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class="vector-toc-numb">2</span> <span>Role of embryology</span> </div> </a> <ul id="toc-Role_of_embryology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Randomizing_access_and_increasing_size" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Randomizing_access_and_increasing_size"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Randomizing access and increasing size</span> </div> </a> <ul id="toc-Randomizing_access_and_increasing_size-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Brain_re-organization" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Brain_re-organization"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Brain re-organization</span> </div> </a> <ul id="toc-Brain_re-organization-sublist" class="vector-toc-list"> </ul> </li> <li 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cortex</span> </div> </a> <ul id="toc-Development_of_the_human_cerebral_cortex-sublist" class="vector-toc-list"> <li id="toc-MCPH1_and_ASPM" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#MCPH1_and_ASPM"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1.1</span> <span>MCPH1 and ASPM</span> </div> </a> <ul id="toc-MCPH1_and_ASPM-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-ZEB2" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#ZEB2"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1.2</span> <span>ZEB2</span> </div> </a> <ul id="toc-ZEB2-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-NOVA1" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#NOVA1"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1.3</span> <span>NOVA1</span> </div> </a> <ul id="toc-NOVA1-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-SRGAP2C_and_neuronal_maturation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#SRGAP2C_and_neuronal_maturation"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>SRGAP2C and neuronal maturation</span> </div> </a> <ul id="toc-SRGAP2C_and_neuronal_maturation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-LRRC37B_and_neuronal_electrical_properties" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#LRRC37B_and_neuronal_electrical_properties"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>LRRC37B and neuronal electrical properties</span> </div> </a> <ul id="toc-LRRC37B_and_neuronal_electrical_properties-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Genome_repair" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Genome_repair"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Genome repair</span> </div> </a> <ul id="toc-Genome_repair-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_factors" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Other_factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.5</span> <span>Other factors</span> </div> </a> <ul id="toc-Other_factors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Recently_evolved_traits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Recently_evolved_traits"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.6</span> <span>Recently evolved traits</span> </div> </a> <ul id="toc-Recently_evolved_traits-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Human_brain_evolution" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Human_brain_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Human brain evolution</span> </div> </a> <button aria-controls="toc-Human_brain_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 Human brain evolution subsection</span> </button> <ul id="toc-Human_brain_evolution-sublist" class="vector-toc-list"> <li id="toc-Evolution_of_the_neocortex" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evolution_of_the_neocortex"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Evolution of the neocortex</span> </div> </a> <ul id="toc-Evolution_of_the_neocortex-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Theories_of_human_brain_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Theories_of_human_brain_evolution"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Theories of human brain evolution</span> </div> </a> <ul id="toc-Theories_of_human_brain_evolution-sublist" class="vector-toc-list"> </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">7</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</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">9</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " 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href="https://bn.wikipedia.org/wiki/%E0%A6%AE%E0%A6%B8%E0%A7%8D%E0%A6%A4%E0%A6%BF%E0%A6%B7%E0%A7%8D%E0%A6%95%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="মস্তিষ্কের বিবর্তন – Bangla" lang="bn" hreflang="bn" data-title="মস্তিষ্কের বিবর্তন" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target"><span>বাংলা</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Evoluci%C3%B3n_del_cerebro" title="Evolución del cerebro – Spanish" lang="es" hreflang="es" data-title="Evolución del cerebro" 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_%D9%85%D8%BA%D8%B2" title="فرگشت مغز – Persian" lang="fa" hreflang="fa" data-title="فرگشت مغز" 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<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">Overview of animal and human brain evolution</div> <p class="mw-empty-elt"> </p> <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-Technical plainlinks metadata ambox ambox-style ambox-technical" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article <b>may be too technical for most readers to understand</b>.<span class="hide-when-compact"> Please <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Evolution_of_the_brain&action=edit">help improve it</a> to <a href="/wiki/Wikipedia:Make_technical_articles_understandable" title="Wikipedia:Make technical articles understandable">make it understandable to non-experts</a>, without removing the technical details.</span> <span class="date-container"><i>(<span class="date">January 2022</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:The_brain_from_ape_to_man;_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_(1928)_(20408211835).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg/220px-The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg" decoding="async" width="220" height="291" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6b/The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg/330px-The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6b/The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg/440px-The_brain_from_ape_to_man%3B_a_contribution_to_the_study_of_the_evolution_and_development_of_the_human_brain_%281928%29_%2820408211835%29.jpg 2x" data-file-width="1598" data-file-height="2116" /></a><figcaption>Evolution of the brain from ape to man</figcaption></figure> <p>The <b>evolution of the brain</b> refers to the progressive development and complexity of <a href="/wiki/Neural_structure" class="mw-redirect" title="Neural structure">neural structures</a> over millions of years, resulting in the diverse range of brain sizes and functions observed across different species today, particularly in <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a>. </p><p>The evolution of the brain has exhibited diverging adaptations within taxonomic classes, such as Mammalia, and even more diverse adaptations across other taxonomic classes. Brain-to-body size scales <a href="/wiki/Allometry" title="Allometry">allometrically</a>.<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> This means that as body size changes, so do other physiological, anatomical, and biochemical connections between the brain and body.<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> Small-bodied mammals tend to have relatively large brains compared to their bodies, while larger mammals (such as whales) have smaller brain-to-body ratios. When brain weight is plotted against body weight for primates, the regression line of the sample points can indicate the brain power of a species. For example, lemurs fall below this line, suggesting that for a primate of their size, a larger brain would be expected. In contrast, humans lie well above this line, indicating they are more <a href="/wiki/Encephalization_Quotient" class="mw-redirect" title="Encephalization Quotient">encephalized</a> than lemurs and, in fact, more <a href="/wiki/Encephalization_Quotient" class="mw-redirect" title="Encephalization Quotient">encephalized</a> than any other primate.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> This suggests that human brains have undergone a larger evolutionary increase in complexity relative to size. Some of these changes have been linked to multiple genetic factors, including proteins and other organelles. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Early_history">Early history</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=1" title="Edit section: Early history"><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">Main article: <a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></div> <style data-mw-deduplicate="TemplateStyles:r1233989161">.mw-parser-output .unsolved{margin:0.5em 0 1em 1em;border:#ccc solid;padding:0.35em 0.35em 0.35em 2.2em;background-color:var(--background-color-interactive-subtle);background-image:url("https://upload.wikimedia.org/wikipedia/commons/2/26/Question%2C_Web_Fundamentals.svg");background-position:top 50%left 0.35em;background-size:1.5em;background-repeat:no-repeat}@media(min-width:720px){.mw-parser-output .unsolved{clear:right;float:right;max-width:25%}}.mw-parser-output .unsolved-label{font-weight:bold}.mw-parser-output .unsolved-body{margin:0.35em;font-style:italic}.mw-parser-output .unsolved-more{font-size:smaller}</style> <div role="note" aria-labelledby="unsolved-label-biology" class="unsolved"> <div><span class="unsolved-label" id="unsolved-label-biology">Unsolved problem in biology</span>:</div> <div class="unsolved-body">How and why did the brain evolve?</div> <div class="unsolved-more"><a href="/wiki/List_of_unsolved_problems_in_biology" title="List of unsolved problems in biology">(more unsolved problems in biology)</a></div> </div> <p>One approach to understanding overall brain evolution is to use a <a href="/wiki/Palaeoarchaeology" title="Palaeoarchaeology">paleoarchaeological</a> timeline to trace the necessity for ever increasing complexity in structures that allow for chemical and electrical signaling. Because brains and other <a href="/wiki/Soft_tissue" title="Soft tissue">soft tissues</a> do not fossilize as readily as <a href="/wiki/Mineralized_tissues" title="Mineralized tissues">mineralized tissues</a>, scientists often look to other structures as evidence in the fossil record to get an understanding of brain evolution. This, however, leads to a dilemma as the emergence of organisms with more complex nervous systems with protective bone or other protective tissues that can then readily fossilize occur in the fossil record before evidence for chemical and electrical signaling.<sup id="cite_ref-Cai_2008_4-0" class="reference"><a href="#cite_note-Cai_2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><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> Evidence from 2008 showed that the ability to transmit electrical and chemical signals existed even before more complex multicellular lifeforms.<sup id="cite_ref-Cai_2008_4-1" class="reference"><a href="#cite_note-Cai_2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>Fossilization of brain tissue, as well as other soft tissue, is nonetheless possible, and scientists can infer that the first brain structure appeared at least 521 million years ago, with fossil brain tissue present in sites of exceptional preservation.<sup id="cite_ref-Park_2018_6-0" class="reference"><a href="#cite_note-Park_2018-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Another approach to understanding brain evolution is to look at extant organisms that do not possess complex nervous systems, comparing anatomical features that allow for chemical or electrical messaging. For example, <a href="/wiki/Choanoflagellate" title="Choanoflagellate">choanoflagellates</a> are organisms that possess various <a href="/wiki/Membrane_channel" title="Membrane channel">membrane channels</a> that are crucial to electrical signaling. The membrane channels of choanoflagellates' are <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous</a> to the ones found in animal cells, and this is supported by the evolutionary connection between early choanoflagellates and the ancestors of animals.<sup id="cite_ref-Cai_2008_4-2" class="reference"><a href="#cite_note-Cai_2008-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Another example of extant organisms with the capacity to transmit electrical signals would be the <a href="/wiki/Hexactinellid" title="Hexactinellid">glass sponge</a>, a multicellular organism, which is capable of propagating electrical impulses without the presence of a nervous system.<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> </p><p>Before the evolutionary development of the brain, <a href="/wiki/Nerve_net" title="Nerve net">nerve nets</a>, the simplest form of a <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a> developed. These nerve nets were a sort of precursor for the more evolutionarily advanced brains. They were first observed in <a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a> and consist of a number of neurons spread apart that allow the organism to respond to physical contact. They are able to rudimentarily detect food and other chemicals, but these nerve nets do not allow them to detect the source of the stimulus. </p><p><a href="/wiki/Ctenophora" title="Ctenophora">Ctenophores</a> also demonstrate this crude precursor to a brain or centralized nervous system, however they phylogenetically diverged before the phylum <a href="/wiki/Porifera" class="mw-redirect" title="Porifera">Porifera</a> (the Sponges) and Cnidaria. There are two current theories on the emergence of nerve nets. One theory is that nerve nets may have developed independently in Ctenophores and Cnidarians. The other theory states that a common ancestor may have developed nerve nets, but they were lost in Porifera. While comparing the average neuron size and the packing density the difference between primate and mammal brains is shown.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p><p>A trend in brain evolution according to a study done with mice, chickens, monkeys and apes concluded that more evolved species tend to preserve the structures responsible for basic behaviors. A long term human study comparing the human brain to the primitive brain found that the modern human brain contains the primitive hindbrain region – what most neuroscientists call the <a href="/wiki/Triune_brain" title="Triune brain">protoreptilian brain</a>. The purpose of this part of the brain is to sustain fundamental homeostatic functions, which are self regulating processes organisms use to help their bodies adapt. The <a href="/wiki/Pons" title="Pons">pons</a> and <a href="/wiki/Medulla_oblongata" title="Medulla oblongata">medulla</a> are major structures found there. A new region of the brain developed in mammals about 250 million years after the appearance of the hindbrain. This region is known as the paleomammalian brain, the major parts of which are the <a href="/wiki/Hippocampus" title="Hippocampus">hippocampi</a> and <a href="/wiki/Amygdala" title="Amygdala">amygdalas</a>, often referred to as the <a href="/wiki/Limbic_system" title="Limbic system">limbic system</a>. The limbic system deals with more complex functions including emotional, sexual and fighting behaviors. Of course, animals that are not <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> also have brains, and their brains have undergone separate evolutionary histories.<sup id="cite_ref-Park_2018_6-1" class="reference"><a href="#cite_note-Park_2018-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The brainstem and limbic system are largely based on <a href="/wiki/Nucleus_(neuroanatomy)" title="Nucleus (neuroanatomy)">nuclei</a>, which are essentially balled-up clusters of tightly packed neurons and the <a href="/wiki/Axon" title="Axon">axon</a> fibers that connect them to each other, as well as to neurons in other locations. The other two major brain areas (the <a href="/wiki/Cerebrum" title="Cerebrum">cerebrum</a> and <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>) are based on a <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cortical</a> architecture. At the outer periphery of the cortex, the neurons are arranged into layers (the number of which vary according to species and function) a few millimeters thick. There are axons that travel between the layers, but the majority of axon mass is <i>below</i> the neurons themselves. Since cortical neurons and most of their axon fiber tracts do not have to compete for space, cortical structures can scale more easily than nuclear ones. A key feature of cortex is that because it scales with surface area, more of it can be fit inside a skull by introducing convolutions, in much the same way that a dinner napkin can be stuffed into a glass by wadding it up. The degree of convolution is generally greater in species with more complex behavior, which benefits from the increased surface area. </p><p>The <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>, or "little brain," is behind the brainstem and below the occipital lobe of the <a href="/wiki/Cerebrum" title="Cerebrum">cerebrum</a> in humans. Its purposes include the coordination of fine sensorimotor tasks, and it may be involved in some cognitive functions, such as language and different motor skills that may involve hands and feet. The cerebellum helps keep equilibrium. Damage to the cerebellum would result in all physical roles in life to be affected. Human cerebellar cortex is finely convoluted, much more so than cerebral cortex. Its interior axon fiber tracts are called the <i>arbor vitae</i>, or <a href="/wiki/Arbor_vitae_(anatomy)" title="Arbor vitae (anatomy)">Tree of Life</a>. </p><p>The area of the brain with the greatest amount of recent evolutionary change is called the <a href="/wiki/Neocortex" title="Neocortex">neocortex</a>. In reptiles and fish, this area is called the <a href="/wiki/Pallium_(neuroanatomy)" title="Pallium (neuroanatomy)">pallium</a> and is smaller and simpler relative to body mass than what is found in mammals. According to research, the cerebrum first developed about 200 million years ago. It is responsible for higher cognitive functions—for example, language, thinking, and related forms of information processing.<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> It is also responsible for processing sensory input (together with the <a href="/wiki/Thalamus" title="Thalamus">thalamus</a>, a part of the limbic system that acts as an information router). The thalamus receives the different sensations before the information is then passed onto the cerebral cortex. Most of its function is <a href="/wiki/Subconscious" title="Subconscious">subconscious</a>, that is, not available for inspection or intervention by the conscious mind. The neocortex is an elaboration, or outgrowth, of structures in the limbic system, with which it is tightly integrated. The neocortex is the main part controlling many brain functions as it covers half of the whole brain in volume. The development of these recent evolutionary changes in the neocortex likely occurred as a result of new neural network formations and positive selections of certain genetic components. </p> <div class="mw-heading mw-heading2"><h2 id="Role_of_embryology">Role of embryology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=2" title="Edit section: Role of embryology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Embryology" title="Embryology">Embryology</a> and <a href="/wiki/Brain_development_timelines" title="Brain development timelines">Brain development timelines</a></div> <p>In addition to studying the <a href="/wiki/Fossil" title="Fossil">fossil record</a>, evolutionary history can be investigated via embryology. An <a href="/wiki/Embryo" title="Embryo">embryo</a> is an unborn/unhatched animal and evolutionary history can be studied by observing how processes in embryonic development are conserved (or not conserved) across species. Similarities between different species may indicate evolutionary connection. One way anthropologists study evolutionary connection between species is by observing orthologs. An ortholog is defined as two or more homologous genes between species that are evolutionarily related by linear descent. By using embryology the evolution of the brain can be tracked between various species. </p><p><a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">Bone morphogenetic protein</a> (BMP), a <a href="/wiki/Growth_factor" title="Growth factor">growth factor</a> that plays a significant role in embryonic neural development, is highly conserved amongst vertebrates, as is <a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">sonic hedgehog</a> (SHH), a morphogen that inhibits BMP to allow neural crest development. Tracking these growth factors with the use of embryology provides a deeper understanding of what areas of the brain diverged in their evolution. Varying levels of these growth factors lead to differing embryonic neural development which then in turn affects the complexity of future neural systems. Studying the brain's development at various embryonic stages across differing species provides additional insight into what evolutionary changes may have historically occurred. This then allows scientists to look into what factors may have caused such changes, such as links to neural network diversity, growth factor production, protein- coding selections, and other genetic factors. </p> <div class="mw-heading mw-heading2"><h2 id="Randomizing_access_and_increasing_size">Randomizing access and increasing size</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=3" title="Edit section: Randomizing access and increasing size"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some animal phyla have gone through major brain enlargement through evolution (e.g. <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a> and <a href="/wiki/Cephalopod" title="Cephalopod">cephalopods</a> both contain many lineages in which brains have grown through evolution) but most animal groups are composed only of species with extremely small brains. Some scientists<sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Manual_of_Style/Words_to_watch#Unsupported_attributions" title="Wikipedia:Manual of Style/Words to watch"><span title="The material near this tag possibly uses too-vague attribution or weasel words. (June 2019)">who?</span></a></i>]</sup> argue that this difference is due to vertebrate and cephalopod neurons having evolved ways of communicating that overcome the scalability problem of <a href="/wiki/Neural_network" title="Neural network">neural networks</a> while most animal groups have not. They argue that the reason why traditional neural networks fail to improve their function when they scale up is because filtering based on previously known probabilities cause <a href="/wiki/Self-fulfilling_prophecy" title="Self-fulfilling prophecy">self-fulfilling prophecy</a>-like biases that create false statistical evidence giving a completely false worldview and that randomized access can overcome this problem and allow brains to be scaled up to more discriminating <a href="/wiki/Conditioned_reflex" class="mw-redirect" title="Conditioned reflex">conditioned reflexes</a> at larger brains that lead to <a href="/wiki/Emergence" title="Emergence">new worldview forming abilities at certain thresholds</a>. This means when neurons scale in a non randomized fashion that their functionality becomes more limited due to their neural networks being unable to process more complex systems without the exposure to new formations. This is explained by randomization allowing the entire brain to eventually get access to all information over the course of many shifts even though instant <a href="/wiki/Privileged_access" class="mw-redirect" title="Privileged access">privileged access</a> is physically impossible. They cite that vertebrate neurons transmit virus-like capsules containing <a href="/wiki/RNA" title="RNA">RNA</a> that are sometimes read in the neuron to which it is transmitted and sometimes passed further on unread which creates randomized access, and that cephalopod neurons make different proteins from the same gene which suggests another mechanism for randomization of concentrated information in neurons, both making it evolutionarily worth scaling up brains.<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><sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Brain_re-organization">Brain re-organization</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=4" title="Edit section: Brain re-organization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>With the use of in vivo Magnetic resonance imaging (<a href="/wiki/MRI" class="mw-redirect" title="MRI">MRI</a>) and tissue sampling, different cortical samples from members of each hominoid species were analyzed. In each species, specific areas were either relatively enlarged or shrunken, which can detail neural organizations. Different sizes in the cortical areas can show specific adaptations, functional specializations and evolutionary events that were changes in how the hominoid brain is organized. In early prediction it was thought that the frontal lobe, a large part of the brain that is generally devoted to behavior and social interaction, predicted the differences in behavior between hominoid and humans. Discrediting this theory was evidence supporting that damage to the frontal lobe in both humans and hominoids show atypical social and emotional behavior; thus, this similarity means that the frontal lobe was not very likely to be selected for reorganization. Instead, it is now believed that evolution occurred in other parts of the brain that are strictly associated with certain behaviors. The reorganization that took place is thought to have been more organizational than volumetric; whereas the brain volumes were relatively the same but specific landmark position of surface anatomical features, for example, the lunate sulcus suggest that the brains had been through a neurological reorganization.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> There is also evidence that the early hominin lineage also underwent a quiescent period, or a period of dormancy, which supports the idea of neural reorganization. </p><p>Dental fossil records for early humans and hominins show that immature hominins, including australopithecines and members of <i>Homo</i>, have a quiescent period (Bown et al. 1987). A quiescent period is a period in which there are no dental eruptions of adult teeth; at this time the child becomes more accustomed to social structure, and development of culture. During this time the child is given an extra advantage over other hominoids, devoting several years into developing speech and learning to cooperate within a community.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> This period is also discussed in relation to encephalization. It was discovered that chimpanzees do not have this neutral dental period, which suggests that a quiescent period occurred in very early hominin evolution. Using the models for neurological reorganization it can be suggested the cause for this period, dubbed middle childhood, is most likely for enhanced foraging abilities in varying seasonal environments. </p> <div class="mw-heading mw-heading2"><h2 id="Genetic_factors_in_recent_evolution">Genetic factors in recent evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=5" title="Edit section: Genetic factors in recent evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Genes involved in the neuro-development and in neuron physiology are extremely conserved between mammalian species (94% of genes expressed in common between humans and chimpanzees, 75% between humans and mice), compared to other organs. Therefore, few genes account for species differences in the human brain development and function.<sup id="cite_ref-Libé-Philippot_2021_15-0" class="reference"><a href="#cite_note-Libé-Philippot_2021-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Development_of_the_human_cerebral_cortex">Development of the human cerebral cortex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=6" title="Edit section: Development of the human cerebral cortex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Main differences rely on the evolution of <a href="/wiki/Non-coding_DNA" title="Non-coding DNA">non-coding genomic regions</a>, involved in the regulation of gene expression. This leads to differential expression of genes during the development of the human brain compared to other species, including chimpanzees. Some of these regions evolved fast in the human genome (<a href="/wiki/Human_accelerated_regions" title="Human accelerated regions">human accelerated regions</a>). The new genes expressed during human <a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a> are notably associated with the <a href="/wiki/Notch_signaling_pathway" title="Notch signaling pathway">NOTCH</a>, <a href="/wiki/Wnt_signaling_pathway" title="Wnt signaling pathway">WNT</a> and <a href="/wiki/MTOR" title="MTOR">mTOR</a> pathways, but are also involved <a href="/wiki/ZEB2" title="ZEB2">ZEB2</a>, <a href="/wiki/PDGFD" title="PDGFD">PDGFD</a> and its receptor <a href="/wiki/PDGFRB" title="PDGFRB">PDGFRβ</a>. The human cerebral cortex is also characterized by a higher gradient of <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a> in the <a href="/wiki/Prefrontal_cortex" title="Prefrontal cortex">prefrontal cortex</a>, leading to higher prefrontal cortex volume. All these differential gene expression lead to higher proliferation of the <a href="/wiki/Progenitor_cell" title="Progenitor cell">neural progenitors</a> leading to more neurons in the human cerebral cortex. Some genes are lost in their expression during the development of the human cerebral cortex like <a href="/wiki/GADD45G" title="GADD45G">GADD45G</a> and <a href="/wiki/FLRT2" title="FLRT2">FLRT2</a>/<a href="/wiki/FLRT3" title="FLRT3">FLRT3</a>.<sup id="cite_ref-Libé-Philippot_2021_15-1" class="reference"><a href="#cite_note-Libé-Philippot_2021-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Another source of molecular novelty rely on new genes in the human or <a href="/wiki/Hominidae" title="Hominidae">hominid</a> genomes through segmental duplication. Around 30 new genes in the hominid genomes are dynamically expressed during human corticogenesis. Some were linked to higher proliferation of neural progenitors: <a href="/wiki/NOTCH2NL" title="NOTCH2NL"><i>NOTCH2NLA</i>/B/C</a>, <i><a href="/wiki/ARHGAP11B" title="ARHGAP11B">ARHGAP11B</a>, <a href="/w/index.php?title=CROCCP2&action=edit&redlink=1" class="new" title="CROCCP2 (page does not exist)">CROCCP2</a>, <a href="/wiki/TBC1D3" title="TBC1D3">TBC1D3</a>, TMEM14B.</i> Patients with deletions with <i>NOTCH2NL</i> genes display <a href="/wiki/Microcephaly" title="Microcephaly">microcephaly</a>, showing the necessity of such duplicated genes, acquired in the human genomes, in the proper <a href="/wiki/Corticogenesis" class="mw-redirect" title="Corticogenesis">corticogenesis</a>.<sup id="cite_ref-Libé-Philippot_2021_15-2" class="reference"><a href="#cite_note-Libé-Philippot_2021-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="MCPH1_and_ASPM">MCPH1 and ASPM</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=7" title="Edit section: MCPH1 and ASPM"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1251242444"><table class="box-Very_long_section plainlinks metadata ambox ambox-style ambox-very_long" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">This section <b>may be <a href="/wiki/Wikipedia:Article_size" title="Wikipedia:Article size">too long</a> to read and navigate comfortably</b>.<span class="hide-when-compact"> Consider <a href="/wiki/Wikipedia:Splitting" title="Wikipedia:Splitting">splitting</a> content into sub-articles, <a href="/wiki/Wikipedia:Summary_style" title="Wikipedia:Summary style">condensing</a> it, or adding <a href="/wiki/Help:Section#Subsections" title="Help:Section">subheadings</a>. Please discuss this issue on the article's <a href="/wiki/Talk:Evolution_of_the_brain" title="Talk:Evolution of the brain">talk page</a>.</span> <span class="date-container"><i>(<span class="date">November 2022</span>)</i></span></div></td></tr></tbody></table> <p>Bruce Lahn, the senior author at the Howard Hughes Medical Center at the University of Chicago and colleagues have suggested that there are specific genes that control the size of the human brain. These genes continue to play a role in brain evolution, implying that the brain is continuing to evolve. The study began with the researchers assessing 214 genes that are involved in brain development. These genes were obtained from humans, macaques, rats and mice. Lahn and the other researchers noted points in the DNA sequences that caused protein alterations. These DNA changes were then scaled to the evolutionary time that it took for those changes to occur. The data showed the genes in the human brain evolved much faster than those of the other species. Once this genomic evidence was acquired, Lahn and his team decided to find the specific gene or genes that allowed for or even controlled this rapid evolution. Two genes were found to control the size of the human brain as it develops. These genes are <a href="/wiki/Microcephalin" title="Microcephalin">Microcephalin (MCPH1)</a> and <a href="/wiki/ASPM_(gene)" title="ASPM (gene)">Abnormal Spindle-like Microcephaly (ASPM)</a>. The researchers at the University of Chicago were able to determine that under the pressures of selection, both of these genes showed significant DNA sequence changes. Lahn's earlier studies displayed that Microcephalin experienced rapid evolution along the primate lineage which eventually led to the emergence of <i>Homo sapiens</i>. After the emergence of humans, Microcephalin seems to have shown a slower evolution rate. On the contrary, ASPM showed its most rapid evolution in the later years of human evolution once the divergence between chimpanzees and humans had already occurred.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Each of the gene sequences went through specific changes that led to the evolution of humans from ancestral relatives. In order to determine these alterations, Lahn and his colleagues used DNA sequences from multiple primates then compared and contrasted the sequences with those of humans. Following this step, the researchers statistically analyzed the key differences between the primate and human DNA to come to the conclusion, that the differences were due to natural selection. The changes in DNA sequences of these genes accumulated to bring about a competitive advantage and higher fitness that humans possess in relation to other primates. This comparative advantage is coupled with a larger brain size which ultimately allows the human mind to have a higher cognitive awareness.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Protein_ZEB2_PDB_2da7.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Protein_ZEB2_PDB_2da7.png/135px-Protein_ZEB2_PDB_2da7.png" decoding="async" width="135" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Protein_ZEB2_PDB_2da7.png/202px-Protein_ZEB2_PDB_2da7.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/82/Protein_ZEB2_PDB_2da7.png/270px-Protein_ZEB2_PDB_2da7.png 2x" data-file-width="485" data-file-height="590" /></a><figcaption>ZEB2 protein</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="ZEB2">ZEB2</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=8" title="Edit section: ZEB2"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>ZEB2 is a protein- coding gene in the Homo sapien species. A 2021 study found that a delayed change in the shape of <a href="/wiki/Neuroepithelial_cell" title="Neuroepithelial cell">early brain cells</a> causes the distinctly large human forebrain compared to other apes and identify <a href="/wiki/ZEB2" title="ZEB2">ZEB2</a> as a genetic regulator of it, whose manipulation lead to acquisition of nonhuman ape cortical architecture in <a href="/wiki/Brain_organoid" class="mw-redirect" title="Brain organoid">brain organoids</a>.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10.1016/j.cell.2021.02.050_19-0" class="reference"><a href="#cite_note-10.1016/j.cell.2021.02.050-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="NOVA1">NOVA1</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=9" title="Edit section: NOVA1"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2021, researchers reported that <a href="/wiki/Brain_organoid" class="mw-redirect" title="Brain organoid">brain organoids</a> created with stem cells into which they reintroduced the archaic gene variant <i><a href="/wiki/NOVA1" title="NOVA1">NOVA1</a></i> present in <a href="/wiki/Neanderthals" class="mw-redirect" title="Neanderthals">Neanderthals</a> and <a href="/wiki/Denisovans" class="mw-redirect" title="Denisovans">Denisovans</a> via <a href="/wiki/CRISPR_gene_editing" title="CRISPR gene editing">CRISPR-Cas9</a> shows that it has a major impact on <a href="/wiki/Neurodevelopment" class="mw-redirect" title="Neurodevelopment">neurodevelopment</a> and that such genetic mutations during the evolution of the human brain underlie traits that <a href="/wiki/Human#Genetics" title="Human">separate modern humans</a> from extinct <i>Homo</i> species. They found that expression of the archaic NOVA1 in cortical organoids leads to "modified synaptic protein interactions, affects <a href="/wiki/Glutamate_receptor" title="Glutamate receptor">glutamatergic signaling</a>, underlies differences in neuronal connectivity, and promotes higher heterogeneity of neurons regarding their <a href="/wiki/Electrophysiology" title="Electrophysiology">electrophysiological</a> profiles".<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> This research suggests positive selection of the modern NOVA1 gene, which may have promoted the randomization of neural scaling. A subsequent study failed to replicate the differences in organoid morphology between the modern human and the archaic NOVA1 variant,<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> consistent with suspected unwanted side effects of CRISPR editing in the original study.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="SRGAP2C_and_neuronal_maturation">SRGAP2C and neuronal maturation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=10" title="Edit section: SRGAP2C and neuronal maturation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Less is known about <a href="/wiki/Neuronal_maturation" class="mw-redirect" title="Neuronal maturation">neuronal maturation</a>. Synaptic gene and protein expression are protracted, in line with the <a href="/wiki/Neoteny_in_humans" title="Neoteny in humans">protracted</a> synaptic maturation of human cortical neurons so called neoteny. This probably relies on the evolution of non-coding genomic regions. The consequence of the neoteny could be an extension of the period of <a href="/wiki/Synaptic_plasticity" title="Synaptic plasticity">synaptic plasticity</a> and therefore of learning. A human-specific duplicated gene, <i><a href="/wiki/SRGAP2C" title="SRGAP2C">SRGAP2C</a></i> accounts for this synaptic neoteny and acts by regulating molecular pathways linked to neurodevelopmental disorders.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> Other genes are deferentially expressed in human neurons during their development such as <i><a href="/w/index.php?title=Osteocrin&action=edit&redlink=1" class="new" title="Osteocrin (page does not exist)">osteocrin</a></i> or <i><a href="/w/index.php?title=Cerebelin-2&action=edit&redlink=1" class="new" title="Cerebelin-2 (page does not exist)">cerebelin-2</a> </i>.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="LRRC37B_and_neuronal_electrical_properties">LRRC37B and neuronal electrical properties</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=11" title="Edit section: LRRC37B and neuronal electrical properties"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Even less is known about molecular specificities linked to the physiology of the human neurons. Human neurons are more divergent in the genes they express compared to chimpanzees than chimpanzees to gorilla, which suggests an acceleration of non-coding genomic regions associated with genes involved in neuronal physiology, in particular linked to the synapses.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> A hominid-specific duplicated gene, <i><a href="/wiki/LRRC37A" title="LRRC37A">LRRC37B</a></i>, codes for a <a href="/wiki/Transmembrane_receptor" class="mw-redirect" title="Transmembrane receptor">transmembrane receptor</a> that is selectively localized at the <a href="/wiki/Axon" title="Axon">axon initial segment</a> of <a href="/wiki/Pyramidal_neurons" class="mw-redirect" title="Pyramidal neurons">human cortical pyramidal neurons</a>.<sup id="cite_ref-doi.org_28-0" class="reference"><a href="#cite_note-doi.org-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> It inhibits their <a href="/wiki/Voltage-gated_sodium_channels" class="mw-redirect" title="Voltage-gated sodium channels">voltage-gated sodium channels</a> that generate the <a href="/wiki/Action_potential" title="Action potential">action potentials</a> leading to a lower neuronal excitability. Human cortical pyramidal neurons display a lower <a href="/wiki/Neuron" title="Neuron">excitability</a> compared to other mammalian species (including <a href="/wiki/Macaque" title="Macaque">macaques</a> and <a href="/wiki/Marmoset" title="Marmoset">marmosets</a>) which could lead to different circuit functions in the human species.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> Therefore, <i>LRRC37B</i> whose expression has been acquired in the human lineage after the separation from the chimpanzees could be a key gene in the function of the human cerebral cortex. LRRC37B binds to secreted <a href="/wiki/FGF13" title="FGF13">FGF13A</a> and <a href="/wiki/SCN1B" title="SCN1B">SCN1B</a> and modulate indirectly the activity of <a href="/wiki/SCN8A" title="SCN8A">SCN8A</a>,<sup id="cite_ref-doi.org_28-1" class="reference"><a href="#cite_note-doi.org-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> all involved in neural disorders such as epilepsy and autism. Therefore, LRRC37B may contribute to human-specific sensitivities to such disorders, both involved defects in neuronal excitability. </p> <div class="mw-heading mw-heading3"><h3 id="Genome_repair">Genome repair</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=12" title="Edit section: Genome repair"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Genome" title="Genome">genomic</a> <a href="/wiki/DNA" title="DNA">DNA</a> of postmitotic neurons ordinarily does not <a href="/wiki/DNA_replication" title="DNA replication">replicate</a>. Protection strategies have evolved to ensure the distinctive longevity of the neuronal genome. Human neurons are reliant on <a href="/wiki/DNA_repair" title="DNA repair">DNA repair</a> processes to maintain function during an individual's life-time. DNA repair tends to occur preferentially at evolutionarily conserved sites that are specifically involved with the <a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">regulation of expression of genes</a> essential for neuronal identity and function.<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Other_factors">Other factors</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=13" title="Edit section: Other factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many other genetics may also be involved in recent evolution of the brain. </p> <ul><li>For instance, scientists showed experimentally, with <a href="/wiki/Cerebral_organoids" class="mw-redirect" title="Cerebral organoids">brain organoids</a> grown from stem cells, how differences between humans and chimpanzees are also substantially caused by <a href="/wiki/Non-coding_DNA" title="Non-coding DNA">non-coding DNA</a> (often discarded as relatively meaningless "junk DNA") – in particular via <a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">CRE</a>-regulated <a href="/wiki/Gene_expression" title="Gene expression">expression</a> of the <i>ZNF558</i> gene for a transcription factor that regulates the <i><b><a rel="nofollow" class="external text" href="https://www.genenames.org/tools/search/#!/genes?query=SPATA18">SPATA18</a></b></i> gene.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> SPATA18 gene encodes a protein and is able to influence lysosome-like organelles that are found within mitochondria that eradicate oxidized mitochondrial proteins. This helps monitor the quality of the mitochondria as the disregulation of its quality control has been linked to cancer and degenerative diseases.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> This example may contribute to illustrations of the complexity and scope of relatively recent evolution to <i>Homo sapiens</i>.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup></li> <li>A change in gene <a href="/wiki/TKTL1" title="TKTL1">TKTL1</a> could be a key factor of recent brain evolution and difference of modern humans to (other) apes and Neanderthals, related to <a href="/wiki/Neocortex" title="Neocortex">neocortex</a>-<a href="/wiki/Neurogenesis" title="Neurogenesis">neurogenesis</a>.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> However, the "archaic" <a href="/wiki/Allele" title="Allele">allele</a> attributed to Neanderthals is present in 0.03% of <i>Homo sapiens</i>, but no resultant <a href="/wiki/Phenotypic_trait" title="Phenotypic trait">phenotypic</a> differences have been reported in these people.<sup id="cite_ref-Herai_2023_37-0" class="reference"><a href="#cite_note-Herai_2023-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Additionally, as Herai et al. contend, more is not always better. In fact, enhanced neuron production "can lead to an abnormally enlarged cortex and layer-specific imbalances in <a href="/wiki/Glia" title="Glia">glia</a>/neuron ratios and neuronal subpopulations during neurodevelopment."<sup id="cite_ref-Herai_2023_37-1" class="reference"><a href="#cite_note-Herai_2023-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Even the original study's authors agree that “any attempt to discuss prefrontal cortex and cognitive advantage of modern humans over Neandertals based on TKTL1 alone is problematic”.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup></li> <li>Some of the prior study's authors reported a similar <a href="/wiki/ARHGAP11B" title="ARHGAP11B">ARHGAP11B</a> mutation in 2016.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup></li> <li><a href="/wiki/Epigenetic" class="mw-redirect" title="Epigenetic">Epigenetics</a> also play a major role in the brain evolution in and to humans.<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag needs further explanation. (November 2022)">further explanation needed</span></a></i>]</sup></li></ul> <div class="mw-heading mw-heading3"><h3 id="Recently_evolved_traits">Recently evolved traits</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=14" title="Edit section: Recently evolved traits"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <dl><dt>Language</dt></dl> <p>A <a href="/wiki/Genome-wide_association_study" title="Genome-wide association study">genome-wide association study</a> <a href="/wiki/Meta-analysis" title="Meta-analysis">meta-analysis</a> reported genetic factors of, the <a href="/wiki/Origin_of_language" title="Origin of language">so far</a> uniquely human, <a href="/wiki/Neurogenetics" title="Neurogenetics">language-related</a> capacities, in particular factors of differences in <a href="/wiki/Skill" title="Skill">skill</a>-levels of five tested traits. It e.g. identified association with <a href="/wiki/Neuroanatomy" title="Neuroanatomy">neuroanatomy</a> of a language-related brain area via <a href="/wiki/Neuroimaging" title="Neuroimaging">neuroimaging</a> correlation. The data contributes to identifying or understanding the biological basis of this recently evolved characteristic capability.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Human_brain_evolution">Human brain evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=15" title="Edit section: Human brain evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Cranial_capacity" class="mw-redirect" title="Cranial capacity">Cranial capacity</a> and <a href="/wiki/Brain-to-body_mass_ratio" class="mw-redirect" title="Brain-to-body mass ratio">Brain-to-body mass ratio</a></div> <p>One of the prominent ways of tracking the evolution of the human brain is through direct evidence in the form of fossils. The evolutionary history of the human brain shows primarily a gradually bigger brain relative to body size during the evolutionary path from early primates to hominids and finally to <i>Homo sapiens</i>. Because fossilized brain tissue is rare, a more reliable approach is to observe anatomical characteristics of the skull that offer insight into brain characteristics. One such method is to observe the endocranial cast (also referred to as <a href="/wiki/Endocast" title="Endocast">endocasts</a>). Endocasts occur when, during the fossilization process, the brain deteriorates away, leaving a space that is filled by surrounding sedimentary material over time. These casts, give an imprint of the lining of the brain cavity, which allows a visualization of what was there.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> This approach, however, is limited in regard to what information can be gathered. Information gleaned from endocasts is primarily limited to the size of the brain (<a href="/wiki/Brain_size" title="Brain size">cranial capacity</a> or <a href="/wiki/Brain_size" title="Brain size">endocranial volume</a>), prominent <a href="/wiki/Sulcus_(neuroanatomy)" title="Sulcus (neuroanatomy)">sulci</a> and <a href="/wiki/Gyrus" title="Gyrus">gyri</a>, and size of dominant lobes or regions of the brain.<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Du_2018_49-0" class="reference"><a href="#cite_note-Du_2018-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> While endocasts are extremely helpful in revealing superficial brain anatomy, they cannot reveal brain structure, particularly of deeper brain areas. By determining scaling metrics of cranial capacity as it relates to total number of neurons present in primates, it is also possible to estimate the number of neurons through fossil evidence.<sup id="cite_ref-Herculano-Houzel_2012_50-0" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:H_georgicus_5_BLACK_PRINT.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/H_georgicus_5_BLACK_PRINT.jpg/220px-H_georgicus_5_BLACK_PRINT.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/H_georgicus_5_BLACK_PRINT.jpg/330px-H_georgicus_5_BLACK_PRINT.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/aa/H_georgicus_5_BLACK_PRINT.jpg/440px-H_georgicus_5_BLACK_PRINT.jpg 2x" data-file-width="2632" data-file-height="2632" /></a><figcaption>Facial reconstruction of a <i><a href="/wiki/Homo_georgicus" class="mw-redirect" title="Homo georgicus">Homo georgicus</a></i> from over 1.5 Mya</figcaption></figure> <p>Despite the limitations to endocasts, they can and do provide a basis for understanding human brain evolution, which shows primarily a gradually bigger brain. The evolutionary history of the human brain shows primarily a gradually bigger brain relative to body size during the evolutionary path from early primates to <a href="/wiki/Hominini" title="Hominini">hominins</a> and finally to <i>Homo sapiens</i>. This trend that has led to the present day human brain size indicates that there has been a 2-3 factor increase in size over the past 3 million years.<sup id="cite_ref-Du_2018_49-1" class="reference"><a href="#cite_note-Du_2018-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> This can be visualized with current data on hominin evolution, starting with <i><a href="/wiki/Australopithecus" title="Australopithecus">Australopithecus</a></i>, a group of hominins from which humans are likely descended.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> After all of the data, all observations concluded that the main development that occurred during evolution was the increase of brain size.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p><p>However, recent research has called into question the hypothesis of a threefold increase in brain size when comparing <i>Homo sapiens</i> with <i>Australopithecus</i> and chimpanzees. For example, in an article published in 2022 compiled a large data set of contemporary humans and found that the smallest human brains are less than twice that of large brained chimpanzees. As the authors write '...the upper limit of chimpanzee brain size is 500g/ml yet numerous modern humans have brain size below 900 g/ml.'<sup id="cite_ref-Clark_2022_53-0" class="reference"><a href="#cite_note-Clark_2022-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> (Note that in this quote, the unit g/ml is to be understood not in the usual way as gram per millilitre but rather as gram <i>or</i> millilitre. This is consistent because brain density is close to 1 g/ml.) Consequently, the authors argue that the notion of an increase in brain size being related to advances in cognition needs to be re-thought in light of global variation in brain size, as the brains of many modern humans with normal cognitive capacities are only 400g/ml larger than chimpanzees. Additionally, much of the increase in brain size - which occurs to a much greater degree in specific modern populations - can be explained by increases in correlated body size related to diet and climatic factors.<sup id="cite_ref-Clark_2022_53-1" class="reference"><a href="#cite_note-Clark_2022-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Australopithecus" title="Australopithecus">Australopiths</a> lived from 3.85 to 2.95 million years ago with the general cranial capacity somewhere near that of the extant chimpanzee—around 300–500 cm<sup>3</sup>.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> Considering that the volume of the modern human brain is around 1,352 cm<sup>3</sup> on average this represents a substantial amount of brain mass evolved.<sup id="cite_ref-56" class="reference"><a href="#cite_note-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> Australopiths are estimated to have a total neuron count of ~30-35 billion.<sup id="cite_ref-Herculano-Houzel_2012_50-1" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>Progressing along the human ancestral timeline, brain size continues to steadily increase (see <i><a href="/wiki/Homininae" title="Homininae">Homininae</a></i>) when moving into the era of <i><a href="/wiki/Homo" title="Homo">Homo</a></i>. For example, <i><a href="/wiki/Homo_habilis" title="Homo habilis">Homo habilis</a></i>, living 2.4 million to 1.4 million years ago and argued to be the first <i>Homo</i> species based on a host of characteristics, had a cranial capacity of around 600 cm<sup>3</sup>.<sup id="cite_ref-The_Smithsonian_Institution's_Human_Origins_Program_2010_57-0" class="reference"><a href="#cite_note-The_Smithsonian_Institution's_Human_Origins_Program_2010-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> <i>Homo habilis</i> is estimated to have had ~40 billion neurons.<sup id="cite_ref-Herculano-Houzel_2012_50-2" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>A little closer to present day, <i><a href="/wiki/Homo_heidelbergensis" title="Homo heidelbergensis">Homo heidelbergensis</a></i> lived from around 700,000 to 200,000 years ago and had a cranial capacity of around 1290 cm<sup>3</sup><sup id="cite_ref-The_Smithsonian_Institution's_Human_Origins_Program_2010_57-1" class="reference"><a href="#cite_note-The_Smithsonian_Institution's_Human_Origins_Program_2010-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> and having around 76 billion neurons.<sup id="cite_ref-Herculano-Houzel_2012_50-3" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p><i><a href="/wiki/Neanderthal" title="Neanderthal">Homo neaderthalensis</a></i>, living 400,000 to 40,000 years ago, had a cranial capacity comparable to that of modern humans at around 1500–1600 cm<sup>3</sup>on average, with some specimens of <a href="/wiki/Neanderthal" title="Neanderthal">Neanderthal</a> having even greater cranial capacity.<sup id="cite_ref-58" class="reference"><a href="#cite_note-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> Neanderthals are estimated to have had around 85 billion neurons.<sup id="cite_ref-Herculano-Houzel_2012_50-4" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> The increase in brain size topped with <a href="/wiki/Neanderthal" title="Neanderthal">Neanderthals</a>, possibly due to their larger visual systems.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p><p>It is also important to note that the measure of brain mass or volume, seen as cranial capacity, or even <a href="/wiki/Encephalization_quotient" title="Encephalization quotient">relative brain size</a>, which is brain mass that is expressed as a percentage of body mass, are not a measure of intelligence, use, or function of regions of the brain.<sup id="cite_ref-Herculano-Houzel_2012_50-5" class="reference"><a href="#cite_note-Herculano-Houzel_2012-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Total neurons, however, also do not indicate a higher ranking in cognitive abilities. Elephants have a higher number of total neurons (257 billion)<sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> compared to humans (100 billion).<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> Relative brain size, overall mass, and total number of neurons are only a few metrics that help scientists follow the evolutionary trend of increased brain to body ratio through the hominin phylogeny. </p><p>In 2021, scientists suggested that the brains of early <i>Homo</i> from Africa and <a href="/wiki/Dmanisi_historic_site" title="Dmanisi historic site">Dmanisi</a>, Georgia, Western Asia "retained a <a href="/wiki/Great_ape" class="mw-redirect" title="Great ape">great ape</a>-like structure of the <a href="/wiki/Frontal_lobe" title="Frontal lobe">frontal lobe</a>" for far longer than previously thought – until about 1.5 million years ago. Their findings imply that <i>Homo</i> <a href="/wiki/Early_human_migrations" title="Early human migrations">first dispersed out of Africa</a> before human brains evolved to roughly their modern anatomical structure in terms of the location and organization of individual brain regions. It also suggests that this evolution occurred – not during – but only long after the Homo lineage evolved ~2.5 million years ago and after they – <i><a href="/wiki/Homo_erectus" title="Homo erectus">Homo erectus</a></i> in particular – evolved to walk upright.<sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> What is the least controversial is that the brain expansion started about 2.6 Ma (about the same as the start of the <a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a>), and ended around 0.2 Ma. </p> <div class="mw-heading mw-heading3"><h3 id="Evolution_of_the_neocortex">Evolution of the neocortex</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=16" title="Edit section: Evolution of the neocortex"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Neocortex" title="Neocortex">Neocortex</a></div><p>In addition to just the size of the brain, scientists have observed changes in the folding of the brain, as well as in the thickness of the <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cortex</a>. The more convoluted the surface of the brain is, the greater the surface area of the cortex which allows for an expansion of cortex. It is the most evolutionarily advanced part of the brain.<sup id="cite_ref-Schnack_2015_67-0" class="reference"><a href="#cite_note-Schnack_2015-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> Greater surface area of the brain is linked to higher intelligence as is the thicker cortex but there is an inverse relationship—the thicker the cortex, the more difficult it is for it to fold. In adult humans, thicker cerebral cortex has been linked to higher intelligence.<sup id="cite_ref-Schnack_2015_67-1" class="reference"><a href="#cite_note-Schnack_2015-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p><p>The neocortex is the most advanced and most evolutionarily young part of the human brain. It is six layers thick and is only present in mammals. It is especially prominent in humans and is the location of most higher level functioning and cognitive ability.<sup id="cite_ref-Rakic_2009_68-0" class="reference"><a href="#cite_note-Rakic_2009-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> The six-layered neocortex found in mammals is evolutionarily derived from a three-layer cortex present in all modern reptiles.<sup id="cite_ref-www.mpg.de_69-0" class="reference"><a href="#cite_note-www.mpg.de-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> This three-layer cortex is still conserved in some parts of the human brain such as the hippocampus and is believed to have evolved in mammals to the neocortex during the transition between the Triassic and Jurassic periods.<sup id="cite_ref-www.mpg.de_69-1" class="reference"><a href="#cite_note-www.mpg.de-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Rakic_2009_68-1" class="reference"><a href="#cite_note-Rakic_2009-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> After looking at history, the mammals had little neocortex compared to the primates as they had more cortex.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> The three layers of this reptilian cortex correlate strongly to the first, fifth and sixth layers of the mammalian neocortex.<sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> Across species of mammals, primates have greater neuronal density compared to rodents of similar brain mass and this may account for increased intelligence.<sup id="cite_ref-Rakic_2009_68-2" class="reference"><a href="#cite_note-Rakic_2009-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Theories_of_human_brain_evolution">Theories of human brain evolution</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=17" title="Edit section: Theories of human brain evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Explanations of the rapid evolution and exceptional size of the human brain can be classified into five groups: instrumental, social, environmental, dietary, and anatomo-physiological. The instrumental hypotheses<sup id="cite_ref-Dunbar_2017_72-0" class="reference"><a href="#cite_note-Dunbar_2017-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> are based on the logic that <a href="/wiki/Natural_selection" title="Natural selection">evolutionary selection</a> for larger brains is beneficial for <a href="/wiki/Species" title="Species">species</a> survival, dominance, and spread, because larger brains facilitate food-finding and mating success. The social hypotheses<sup id="cite_ref-Dunbar_2017_72-1" class="reference"><a href="#cite_note-Dunbar_2017-72"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> suggest that social behavior stimulates evolutionary expansion of brain size.<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> Similarly, the environmental hypotheses suppose that <a href="/wiki/Encephalization_quotient" title="Encephalization quotient">encephalization</a> is promoted by environmental factors such as stress, variability, and consistency.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> The dietary theories maintain that food quality and certain nutritional components directly contributed to the brain growth in the <i><a href="/wiki/Homo" title="Homo">Homo</a></i> genus.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> The anatomo-physiologic concepts, such as <a href="/wiki/Human_head" title="Human head">cranio-cerebral</a> vascular <a href="/wiki/Hypertension" title="Hypertension">hypertension</a> due to head-down posture of the <a href="/wiki/Simian" title="Simian">anthropoid</a> fetus during pregnancy, are primarily focused on anatomic-functional changes that predispose to brain enlargement. </p><p>No single theory can completely account for human brain evolution. Multiple selective pressures in combination seems to have been involved.<sup id="cite_ref-76" class="reference"><a href="#cite_note-76"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> Synthetic theories have been proposed,<sup id="cite_ref-77" class="reference"><a href="#cite_note-77"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> but have not clearly explained reasons for the uniqueness of the human brain. Puzzlingly, brain enlargement has been found to have occurred independently in different primate lineages,<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> but only human lineage ended up with an exceptional brain capacity. <a href="/wiki/Medical_imaging_in_pregnancy" title="Medical imaging in pregnancy">Fetal head-down posture</a> may be an explanation of this conundrum <a rel="nofollow" class="external autonumber" href="https://ars.els-cdn.com/content/image/1-s2.0-S0006899323003293-ga1_lrg.jpg">[1]</a> because <i><a href="/wiki/Human" title="Human">Homo sapiens</a></i> is the only primate <a href="/wiki/Facultative_bipedalism" title="Facultative bipedalism">obligatory biped</a> with <a href="/wiki/Orthograde_posture" title="Orthograde posture">upright posture</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_brain&action=edit&section=18" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Bilateria" title="Bilateria">Bilateria</a></li> <li><a href="/wiki/Brain" title="Brain">Brain</a></li> <li><a href="/wiki/Evolution" title="Evolution">Evolution</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_brain&action=edit&section=19" 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 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Series B, Biological Sciences</i>. <b>370</b> (1663): 20140064. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.2014.0064">10.1098/rstb.2014.0064</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/PMC4305165">4305165</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/25602068">25602068</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+of+London.+Series+B%2C+Biological+Sciences&rft.atitle=A+synthesis+of+the+theories+and+concepts+of+early+human+evolution&rft.volume=370&rft.issue=1663&rft.pages=20140064&rft.date=2015-03&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4305165%23id-name%3DPMC&rft_id=info%3Apmid%2F25602068&rft_id=info%3Adoi%2F10.1098%2Frstb.2014.0064&rft.aulast=Maslin&rft.aufirst=MA&rft.au=Shultz%2C+S&rft.au=Trauth%2C+MH&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4305165&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></span> </li> <li id="cite_note-78"><span class="mw-cite-backlink"><b><a href="#cite_ref-78">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHalleyDeacon2017" class="citation cs2">Halley AC, Deacon TW (2017), <a rel="nofollow" class="external text" href="https://dx.doi.org/10.1016/b978-0-12-804042-3.00135-4">"The Developmental Basis of Evolutionary Trends in Primate Encephalization"</a>, <i>Evolution of Nervous Systems</i>, Elsevier, pp. <span class="nowrap">149–</span>162, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fb978-0-12-804042-3.00135-4">10.1016/b978-0-12-804042-3.00135-4</a>, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-804096-6" title="Special:BookSources/978-0-12-804096-6"><bdi>978-0-12-804096-6</bdi></a><span class="reference-accessdate">, retrieved <span class="nowrap">2024-03-05</span></span></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Evolution+of+Nervous+Systems&rft.atitle=The+Developmental+Basis+of+Evolutionary+Trends+in+Primate+Encephalization&rft.pages=%3Cspan+class%3D%22nowrap%22%3E149-%3C%2Fspan%3E162&rft.date=2017&rft_id=info%3Adoi%2F10.1016%2Fb978-0-12-804042-3.00135-4&rft.isbn=978-0-12-804096-6&rft.aulast=Halley&rft.aufirst=AC&rft.au=Deacon%2C+TW&rft_id=http%3A%2F%2Fdx.doi.org%2F10.1016%2Fb978-0-12-804042-3.00135-4&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></span> </li> </ol></div></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_brain&action=edit&section=20" 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="CITEREFFalk2011" class="citation book cs1">Falk D (2011). <i>The Fossil Chronicles: How Two Controversial Discoveries Changed Our View of Human Evolution</i>. University of California Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-26670-4" title="Special:BookSources/978-0-520-26670-4"><bdi>978-0-520-26670-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Fossil+Chronicles%3A+How+Two+Controversial+Discoveries+Changed+Our+View+of+Human+Evolution.&rft.pub=University+of+California+Press&rft.date=2011&rft.isbn=978-0-520-26670-4&rft.aulast=Falk&rft.aufirst=D&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRaichlenPolk2013" class="citation journal cs1">Raichlen DA, Polk JD (January 2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3574441">"Linking brains and brawn: exercise and the evolution of human neurobiology"</a>. <i>Proceedings. Biological Sciences</i>. <b>280</b> (1750): 20122250. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frspb.2012.2250">10.1098/rspb.2012.2250</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/PMC3574441">3574441</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/23173208">23173208</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings.+Biological+Sciences&rft.atitle=Linking+brains+and+brawn%3A+exercise+and+the+evolution+of+human+neurobiology&rft.volume=280&rft.issue=1750&rft.pages=20122250&rft.date=2013-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3574441%23id-name%3DPMC&rft_id=info%3Apmid%2F23173208&rft_id=info%3Adoi%2F10.1098%2Frspb.2012.2250&rft.aulast=Raichlen&rft.aufirst=DA&rft.au=Polk%2C+JD&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3574441&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStriedter2005" class="citation book cs1">Striedter GF (2005). <i>Principles of Brain Evolution</i>. Sinauer Associates.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Principles+of+Brain+Evolution.&rft.pub=Sinauer+Associates&rft.date=2005&rft.aulast=Striedter&rft.aufirst=GF&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEccles1989" class="citation book cs1">Eccles JC (1989). <i>Evolution of the Brain</i>. Routledge.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution+of+the+Brain&rft.pub=Routledge&rft.date=1989&rft.aulast=Eccles&rft.aufirst=JC&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+brain" class="Z3988"></span></li></ul> </div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul 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aria-labelledby="Evolutionary_biology753" 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"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology753" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a 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 href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">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 class="mw-selflink selflink">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 class="mw-selflink selflink">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 href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">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 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