-redirect" title="Origin of Species">Origin of Species</a></i>, that the evolution of the eye by natural selection seemed at first glance "absurd in the highest possible degree".<sup id="cite_ref-origin_3-0" class="reference"><a href="#cite_note-origin-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> However, he went on that despite the difficulty in imagining it, its evolution was perfectly feasible: </p> <blockquote><p>... if numerous gradations from a simple and imperfect eye to one complex and perfect can be shown to exist, each grade being useful to its possessor, as is certainly the case; if further, the eye ever varies and the variations be inherited, as is likewise certainly the case and if such variations should be useful to any animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, should not be considered as subversive of the theory.<sup id="cite_ref-origin_3-1" class="reference"><a href="#cite_note-origin-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></p></blockquote> <p>He suggested a stepwise evolution from "an optic nerve merely coated with pigment, and without any other mechanism" to "a moderately high stage of perfection", and gave examples of existing intermediate.<sup id="cite_ref-origin_3-2" class="reference"><a href="#cite_note-origin-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Current research is investigating the genetic mechanisms underlying eye development and evolution.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>Biologist <a href="/w/index.php?title=D.E._Nilsson&action=edit&redlink=1" class="new" title="D.E. Nilsson (page does not exist)">D.E. Nilsson</a> has independently theorized about four general stages in the evolution of a vertebrate eye from a patch of photoreceptors.<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> Nilsson and S. Pelger estimated in a classic paper that only a few hundred thousand generations are needed to evolve a complex eye in vertebrates.<sup id="cite_ref-Nilsson,_D_1994_6-0" class="reference"><a href="#cite_note-Nilsson,_D_1994-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Another researcher, G.C. Young, has used the fossil record to infer evolutionary conclusions, based on the structure of eye orbits and openings in fossilized skulls for blood vessels and nerves to go through.<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> All this adds to the growing amount of evidence that supports Darwin's theory. </p> <div class="mw-heading mw-heading2"><h2 id="Rate_of_evolution">Rate of evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=2" title="Edit section: Rate of evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first possible fossils of eyes found to date are from the <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> period (about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=555">555</a> million years ago</span>),<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> while the oldest certain fossilized eye is from a <i><a href="/wiki/Schmidtiellus" title="Schmidtiellus">Schmidtiellus reetae</a></i> fossil from 530 mya, collected in <a href="/wiki/Saviranna" title="Saviranna">Saviranna</a> in northern Estonia. The structure is similar to the <a href="/wiki/Compound_eye" title="Compound eye">compound eyes</a> of modern-day dragonflies and bees, but with (~100) <a href="/wiki/Ommatidium" title="Ommatidium">ommatidia</a> spaced further apart, and without a <a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">lens</a>.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The lower Cambrian had a burst of apparently rapid evolution, called the "<a href="/wiki/Cambrian_explosion" title="Cambrian explosion">Cambrian explosion</a>". One of the many hypotheses for "causes" of the Cambrian explosion is the "Light Switch" theory of <a href="/wiki/Andrew_Parker_(zoologist)" title="Andrew Parker (zoologist)">Andrew Parker</a>: it holds that the evolution of advanced eyes started an <a href="/wiki/Red_Queen_hypothesis" title="Red Queen hypothesis">arms race</a> that accelerated evolution.<sup id="cite_ref-parker2003_11-0" class="reference"><a href="#cite_note-parker2003-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Before the Cambrian explosion, animals may have sensed light, but did not use it for fast locomotion or navigation by vision. </p><p>The rate of eye evolution is difficult to estimate because the fossil record, particularly of the lower Cambrian, is poor. How fast a circular patch of photoreceptor cells can evolve into a fully functional vertebrate eye has been estimated based on rates of mutation, relative advantage to the organism, and natural selection. However, the time needed for each state was consistently overestimated and the <a href="/wiki/Generation_time" title="Generation time">generation time</a> was set to one year, which is common in small animals. Even with these pessimistic values, the vertebrate eye could still evolve from a patch of photoreceptor cells in less than 364,000 years.<sup id="cite_ref-Nilsson1994_12-0" class="reference"><a href="#cite_note-Nilsson1994-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>note 1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Origins_of_the_eye">Origins of the eye</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=3" title="Edit section: Origins of the eye"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Whether the eye evolved once or many times depends on the definition of an eye. All eyed animals share much of the genetic machinery for eye development. This suggests that the ancestor of eyed animals had some form of light-sensitive machinery – even if it was not a dedicated optical organ. However, even photoreceptor cells may have evolved more than once from molecularly similar chemoreceptor cells. Probably, photoreceptor cells existed long before the Cambrian explosion.<sup id="cite_ref-Nilsson1996_14-0" class="reference"><a href="#cite_note-Nilsson1996-14"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Higher-level similarities – such as the use of the protein <a href="/wiki/Crystallin" title="Crystallin">crystallin</a> in the independently derived cephalopod and vertebrate lenses<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> – reflect the <a href="/wiki/Exaptation" title="Exaptation">co-option</a> of a more fundamental protein to a new function within the eye.<sup id="cite_ref-Scotland2010_16-0" class="reference"><a href="#cite_note-Scotland2010-16"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>A shared trait common to all light-sensitive organs are <a href="/wiki/Opsin" title="Opsin">opsins</a>. Opsins belong to a family of photo-sensitive proteins and fall into nine groups, which already existed in the <a href="/wiki/Urbilaterian" title="Urbilaterian">urbilaterian</a>, the last common ancestor of all <a href="/wiki/Bilateria" title="Bilateria">bilaterally symmetrical animals</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Additionally, the genetic toolkit for positioning eyes is shared by all animals: The <a href="/wiki/PAX6" title="PAX6">PAX6 gene</a> controls where eyes develop in animals ranging from octopuses<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> to mice and <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">fruit flies</a>.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Such high-level genes are, by implication, much older than many of the structures that they control today; they must originally have served a different purpose, before they were co-opted for eye development.<sup id="cite_ref-Scotland2010_16-1" class="reference"><a href="#cite_note-Scotland2010-16"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Eyes and other sensory organs probably evolved before the brain: There is no need for an information-processing organ (brain) before there is information to process.<sup id="cite_ref-Gehring_22-0" class="reference"><a href="#cite_note-Gehring-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> A living example are <a href="/wiki/Cubozoan" class="mw-redirect" title="Cubozoan">cubozoan</a> jellyfish that possess eyes comparable to <a href="/wiki/Vertebrate" title="Vertebrate">vertebrate</a> and <a href="/wiki/Cephalopod" title="Cephalopod">cephalopod</a> <a href="/wiki/Cephalopod_eye" title="Cephalopod eye">camera eyes</a> despite lacking a brain.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Stages_of_evolution">Stages of evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=4" title="Edit section: Stages of evolution"><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:Augentierchen_440x600.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Augentierchen_440x600.jpg/170px-Augentierchen_440x600.jpg" decoding="async" width="170" height="231" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/17/Augentierchen_440x600.jpg/255px-Augentierchen_440x600.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/17/Augentierchen_440x600.jpg/340px-Augentierchen_440x600.jpg 2x" data-file-width="440" data-file-height="599" /></a><figcaption>The <i>stigma</i> (2) of the <a href="/wiki/Euglena" title="Euglena">euglena</a> hides a light-sensitive spot.</figcaption></figure> <p>The earliest predecessors of the eye were photoreceptor proteins that sense light, found even in unicellular organisms, called "<a href="/wiki/Eyespot_apparatus" title="Eyespot apparatus">eyespots</a>".<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Eyespots can sense only ambient brightness: they can distinguish light from dark, sufficient for <a href="/wiki/Photoperiodism" title="Photoperiodism">photoperiodism</a> and daily synchronization of <a href="/wiki/Circadian_rhythm" title="Circadian rhythm">circadian rhythms</a>. They are insufficient for vision, as they cannot distinguish shapes or determine the direction light is coming from. Eyespots are found in nearly all major animal groups, and are common among unicellular organisms, including <a href="/wiki/Euglena" title="Euglena">euglena</a>. The euglena's eyespot, called a <i>stigma</i>, is located at its anterior end. It is a small splotch of red pigment which shades a collection of light sensitive crystals. Together with the leading <a href="/wiki/Flagellum" title="Flagellum">flagellum</a>, the eyespot allows the organism to move in response to light, often toward the light to assist in <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>,<sup id="cite_ref-Land1992_25-0" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> and to predict day and night, the primary function of circadian rhythms. Visual pigments are located in the brains of more complex organisms, and are thought to have a role in synchronising spawning with lunar cycles. By detecting the subtle changes in night-time illumination, organisms could synchronise the release of sperm and eggs to maximise the probability of fertilisation.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p><p>Vision itself relies on a basic biochemistry which is common to all eyes. However, how this biochemical toolkit is used to interpret an organism's environment varies widely: eyes have a wide range of structures and forms, all of which have evolved quite late relative to the underlying proteins and molecules.<sup id="cite_ref-Land1992_25-1" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>At a cellular level, there appear to be two main types of eyes, one possessed by the <a href="/wiki/Protostomes" class="mw-redirect" title="Protostomes">protostomes</a> (<a href="/wiki/Mollusc" class="mw-redirect" title="Mollusc">molluscs</a>, <a href="/wiki/Annelids" class="mw-redirect" title="Annelids">annelid worms</a> and <a href="/wiki/Arthropod" title="Arthropod">arthropods</a>), the other by the <a href="/wiki/Deuterostomes" class="mw-redirect" title="Deuterostomes">deuterostomes</a> (<a href="/wiki/Chordate" title="Chordate">chordates</a> and <a href="/wiki/Echinoderm" title="Echinoderm">echinoderms</a>).<sup id="cite_ref-Land1992_25-2" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>The functional unit of the eye is the photoreceptor cell, which contains the opsin proteins and responds to light by initiating a nerve impulse. The light sensitive opsins are borne on a hairy layer, to maximise the surface area. The nature of these "hairs" differs, with two basic forms underlying photoreceptor structure: <a href="/wiki/Microvillus" title="Microvillus">microvilli</a> and <a href="/wiki/Cilium" title="Cilium">cilia</a>.<sup id="cite_ref-Autrum1979_27-0" class="reference"><a href="#cite_note-Autrum1979-27"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> In the eyes of protostomes, they are microvilli: extensions or protrusions of the cellular membrane. But in the eyes of deuterostomes, they are derived from cilia, which are separate structures.<sup id="cite_ref-Land1992_25-3" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> However, outside the eyes an organism may use the other type of photoreceptor cells, for instance the clamworm <i><a href="/wiki/Platynereis_dumerilii" title="Platynereis dumerilii">Platynereis dumerilii</a></i> uses microvilliar cells in the eyes but has additionally deep brain ciliary photoreceptor cells.<sup id="cite_ref-Arendt2004_28-0" class="reference"><a href="#cite_note-Arendt2004-28"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> The actual derivation may be more complicated, as some microvilli contain traces of cilia – but other observations appear to support a fundamental difference between protostomes and deuterostomes.<sup id="cite_ref-Land1992_25-4" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> These considerations centre on the response of the cells to light – some use sodium to cause the electric signal that will form a nerve impulse, and others use potassium; further, protostomes on the whole construct a signal by allowing <i>more</i> sodium to pass through their cell walls, whereas deuterostomes allow less through.<sup id="cite_ref-Land1992_25-5" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>This suggests that when the two lineages diverged in the Precambrian, they had only very primitive light receptors, which developed into more complex eyes independently. </p> <div class="mw-heading mw-heading3"><h3 id="Early_eyes">Early eyes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=5" title="Edit section: Early eyes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The basic light-processing unit of eyes is the <a href="/wiki/Photoreceptor_cell" title="Photoreceptor cell">photoreceptor cell</a>, a specialized cell containing two types of molecules bound to each other and located in a membrane: the <a href="/wiki/Opsin" title="Opsin">opsin</a>, a light-sensitive <a href="/wiki/Protein" title="Protein">protein</a>; and a <a href="/wiki/Chromophore" title="Chromophore">chromophore</a>, the <a href="/wiki/Pigment" title="Pigment">pigment</a> that absorbs light. Groups of such cells are termed "eyespots", and have evolved independently somewhere between 40 and 65 times. These eyespots permit animals to gain only a basic sense of the direction and intensity of light, but not enough to discriminate an object from its surroundings.<sup id="cite_ref-Land1992_25-6" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>Developing an optical system that can discriminate the direction of light to within a few degrees is apparently much more difficult, and only six of the thirty-some phyla<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>note 2<span class="cite-bracket">]</span></a></sup> possess such a system. However, these phyla account for 96% of living species.<sup id="cite_ref-Land1992_25-7" class="reference"><a href="#cite_note-Land1992-25"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Smed.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Smed.jpg/220px-Smed.jpg" decoding="async" width="220" height="91" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Smed.jpg/330px-Smed.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Smed.jpg/440px-Smed.jpg 2x" data-file-width="2116" data-file-height="880" /></a><figcaption>The <a href="/wiki/Planarian" title="Planarian">planarian</a> has "cup" eyespots that can slightly distinguish light direction.</figcaption></figure> <p>These complex optical systems started out as the multicellular eyepatch gradually depressed into a cup, which first granted the ability to discriminate brightness in directions, then in finer and finer directions as the pit deepened. While flat eyepatches were ineffective at determining the direction of light, as a beam of light would activate exactly the same patch of photo-sensitive cells regardless of its direction, the "cup" shape of the pit eyes allowed limited directional differentiation by changing which cells the lights would hit depending upon the light's angle. Pit eyes, which had arisen by the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> period, were seen in ancient <a href="/wiki/Snail" title="Snail">snails</a>,<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (December 2008)">clarification needed</span></a></i>]</sup> and are found in some snails and other invertebrates living today, such as <a href="/wiki/Planaria" title="Planaria">planaria</a>. Planaria can slightly differentiate the direction and intensity of light because of their cup-shaped, heavily pigmented <a href="/wiki/Retina" title="Retina">retina</a> cells, which shield the light-sensitive cells from exposure in all directions except for the single opening for the light. However, this proto-eye is still much more useful for detecting the absence or presence of light than its direction; this gradually changes as the eye's pit deepens and the number of photoreceptive cells grows, allowing for increasingly precise visual information.<sup id="cite_ref-ee_30-0" class="reference"><a href="#cite_note-ee-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>When a <a href="/wiki/Photon" title="Photon">photon</a> is absorbed by the chromophore, a chemical reaction causes the photon's energy to be transduced into electrical energy and relayed, in higher animals, to the <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a>. These photoreceptor cells form part of the <a href="/wiki/Retina" title="Retina">retina</a>, a thin layer of cells that relays <a href="/wiki/Visual_perception" title="Visual perception">visual</a> information,<sup id="cite_ref-photons_31-0" class="reference"><a href="#cite_note-photons-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> including the light and day-length information needed by the circadian rhythm system, to the brain. However, some <a href="/wiki/Jellyfish" title="Jellyfish">jellyfish</a>, such as <i><a href="/wiki/Cladonema_(hydrozoan)" title="Cladonema (hydrozoan)">Cladonema</a></i> (<a href="/wiki/Cladonematidae" title="Cladonematidae">Cladonematidae</a>), have elaborate eyes but no brain. Their eyes transmit a message directly to the muscles without the intermediate processing provided by a brain.<sup id="cite_ref-Gehring_22-1" class="reference"><a href="#cite_note-Gehring-22"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>During the <a href="/wiki/Cambrian_explosion" title="Cambrian explosion">Cambrian explosion</a>, the development of the eye accelerated rapidly, with radical improvements in image-processing and detection of light direction.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Nautilus_pompilius_(head).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Nautilus_pompilius_%28head%29.jpg/220px-Nautilus_pompilius_%28head%29.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Nautilus_pompilius_%28head%29.jpg/330px-Nautilus_pompilius_%28head%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9c/Nautilus_pompilius_%28head%29.jpg/440px-Nautilus_pompilius_%28head%29.jpg 2x" data-file-width="4272" data-file-height="2848" /></a><figcaption>The <a href="/wiki/Nautilus" title="Nautilus">nautilus</a> eye functions similarly to a <a href="/wiki/Pinhole_camera" title="Pinhole camera">pinhole camera</a>.</figcaption></figure> <p>After the photosensitive cell region <a href="/wiki/Invagination" title="Invagination">invaginated</a>, there came a point when reducing the width of the light opening became more efficient at increasing visual resolution than continued deepening of the cup.<sup id="cite_ref-Nilsson1994_12-1" class="reference"><a href="#cite_note-Nilsson1994-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> By reducing the size of the opening, organisms achieved true imaging, allowing for fine directional sensing and even some shape-sensing. Eyes of this nature are currently found in the <a href="/wiki/Nautilus" title="Nautilus">nautilus</a>. Lacking a cornea or lens, they provide poor resolution and dim imaging, but are still, for the purpose of vision, a major improvement over the early eyepatches.<sup id="cite_ref-dawkins_33-0" class="reference"><a href="#cite_note-dawkins-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p><p>Overgrowths of transparent cells prevented contamination and parasitic infestation. The chamber contents, now segregated, could slowly specialize into a transparent humour, for optimizations such as colour filtering, higher <a href="/wiki/Refractive_index" title="Refractive index">refractive index</a>, blocking of <a href="/wiki/Ultraviolet" title="Ultraviolet">ultraviolet</a> radiation, or the ability to operate in and out of water. The layer may, in certain classes, be related to the <a href="/wiki/Ecdysis" title="Ecdysis">moulting</a> of the organism's shell or skin. An example of this can be observed in <a href="/wiki/Onychophorans" class="mw-redirect" title="Onychophorans">Onychophorans</a> where the cuticula of the shell continues to the cornea. The cornea is composed of either one or two cuticular layers depending on how recently the animal has moulted.<sup id="cite_ref-Schoenemann2008_34-0" class="reference"><a href="#cite_note-Schoenemann2008-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Along with the lens and two humors, the cornea is responsible for converging light and aiding the focusing of it on the back of the retina. The cornea protects the eyeball while at the same time accounting for approximately 2/3 of the eye's total refractive power.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> </p><p>It is likely that a key reason eyes specialize in detecting a specific, narrow range of wavelengths on the <a href="/wiki/Electromagnetic_spectrum" title="Electromagnetic spectrum">electromagnetic spectrum</a>—the <a href="/wiki/Visible_spectrum" title="Visible spectrum">visible spectrum</a>—is that the earliest species to develop <a href="/wiki/Photosensitivity" title="Photosensitivity">photosensitivity</a> were aquatic, and water filters out <a href="/wiki/Electromagnetic_radiation" title="Electromagnetic radiation">electromagnetic radiation</a> except for a range of wavelengths, the shorter of which we refer to as blue, through to longer wavelengths we identify as red. This same light-filtering property of water also influenced the photosensitivity of plants.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Lens_formation_and_diversification">Lens formation and diversification</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=6" title="Edit section: Lens formation and diversification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Focus_in_an_eye.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Focus_in_an_eye.svg/220px-Focus_in_an_eye.svg.png" decoding="async" width="220" height="74" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Focus_in_an_eye.svg/330px-Focus_in_an_eye.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/Focus_in_an_eye.svg/440px-Focus_in_an_eye.svg.png 2x" data-file-width="800" data-file-height="270" /></a><figcaption>Light from a distant object and a near object being focused by changing the curvature of the <a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">lens</a></figcaption></figure> <p>In a lensless eye, the light emanating from a distant point hits the back of the eye with about the same size as the eye's <a href="/wiki/Aperture" title="Aperture">aperture</a>. With the addition of a lens this incoming light is concentrated on a smaller surface area, without reducing the overall intensity of the stimulus.<sup id="cite_ref-Nilsson,_D_1994_6-1" class="reference"><a href="#cite_note-Nilsson,_D_1994-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The focal length of an early <a href="/wiki/Lobopod" class="mw-redirect" title="Lobopod">lobopod</a> with lens-containing simple eyes focused the image <i>behind</i> the retina, so while no part of the image could be brought into focus, the intensity of light allowed the organism to see in deeper (and therefore darker) waters.<sup id="cite_ref-Schoenemann2008_34-1" class="reference"><a href="#cite_note-Schoenemann2008-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> A subsequent increase of the lens's <a href="/wiki/Refractive_index" title="Refractive index">refractive index</a> probably resulted in an in-focus image being formed.<sup id="cite_ref-Schoenemann2008_34-2" class="reference"><a href="#cite_note-Schoenemann2008-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Note that this optical layout has not been found, nor is it expected to be found. <a href="/wiki/Fossil" title="Fossil">Fossilization</a> rarely preserves soft tissues, and even if it did, the new humour would almost certainly close as the remains desiccated, or as sediment overburden forced the layers together, making the fossilized eye resemble the previous layout. </p> <div class="mw-heading mw-heading3"><h3 id="Crystallins">Crystallins</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=7" title="Edit section: Crystallins"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Krilleyekils.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Krilleyekils.jpg/220px-Krilleyekils.jpg" decoding="async" width="220" height="210" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/64/Krilleyekils.jpg/330px-Krilleyekils.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/64/Krilleyekils.jpg/440px-Krilleyekils.jpg 2x" data-file-width="1574" data-file-height="1505" /></a><figcaption>Compound eye of <a href="/wiki/Antarctic_krill" title="Antarctic krill">Antarctic krill</a></figcaption></figure> <p>Vertebrate <a href="/wiki/Lens_(anatomy)" class="mw-redirect" title="Lens (anatomy)">lenses</a> are composed of adapted <a href="/wiki/Epithelium" title="Epithelium">epithelial</a> cells which have high concentrations of the protein <a href="/wiki/Crystallin" title="Crystallin">crystallin</a>. These crystallins belong to two major families, the α-crystallins and the βγ-crystallins. Both categories of proteins were originally used for other functions in organisms, but eventually adapted for vision in animal eyes.<sup id="cite_ref-Slingsby,_C._2013_39-0" class="reference"><a href="#cite_note-Slingsby,_C._2013-39"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> In the embryo, the lens is living tissue, but the cellular machinery is not transparent so must be removed before the organism can see. Removing the machinery means the lens is composed of dead cells, packed with crystallins. These crystallins are special because they have the unique characteristics required for transparency and function in the lens such as tight packing, resistance to crystallization, and extreme longevity, as they must survive for the entirety of the organism's life.<sup id="cite_ref-Slingsby,_C._2013_39-1" class="reference"><a href="#cite_note-Slingsby,_C._2013-39"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> The <a href="/wiki/Refractive_index" title="Refractive index">refractive index</a> <a href="/wiki/Gradient_index_optics" class="mw-redirect" title="Gradient index optics">gradient</a> which makes the lens useful is caused by the radial shift in crystallin concentration in different parts of the lens, rather than by the specific type of protein: it is not the presence of crystallin, but the relative distribution of it, that renders the lens useful.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p><p>It is biologically difficult to maintain a transparent layer of cells.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Deposition of transparent, nonliving material eased the need for nutrient supply and waste removal. It's a common assumption that <a href="/wiki/Trilobites" class="mw-redirect" title="Trilobites">Trilobites</a> used <a href="/wiki/Calcite" title="Calcite">calcite</a>, a mineral which today is known to be used for vision only in a single species of <a href="/wiki/Brittle_star" title="Brittle star">brittle star</a>.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> Studies of eyes from 55 million years old crane fly fossils from the <a href="/wiki/Fur_Formation" title="Fur Formation">Fur Formation</a> indicates that the calcite in the eyes of trilobites is a result of taphonomic and diagenetic processes and not an original feature.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> In other <a href="/wiki/Compound_eye" title="Compound eye">compound eyes</a> and camera eyes, the material is <a href="/wiki/Crystallin" title="Crystallin">crystallin</a>.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> A gap between tissue layers naturally forms a biconvex shape, which is optically and mechanically ideal for substances of normal<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (March 2017)">clarification needed</span></a></i>]</sup> refractive index. A biconvex lens confers not only optical resolution, but aperture and low-light ability, as resolution is now decoupled from hole size – which slowly increases again, free from the circulatory constraints. </p> <div class="mw-heading mw-heading3"><h3 id="Aqueous_humor,_iris,_and_cornea"><span id="Aqueous_humor.2C_iris.2C_and_cornea"></span>Aqueous humor, iris, and cornea</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=8" title="Edit section: Aqueous humor, iris, and cornea"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Independently, a transparent layer and a nontransparent layer may split forward from the lens: a separate <a href="/wiki/Cornea" title="Cornea">cornea</a> and <a href="/wiki/Iris_(anatomy)" title="Iris (anatomy)">iris</a>. (These may happen before or after crystal deposition, or not at all.) Separation of the forward layer again forms a humour, the <a href="/wiki/Aqueous_humour" title="Aqueous humour">aqueous humour</a>. This increases refractive power and again eases circulatory problems. Formation of a nontransparent ring allows more blood vessels, more circulation, and larger eye sizes. This flap around the perimeter of the lens also masks optical imperfections, which are more common at lens edges. The need to mask lens imperfections gradually increases with lens curvature and power, overall lens and eye size, and the resolution and aperture needs of the organism, driven by hunting or survival requirements. This type is now functionally identical to the eye of most vertebrates, including humans. Indeed, "the basic pattern of all vertebrate eyes is similar."<sup id="cite_ref-Ali&Klyne1985_45-0" class="reference"><a href="#cite_note-Ali&Klyne1985-45"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Other_developments">Other developments</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=9" title="Edit section: Other developments"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Color_vision">Color vision</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=10" title="Edit section: Color vision"><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">Further information: <a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Evolution of color vision</a></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-More_citations_needed_section plainlinks metadata ambox ambox-content ambox-Refimprove" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Question_book-new.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/50px-Question_book-new.svg.png" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/75px-Question_book-new.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/99/Question_book-new.svg/100px-Question_book-new.svg.png 2x" data-file-width="512" data-file-height="399" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>needs additional citations for <a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability">verification</a></b>.<span class="hide-when-compact"> Please help <a href="/wiki/Special:EditPage/Evolution_of_the_eye" title="Special:EditPage/Evolution of the eye">improve this article</a> by <a href="/wiki/Help:Referencing_for_beginners" title="Help:Referencing for beginners">adding citations to reliable sources</a> in this section. Unsourced material may be challenged and removed.</span> <span class="date-container"><i>(<span class="date">October 2016</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>Five classes of visual <a href="/wiki/Opsin" title="Opsin">opsins</a> are found in vertebrates. All but one of these developed prior to the divergence of <a href="/wiki/Cyclostomata" class="mw-redirect" title="Cyclostomata">Cyclostomata</a> and fish.<sup id="cite_ref-Osorio&Vorobyev_46-0" class="reference"><a href="#cite_note-Osorio&Vorobyev-46"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> The five opsin classes are variously adapted depending on the light spectrum encountered. As <a href="/wiki/Color_of_water" title="Color of water">light travels through water</a>, longer wavelengths, such as reds and yellows, are absorbed more quickly than the shorter <a href="/wiki/Wavelength" title="Wavelength">wavelengths</a> of the greens and blues. This creates a gradient in the <a href="/wiki/Spectral_density" title="Spectral density">spectral power density</a>, with the average wavelength becoming shorter as water depth increases.<sup id="cite_ref-Guehmann2015_47-0" class="reference"><a href="#cite_note-Guehmann2015-47"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> The visual opsins in fish are more sensitive to the range of light in their habitat and depth. However, land environments do not vary in wavelength composition, so that the opsin sensitivities among land vertebrates does not vary much. This directly contributes to the significant presence of communication colors.<sup id="cite_ref-Osorio&Vorobyev_46-1" class="reference"><a href="#cite_note-Osorio&Vorobyev-46"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Color vision gives distinct <a href="/wiki/Selection_(biology)" class="mw-redirect" title="Selection (biology)">selective</a> advantages, such as better recognition of predators, food, and mates. Indeed, it is possible that simple sensory-neural mechanisms may selectively control general behavior patterns, such as escape, foraging, and hiding. Many examples of wavelength-specific behaviors have been identified, in two primary groups: Below 450 nm, associated with direct light, and above 450 nm, associated with reflected light.<sup id="cite_ref-Menzel1979_48-0" class="reference"><a href="#cite_note-Menzel1979-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> As opsin molecules were tuned to detect different wavelengths of light, at some point <a href="/wiki/Color_vision" title="Color vision">color vision</a> developed when the photoreceptor cells used differently tuned opsins.<sup id="cite_ref-photons_31-1" class="reference"><a href="#cite_note-photons-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> This may have happened at any of the early stages of the eye's evolution, and may have disappeared and reevolved as relative selective pressures on the lineage varied. </p> <div class="mw-heading mw-heading4"><h4 id="Polarization_vision">Polarization vision</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=11" title="Edit section: Polarization vision"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Polarization_(waves)" title="Polarization (waves)">Polarization</a> is the organization of disordered light into linear arrangements, which occurs when light passes through slit like filters, as well as when passing into a new medium. Sensitivity to polarized light is especially useful for organisms whose habitats are located more than a few meters under water. In this environment, color vision is less dependable, and therefore a weaker selective factor. While most photoreceptors have the ability to distinguish partially polarized light, terrestrial vertebrates' membranes are orientated perpendicularly, such that they are insensitive to polarized light.<sup id="cite_ref-Cronin_et_al_49-0" class="reference"><a href="#cite_note-Cronin_et_al-49"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> However, some fish can discern polarized light, demonstrating that they possess some linear photoreceptors. Additionally, cuttlefish are capable of perceiving the polarization of light with high visual fidelity, although they appear to lack any significant capacity for color differentiation.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Like color vision, sensitivity to polarization can aid in an organism's ability to differentiate surrounding objects and individuals. Because of the marginal reflective interference of polarized light, it is often used for orientation and navigation, as well as distinguishing concealed objects, such as disguised prey.<sup id="cite_ref-Cronin_et_al_49-1" class="reference"><a href="#cite_note-Cronin_et_al-49"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Focusing_mechanism">Focusing mechanism</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=12" title="Edit section: Focusing mechanism"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By utilizing the iris <a href="/wiki/Sphincter" title="Sphincter">sphincter</a> muscle and the <a href="/wiki/Ciliary_body" title="Ciliary body">ciliary body</a>, some species move the lens back and forth, some <a href="/wiki/Accommodation_(eye)" class="mw-redirect" title="Accommodation (eye)">stretch the lens</a> flatter. Another mechanism regulates focusing chemically and independently of these two, by controlling growth of the eye and maintaining focal length. In addition, the pupil shape can be used to predict the focal system being utilized. A slit pupil can indicate the common multifocal system, while a circular pupil usually specifies a monofocal system. When using a circular form, the pupil will constrict under bright light, increasing the f-number, and will dilate when dark in order to decrease the depth of focus.<sup id="cite_ref-Malstrom&Kroger_51-0" class="reference"><a href="#cite_note-Malstrom&Kroger-51"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> Note that a focusing method is not a requirement. As photographers know, focal errors increase as <a href="/wiki/Aperture" title="Aperture">aperture</a> increases. Thus, countless organisms with small eyes are active in direct sunlight and survive with no focus mechanism at all. As a species grows larger, or transitions to dimmer environments, a means of focusing need only appear gradually. </p> <div class="mw-heading mw-heading4"><h4 id="Placement">Placement</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=13" title="Edit section: Placement"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Predators generally have eyes on the front of their heads for better <a href="/wiki/Depth_perception" title="Depth perception">depth perception</a> to focus on prey. Prey animals' eyes tend to be on the side of the head giving a wide field of view to detect predators from any direction.<sup id="cite_ref-BLM_52-0" class="reference"><a href="#cite_note-BLM-52"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Flatfish" title="Flatfish">Flatfish</a> are predators which lie on their side on the bottom, and have eyes placed asymmetrically on the same side of the head. A <a href="/wiki/Transitional_fossil" title="Transitional fossil">transitional fossil</a> from the common symmetric position to the asymmetric position is <a href="/wiki/Amphistium" title="Amphistium">Amphistium</a>. </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=Evolution_of_the_eye&action=edit&section=14" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Ocelloid" title="Ocelloid">Ocelloid</a></li> <li><a href="/wiki/Sensory_organs_of_gastropods#Eyes" title="Sensory organs of gastropods">Sensory organs of gastropods § Eyes</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="Explanatory_footnotes">Explanatory footnotes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=15" title="Edit section: Explanatory footnotes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text"><a href="/wiki/David_Berlinski" title="David Berlinski">David Berlinski</a>, an intelligent design proponent, questioned the basis of the calculations, and the author of the original paper refuted Berlinski's criticism. <ul><li><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="CITEREFBerlinski,_David2001" class="citation magazine cs1"><a href="/wiki/David_Berlinski" title="David Berlinski">Berlinski, David</a> (April 2001). <a rel="nofollow" class="external text" href="http://www.discovery.org/a/1360">"The Vexing Eye"</a>. <i>Commentary</i>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Commentary&rft.atitle=The+Vexing+Eye&rft.date=2001-04&rft.au=Berlinski%2C+David&rft_id=http%3A%2F%2Fwww.discovery.org%2Fa%2F1360&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNilsson,_Dan-E." class="citation web cs1">Nilsson, Dan-E. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20030915124605/http://www.talkreason.org/articles/blurred.cfm#lund">"Beware of Pseudo-science: a response to David Berlinski's attack on my calculation of how long it takes for an eye to evolve"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.talkreason.org/articles/blurred.cfm#lund">the original</a> on 15 September 2003.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Beware+of+Pseudo-science%3A+a+response+to+David+Berlinski%27s+attack+on+my+calculation+of+how+long+it+takes+for+an+eye+to+evolve&rft.au=Nilsson%2C+Dan-E.&rft_id=http%3A%2F%2Fwww.talkreason.org%2Farticles%2Fblurred.cfm%23lund&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://www.pbs.org/wgbh/evolution/library/01/1/l_011_01.html">"Evolution of the Eye" on PBS</a></li></ul> </span></li> <li id="cite_note-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-29">^</a></b></span> <span class="reference-text">The precise number varies from author to author.</span> </li> </ol></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=Evolution_of_the_eye&action=edit&section=16" 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: 30em;"> <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">""An important cue for understanding eye evolution is the distinction between different types of photoreceptor cells. Salvini-Plawen and Mayr ( 1977 ) noted a remarkable diversity of photoreceptor cell morphology across the animal kingdom, and suggested that photoreceptors evolved independently numerous times." – Land, M.F. and Nilsson, D.-E., <i>Animal Eyes</i>(2-nd ed.), Oxford University Press, Oxford (2012), p. 7.</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"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeeJagoGarcia-BellidoEdgecombe2011" class="citation journal cs1">Lee, M. S. Y.; Jago, J. B.; Garcia-Bellido, D. C.; Edgecombe, G. E.; Gehling, J. G; Paterson, J. R. (2011). 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"pupil shape and lens optics in the eyes of terrestrial vertebrates". <i>The Journal of Experimental Biology</i>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Experimental+Biology&rft.atitle=pupil+shape+and+lens+optics+in+the+eyes+of+terrestrial+vertebrates&rft.date=2006&rft.aulast=Malstrom&rft.aufirst=T&rft.au=Kroger%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></span> </li> <li id="cite_note-BLM-52"><span class="mw-cite-backlink"><b><a href="#cite_ref-BLM_52-0">^</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/20110614220937/http://www.blm.gov/id/st/en/prog/wildlife/carnivores.html">"Carnivores"</a>. U.S. Department of the Interior, Bureau of Land Management. 14 December 2009. Archived from <a rel="nofollow" class="external text" href="https://www.blm.gov/id/st/en/prog/wildlife/carnivores.html">the original</a> on 14 June 2011<span class="reference-accessdate">. Retrieved <span class="nowrap">28 March</span> 2011</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Carnivores&rft.pub=U.S.+Department+of+the+Interior%2C+Bureau+of+Land+Management&rft.date=2009-12-14&rft_id=http%3A%2F%2Fwww.blm.gov%2Fid%2Fst%2Fen%2Fprog%2Fwildlife%2Fcarnivores.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></span> </li> <li id="cite_note-53"><span class="mw-cite-backlink"><b><a href="#cite_ref-53">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBoroditsky1999" class="citation web cs1">Boroditsky, Lera (24 June 1999). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20100705103809/http://www-psych.stanford.edu/~lera/psych115s/notes/lecture2/">"Light & Eyes: Lecture Notes"</a>. <i>Lecture Notes</i>. <a href="/wiki/Stanford" class="mw-redirect" title="Stanford">Stanford</a>. Archived from <a rel="nofollow" class="external text" href="http://www-psych.stanford.edu/~lera/psych115s/notes/lecture2/">the original</a> on 5 July 2010<span class="reference-accessdate">. Retrieved <span class="nowrap">11 May</span> 2010</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Lecture+Notes&rft.atitle=Light+%26+Eyes%3A+Lecture+Notes&rft.date=1999-06-24&rft.aulast=Boroditsky&rft.aufirst=Lera&rft_id=http%3A%2F%2Fwww-psych.stanford.edu%2F~lera%2Fpsych115s%2Fnotes%2Flecture2%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=17" title="Edit section: Further reading"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLambCollinPugh2007" class="citation journal cs1">Lamb TD, Collin SP, Pugh EN (December 2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3143066">"Evolution of the vertebrate eye: opsins, photoreceptors, retina and eye cup"</a>. <i>Nature Reviews Neuroscience</i>. <b>8</b> (12): <span class="nowrap">960–</span>76. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnrn2283">10.1038/nrn2283</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/PMC3143066">3143066</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18026166">18026166</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews+Neuroscience&rft.atitle=Evolution+of+the+vertebrate+eye%3A+opsins%2C+photoreceptors%2C+retina+and+eye+cup&rft.volume=8&rft.issue=12&rft.pages=%3Cspan+class%3D%22nowrap%22%3E960-%3C%2Fspan%3E76&rft.date=2007-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3143066%23id-name%3DPMC&rft_id=info%3Apmid%2F18026166&rft_id=info%3Adoi%2F10.1038%2Fnrn2283&rft.aulast=Lamb&rft.aufirst=TD&rft.au=Collin%2C+SP&rft.au=Pugh%2C+EN&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3143066&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span> <a rel="nofollow" class="external text" href="http://www.nature.com/nrn/journal/v8/n12/fig_tab/nrn2283_F4.html">Illustration.</a> <a rel="nofollow" class="external text" href="http://scienceblogs.com/pharyngula/2007/12/21/evolution-of-vertebrate-eyes">Review</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLamb2011" class="citation journal cs1">Lamb, TD (2011). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20131212204529/http://physics.okstate.edu/axie/courses/4313/2012fall/C7_B2_reading-evolution-of-eye-2011.pdf">"Evolution of the Eye"</a> <span class="cs1-format">(PDF)</span>. <i>Scientific American</i>. <b>305</b> (1): <span class="nowrap">64–</span>69. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2011SciAm.305f..64L">2011SciAm.305f..64L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fscientificamerican0711-64">10.1038/scientificamerican0711-64</a>. Archived from <a rel="nofollow" class="external text" href="http://physics.okstate.edu/axie/courses/4313/2012fall/C7_B2_reading-evolution-of-eye-2011.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 12 December 2013<span class="reference-accessdate">. Retrieved <span class="nowrap">28 April</span> 2013</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scientific+American&rft.atitle=Evolution+of+the+Eye&rft.volume=305&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E64-%3C%2Fspan%3E69&rft.date=2011&rft_id=info%3Adoi%2F10.1038%2Fscientificamerican0711-64&rft_id=info%3Abibcode%2F2011SciAm.305f..64L&rft.aulast=Lamb&rft.aufirst=TD&rft_id=http%3A%2F%2Fphysics.okstate.edu%2Faxie%2Fcourses%2F4313%2F2012fall%2FC7_B2_reading-evolution-of-eye-2011.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLandNilsson2012" class="citation book cs1">Land, Michael F.; Nilsson, Dan-Eric (2012). "The origin of vision". <i>Animal Eyes</i> (2 ed.). Oxford: Oxford University Press. pp. <span class="nowrap">1–</span>22. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-958114-6" title="Special:BookSources/978-0-19-958114-6"><bdi>978-0-19-958114-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=The+origin+of+vision&rft.btitle=Animal+Eyes&rft.place=Oxford&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1-%3C%2Fspan%3E22&rft.edition=2&rft.pub=Oxford+University+Press&rft.date=2012&rft.isbn=978-0-19-958114-6&rft.aulast=Land&rft.aufirst=Michael+F.&rft.au=Nilsson%2C+Dan-Eric&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://link.springer.com/journal/12052/volumes-and-issues/1-4">Special Issue "The Evolution of Eyes"</a> in <i>Evolution: Education and Outreach</i>, Volume 1, Number 4 / October 2008. 26 articles, free access.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFIvan_R._Schwab2012" class="citation book cs1">Ivan R. Schwab (2012). <i>Evolution's Witness: How Eyes Evolved</i>. New York: Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-536974-8" title="Special:BookSources/978-0-19-536974-8"><bdi>978-0-19-536974-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%27s+Witness%3A+How+Eyes+Evolved&rft.place=New+York&rft.pub=Oxford+University+Press&rft.date=2012&rft.isbn=978-0-19-536974-8&rft.au=Ivan+R.+Schwab&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHayakawaTakakuHwangHoriguchi2015" class="citation journal cs1">Hayakawa S, Takaku Y, Hwang JS, Horiguchi T, Suga H, Gehring W, et al. 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Archived from <a rel="nofollow" class="external text" href="http://repository.kaust.edu.sa/kaust/handle/10754/566109">the original</a> on 12 April 2018<span class="reference-accessdate">. Retrieved <span class="nowrap">2 July</span> 2019</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Eye-like+ocelloids+are+built+from+different+endosymbiotically+acquired+components&rft.volume=523&rft.issue=7559&rft.pages=%3Cspan+class%3D%22nowrap%22%3E204-%3C%2Fspan%3E7&rft.date=2015&rft_id=info%3Ahdl%2F10754%2F566109&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4462376%23id-name%3DS2CID&rft_id=info%3Abibcode%2F2015Natur.523..204G&rft_id=info%3Apmid%2F26131935&rft_id=info%3Adoi%2F10.1038%2Fnature14593&rft.au=Gregory+S.+Gavelis&rft.au=Shiho+Hayakawa&rft.au=Richard+A.+White+III&rft.au=Takashi+Gojobori&rft.au=Curtis+A.+Suttle&rft.au=Patrick+J.+Keeling&rft.au=Brian+S.+Leander&rft_id=http%3A%2F%2Frepository.kaust.edu.sa%2Fkaust%2Fhandle%2F10754%2F566109&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOakleySpeiser2015" class="citation journal cs1">Oakley, Todd H.; Speiser, Daniel I. (2015). <a rel="nofollow" class="external text" href="https://www.biorxiv.org/content/biorxiv/early/2015/03/26/017129.full.pdf">"How Complexity Originates: The Evolution of Animal Eyes"</a> <span class="cs1-format">(PDF)</span>. <i>Annual Review of Ecology, Evolution, and Systematics</i>. <b>46</b>: <span class="nowrap">237–</span>260. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev-ecolsys-110512-135907">10.1146/annurev-ecolsys-110512-135907</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Ecology%2C+Evolution%2C+and+Systematics&rft.atitle=How+Complexity+Originates%3A+The+Evolution+of+Animal+Eyes&rft.volume=46&rft.pages=%3Cspan+class%3D%22nowrap%22%3E237-%3C%2Fspan%3E260&rft.date=2015&rft_id=info%3Adoi%2F10.1146%2Fannurev-ecolsys-110512-135907&rft.aulast=Oakley&rft.aufirst=Todd+H.&rft.au=Speiser%2C+Daniel+I.&rft_id=https%3A%2F%2Fwww.biorxiv.org%2Fcontent%2Fbiorxiv%2Fearly%2F2015%2F03%2F26%2F017129.full.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEd_Youngphotographs_by_David_Liittschwager2016" class="citation journal cs1">Ed Young; photographs by David Liittschwager (February 2016). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170624112924/http://www.nationalgeographic.com/magazine/2016/02/evolution-of-eyes/">"Inside the Eye: Nature's Most Exquisite Creation"</a>. <i><a href="/wiki/National_Geographic_(magazine)" class="mw-redirect" title="National Geographic (magazine)">National Geographic</a></i>. <b>229</b> (2): <span class="nowrap">30–</span>57. Archived from <a rel="nofollow" class="external text" href="https://www.nationalgeographic.com/magazine/2016/02/evolution-of-eyes/">the original</a> on 24 June 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=National+Geographic&rft.atitle=Inside+the+Eye%3A+Nature%27s+Most+Exquisite+Creation&rft.volume=229&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E30-%3C%2Fspan%3E57&rft.date=2016-02&rft.au=Ed+Young&rft.au=photographs+by+David+Liittschwager&rft_id=https%3A%2F%2Fwww.nationalgeographic.com%2Fmagazine%2F2016%2F02%2Fevolution-of-eyes%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_eye&action=edit&section=18" title="Edit section: External links"><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 class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.pbs.org/wgbh/evolution/library/01/1/l_011_01.html">"Evolution of the Eye"</a>. <i>WGBH Educational Foundation and Clear Blue Sky Productions</i>. PBS. 2001.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=WGBH+Educational+Foundation+and+Clear+Blue+Sky+Productions&rft.atitle=Evolution+of+the+Eye&rft.date=2001&rft_id=https%3A%2F%2Fwww.pbs.org%2Fwgbh%2Fevolution%2Flibrary%2F01%2F1%2Fl_011_01.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+eye" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=fOtP7HEuDYA">Creationism Disproved?</a> Video from the <a href="/wiki/National_Center_for_Science_Education" title="National Center for Science Education">National Center for Science Education</a> on the evolution of the eye</li> <li><a rel="nofollow" class="external text" href="https://evolution-outreach.biomedcentral.com/articles/10.1007/s12052-008-0081-4"><i>Evolution: Education and Outreach</i> Special Issue: Evolution and Eyes</a> volume 1, number 4, October 2008, pages 351–559. 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href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</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/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</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/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</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/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</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/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</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_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a class="mw-selflink selflink">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</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/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</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/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</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/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</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/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</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/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</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/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></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:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</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="Evolutionary_psychology1033" 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:Evolutionary_psychology" title="Template:Evolutionary psychology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_psychology" title="Template talk:Evolutionary psychology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_psychology" title="Special:EditPage/Template:Evolutionary psychology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_psychology1033" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</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/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</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/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</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/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a href="/wiki/Motor_control" title="Motor control">Motor control</a>/<a href="/wiki/Motor_skill" title="Motor skill">skill</a></li> <li><a href="/wiki/Human_multitasking" title="Human multitasking">Multitasking</a></li> <li><a href="/wiki/Neuroscience_of_sleep" title="Neuroscience of sleep">Sleep</a></li> <li><a href="/wiki/Visual_perception" title="Visual perception">Visual perception</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">Color vision</a></li> <li><a class="mw-selflink selflink">Eye</a></li> <li><a href="/wiki/Na%C3%AFve_physics" title="Naïve physics">Naïve physics</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_psychology_and_culture" title="Evolutionary psychology and culture">Culture</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/Evolutionary_aesthetics" title="Evolutionary aesthetics">Aesthetics</a> <ul><li><a href="/wiki/Darwinian_literary_studies" title="Darwinian literary studies">Literary criticism</a></li> <li><a href="/wiki/Evolutionary_musicology" title="Evolutionary musicology">Musicology</a></li></ul></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Anthropology</a> <ul><li><a href="/wiki/Biological_anthropology" title="Biological anthropology">Biological</a></li></ul></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Crime</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Language</a> <ul><li><a href="/wiki/Origin_of_language" title="Origin of language">Origin</a></li> <li><a href="/wiki/Evolutionary_psychology_of_language" title="Evolutionary psychology of language">Psychology</a></li> <li><a href="/wiki/Origin_of_speech" title="Origin of speech">Speech</a></li></ul></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a> <ul><li><a href="/wiki/Moral_foundations_theory" title="Moral foundations theory">Moral foundations</a></li></ul></li> <li><a href="/wiki/Evolutionary_psychology_of_religion" title="Evolutionary psychology of religion">Religion</a> <ul><li><a href="/wiki/Evolutionary_origin_of_religions" class="mw-redirect" title="Evolutionary origin of religions">Origin</a></li></ul></li> <li><a href="/wiki/Cultural_universal" title="Cultural universal">Universals</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolutionary_developmental_psychology" title="Evolutionary developmental psychology">Development</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/Attachment_theory" title="Attachment theory">Attachment</a></li> <li><a href="/wiki/Human_bonding" title="Human bonding">Bonding</a></li> <li><a href="/wiki/Affectional_bond" title="Affectional bond">Affectional</a>/<a href="/wiki/Maternal_bond" title="Maternal bond">maternal</a>/<a href="/wiki/Paternal_bond" title="Paternal bond">paternal bond</a></li> <li><a href="/wiki/Maternal_deprivation" title="Maternal deprivation">Caregiver deprivation</a></li> <li><a href="/wiki/Attachment_in_children" title="Attachment in children">Childhood attachment</a></li> <li><a href="/wiki/Cinderella_effect" title="Cinderella effect">Cinderella effect</a></li> <li><a href="/wiki/Cognitive_development" title="Cognitive development">Cognitive development</a></li> <li><a href="/wiki/Evolutionary_educational_psychology" title="Evolutionary educational psychology">Education</a></li> <li><a href="/wiki/Language_acquisition" title="Language acquisition">Language acquisition</a></li> <li><a href="/wiki/Personality_development" title="Personality development">Personality development</a></li> <li><a href="/wiki/Socialization" title="Socialization">Socialization</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Ergonomics" title="Ergonomics">Human factors</a> /<br /><a href="/wiki/Evolutionary_psychiatry" title="Evolutionary psychiatry">Mental health</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/Cognitive_ergonomics" title="Cognitive ergonomics">Cognitive ergonomics</a> <ul><li><a href="/wiki/Computer-mediated_communication" title="Computer-mediated communication">Computer-mediated communication</a></li> <li><a href="/wiki/Engineering_psychology" title="Engineering psychology">Engineering psychology</a></li> <li><a href="/wiki/Human%E2%80%93computer_interaction" title="Human–computer interaction">Human–computer interaction</a></li> <li><a href="/wiki/Media_naturalness_theory" title="Media naturalness theory">Media naturalness theory</a></li> <li><a href="/wiki/Neuroergonomics" title="Neuroergonomics">Neuroergonomics</a></li></ul></li> <li><a href="/wiki/Evolutionary_approaches_to_depression" title="Evolutionary approaches to depression">Depression</a></li> <li><a href="/wiki/Digital_media_use_and_mental_health" title="Digital media use and mental health">Digital media use and mental health</a></li> <li><a href="/wiki/Accident-proneness" title="Accident-proneness">Hypophobia</a></li> <li><a href="/wiki/Imprinted_brain_hypothesis" title="Imprinted brain hypothesis">Imprinted brain hypothesis</a></li> <li><a href="/wiki/Mind-blindness" title="Mind-blindness">Mind-blindness</a></li> <li><a href="/wiki/Psychological_effects_of_Internet_use" title="Psychological effects of Internet use">Psychological effects of Internet use</a></li> <li><a href="/wiki/Rank_theory_of_depression" title="Rank theory of depression">Rank theory of depression</a></li> <li><a href="/wiki/Evolution_of_schizophrenia" title="Evolution of schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Screen_time" title="Screen time">Screen time</a></li> <li><a href="/wiki/Smartphones_and_pedestrian_safety" title="Smartphones and pedestrian safety">Smartphones and pedestrian safety</a></li> <li><a href="/wiki/Social_aspects_of_television" title="Social aspects of television">Social aspects of television</a></li> <li><a href="/wiki/Societal_effects_of_cars" title="Societal effects of cars">Societal effects of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</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/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</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/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</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%">Academic disciplines</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_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research 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/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li> <li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</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/Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li> <li><a href="/wiki/Determinism" title="Determinism">Determinism</a>/<a href="/wiki/Indeterminism" title="Indeterminism">indeterminism</a> <ul><li><a href="/wiki/Biological_determinism" title="Biological determinism">Biological determinism</a></li> <li><a href="/wiki/Connectionism" title="Connectionism">Connectionism</a></li> <li><a href="/wiki/Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li> <li><a href="/wiki/Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li> <li><a href="/wiki/Social_constructionism" title="Social constructionism">Social constructionism</a></li> <li><a href="/wiki/Social_determinism" title="Social determinism">Social determinism</a></li> <li><a href="/wiki/Standard_social_science_model" title="Standard social science model">Standard social science model</a></li></ul></li> <li><a href="/wiki/Functionalism_(philosophy_of_mind)" title="Functionalism (philosophy of mind)">Functionalism</a></li> <li><a href="/wiki/Memetics" title="Memetics">Memetics</a></li> <li><a href="/wiki/Multilineal_evolution" title="Multilineal evolution">Multilineal evolution</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Neoevolutionism" title="Neoevolutionism">Neoevolutionism</a></li> <li><a href="/wiki/Sociocultural_evolution" title="Sociocultural evolution">Sociocultural evolution</a></li> <li><a href="/wiki/Unilineal_evolution" title="Unilineal evolution">Unilineal evolution</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><b><a href="/wiki/List_of_evolutionary_psychologists" title="List of evolutionary psychologists">Evolutionary psychologists</a></b></li> <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:Evolutionary_psychology" title="Category:Evolutionary psychology">Evolutionary psychology</a></b></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <b><a href="/wiki/Portal:Psychology" title="Portal:Psychology">Psychology portal</a></b></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <b><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</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="Vision_in_animals86" 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_animals86" 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 class="mw-selflink selflink">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 href="/wiki/Eyespot_(mimicry)" title="Eyespot (mimicry)">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 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