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subsection</span> </button> <ul id="toc-Neural_control_of_muscle_force-sublist" class="vector-toc-list"> <li id="toc-Motor_units_and_force_production" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Motor_units_and_force_production"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Motor units and force production</span> </div> </a> <ul id="toc-Motor_units_and_force_production-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Recruitment_order" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Recruitment_order"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Recruitment order</span> </div> </a> <ul id="toc-Recruitment_order-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Computational_issues_of_motor_control" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Computational_issues_of_motor_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Computational issues of motor control</span> </div> </a> <ul id="toc-Computational_issues_of_motor_control-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Model_systems_for_motor_control" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Model_systems_for_motor_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Model systems for motor control</span> </div> </a> <ul id="toc-Model_systems_for_motor_control-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Sensorimotor_feedback" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Sensorimotor_feedback"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Sensorimotor feedback</span> </div> </a> <button aria-controls="toc-Sensorimotor_feedback-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 Sensorimotor feedback subsection</span> </button> <ul id="toc-Sensorimotor_feedback-sublist" class="vector-toc-list"> <li id="toc-Response_to_stimuli" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Response_to_stimuli"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Response to stimuli</span> </div> </a> <ul id="toc-Response_to_stimuli-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Closed_loop_control" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Closed_loop_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Closed loop control</span> </div> </a> <ul id="toc-Closed_loop_control-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Open_loop_control" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Open_loop_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Open loop control</span> </div> </a> <ul id="toc-Open_loop_control-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Coordination" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Coordination"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Coordination</span> </div> </a> <button aria-controls="toc-Coordination-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 Coordination subsection</span> </button> <ul id="toc-Coordination-sublist" class="vector-toc-list"> <li id="toc-Reflexes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Reflexes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.1</span> <span>Reflexes</span> </div> </a> <ul id="toc-Reflexes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Synergies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Synergies"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.2</span> <span>Synergies</span> </div> </a> <ul id="toc-Synergies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Motor_Programs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Motor_Programs"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.3</span> <span>Motor Programs</span> </div> </a> <ul id="toc-Motor_Programs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Redundancy" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Redundancy"> <div class="vector-toc-text"> <span class="vector-toc-numb">5.4</span> <span>Redundancy</span> </div> </a> <ul id="toc-Redundancy-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Perception_in_motor_control" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Perception_in_motor_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Perception in motor control</span> </div> </a> <button aria-controls="toc-Perception_in_motor_control-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 Perception in motor control subsection</span> </button> <ul id="toc-Perception_in_motor_control-sublist" class="vector-toc-list"> <li id="toc-Model_based_control_strategies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Model_based_control_strategies"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>Model based control strategies</span> </div> </a> <ul id="toc-Model_based_control_strategies-sublist" class="vector-toc-list"> <li id="toc-Inference_and_indirect_perception" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Inference_and_indirect_perception"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1.1</span> <span>Inference and indirect perception</span> </div> </a> <ul id="toc-Inference_and_indirect_perception-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Forward_models" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Forward_models"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1.2</span> <span>Forward models</span> </div> </a> <ul id="toc-Forward_models-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Inverse_models" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Inverse_models"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1.3</span> <span>Inverse models</span> </div> </a> <ul id="toc-Inverse_models-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Information_based_control" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Information_based_control"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Information based control</span> </div> </a> <ul id="toc-Information_based_control-sublist" class="vector-toc-list"> <li id="toc-Direct_perception" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Direct_perception"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2.1</span> <span>Direct perception</span> </div> </a> <ul id="toc-Direct_perception-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Behavioral_dynamics" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Behavioral_dynamics"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2.2</span> <span>Behavioral dynamics</span> </div> </a> <ul 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</div> </a> <ul id="toc-Individual_movement_optimization-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Multi-component_movements" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Multi-component_movements"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Multi-component movements</span> </div> </a> <ul id="toc-Multi-component_movements-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Further reading</span> </div> </a> <button aria-controls="toc-Further_reading-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Further reading subsection</span> </button> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> <li id="toc-Research_in_athletes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Research_in_athletes"> <div class="vector-toc-text"> <span class="vector-toc-numb">10.1</span> <span>Research in athletes</span> </div> </a> <ul id="toc-Research_in_athletes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </div> </div> </nav> 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href="https://de.wikipedia.org/wiki/Bewegungskontrolle" title="Bewegungskontrolle – German" lang="de" hreflang="de" data-title="Bewegungskontrolle" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%A9%D9%86%D8%AA%D8%B1%D9%84_%D9%85%D9%88%D8%AA%D9%88%D8%B1" title="کنترل موتور – Persian" lang="fa" hreflang="fa" data-title="کنترل موتور" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Contr%C3%B4le_moteur" title="Contrôle moteur – French" lang="fr" hreflang="fr" data-title="Contrôle moteur" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Fun%C3%A7%C3%A3o_motora" title="Função motora – Portuguese" lang="pt" hreflang="pt" data-title="Função motora" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Motoriset_toiminnot" title="Motoriset toiminnot – Finnish" lang="fi" hreflang="fi" data-title="Motoriset toiminnot" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Motor_kontrol" title="Motor kontrol – Turkish" lang="tr" hreflang="tr" data-title="Motor kontrol" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E9%81%8B%E5%8B%95%E6%8E%A7%E5%88%B6" title="運動控制 – Cantonese" lang="yue" hreflang="yue" data-title="運動控制" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target"><span>粵語</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E9%81%8B%E5%8B%95%E6%8E%A7%E5%88%B6_(%E7%94%9F%E7%89%A9%E9%AB%94)" title="運動控制 (生物體) – Chinese" lang="zh" hreflang="zh" data-title="運動控制 (生物體)" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target"><span>中文</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q2996165#sitelinks-wikipedia" title="Edit interlanguage links" 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dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Regulation of movement within organisms possessing a nervous system</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">This article is about motor control by humans and other animals. For motor control by machines and robots, see <a href="/wiki/Motor_controller" title="Motor controller">Motor controller</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Motor function" redirects here. Not to be confused with <a href="/wiki/Motor_Function" title="Motor Function">Motor Function</a>.</div> <p><b>Motor control</b> is the regulation of <a href="/wiki/Motility" title="Motility">movements</a> in <a href="/wiki/Organisms" class="mw-redirect" title="Organisms">organisms</a> that possess a <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a>. Motor control includes conscious <a href="/wiki/Motor_skill" title="Motor skill">voluntary movements</a>, subconscious <a href="/wiki/Muscle_memory" title="Muscle memory">muscle memory</a> and involuntary <a href="/wiki/Reflexes" class="mw-redirect" title="Reflexes">reflexes</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> as well as <a href="/wiki/Instinct" title="Instinct">instinctual</a> <a href="/wiki/Taxis" title="Taxis">taxis</a>. </p><p>To control movement, the nervous system must integrate multimodal <a href="/wiki/Sensory_information" class="mw-redirect" title="Sensory information">sensory information</a> (both from the external world as well as <a href="/wiki/Proprioception" title="Proprioception">proprioception</a>) and elicit the necessary signals to recruit <a href="/wiki/Muscles" class="mw-redirect" title="Muscles">muscles</a> to carry out a goal. This pathway spans many disciplines, including <a href="/wiki/Multisensory_integration" title="Multisensory integration">multisensory integration</a>, <a href="/wiki/Signal_processing" title="Signal processing">signal processing</a>, <a href="/wiki/Motor_coordination" title="Motor coordination">coordination</a>, <a href="/wiki/Biomechanics" title="Biomechanics">biomechanics</a>, and <a href="/wiki/Cognition" title="Cognition">cognition</a>,<sup id="cite_ref-Rosenbaum1991_2-0" class="reference"><a href="#cite_note-Rosenbaum1991-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Wise2002_3-0" class="reference"><a href="#cite_note-Wise2002-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> and the computational challenges are often discussed under the term sensorimotor control.<sup id="cite_ref-Franklin_425–442_4-0" class="reference"><a href="#cite_note-Franklin_425–442-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Successful motor control is crucial to interacting with the world to carry out goals as well as for posture, balance, and stability. </p><p>Some researchers (mostly neuroscientists studying movement, such as <a href="/wiki/Daniel_Wolpert" title="Daniel Wolpert">Daniel Wolpert</a> and <a href="/wiki/Randy_Flanagan" title="Randy Flanagan">Randy Flanagan</a>) argue that motor control is the reason brains exist at all.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Neural_control_of_muscle_force">Neural control of muscle force</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=1" title="Edit section: Neural control of muscle force"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>All movements, e.g. touching your nose, require <a href="/wiki/Motor_neuron" title="Motor neuron">motor neurons</a> to fire <a href="/wiki/Action_potential" title="Action potential">action potentials</a> that results in contraction of <a href="/wiki/Muscle" title="Muscle">muscles</a>. In humans, ~150,000 motor neurons control the contraction of ~600 muscles. To produce movements, a subset of 600 muscles must contract in a temporally precise pattern to produce the right force at the right time.<sup id="cite_ref-Kernell,_Daniel._2006_6-0" class="reference"><a href="#cite_note-Kernell,_Daniel._2006-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Motor_units_and_force_production">Motor units and force production</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=2" title="Edit section: Motor units and force production"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A single motor neuron and the <a href="/wiki/Myocyte" class="mw-redirect" title="Myocyte">muscle fibers</a> it innervates are called a <a href="/wiki/Motor_unit" title="Motor unit">motor unit</a>. For example, <a href="/wiki/Rectus_femoris_muscle" title="Rectus femoris muscle">the rectus femoris</a> contains approximately 1 million muscle fibers, which are controlled by around 1000 motor neurons. Activity in the motor neuron causes contraction in all of the innervated muscle fibers so that they function as a unit. Increasing action potential frequency (spike rate) in the motor neuron increases the muscle fiber contraction force, up to the maximal force.<sup id="cite_ref-Kernell,_Daniel._2006_6-1" class="reference"><a href="#cite_note-Kernell,_Daniel._2006-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><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> The maximal force depends on the contractile properties of the muscle fibers. Within a motor unit, all the muscle fibers are of the same type (e.g. <a href="/wiki/Myocyte" class="mw-redirect" title="Myocyte">type I (slow twitch) or Type II fibers (fast twitch)</a>), and motor units of multiple types make up a given muscle. Motor units of a given muscle are collectively referred to as a motor pool. </p><p>The force produced in a given muscle thus depends on: 1) How many motor neurons are active, and their spike rates; 2) the contractile properties and number of muscle fibers innervated by the active neurons. To generate more force, increase the spike rates of active motor neurons and/or recruiting more and stronger motor units. In turn, how the muscle force produces limb movement depends on the limb <a href="/wiki/Biomechanics" title="Biomechanics">biomechanics</a>, e.g. where the tendon and muscle originate (which bone, and precise location) and where the muscle inserts on the bone that it moves. </p> <div class="mw-heading mw-heading3"><h3 id="Recruitment_order">Recruitment order</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=3" title="Edit section: Recruitment order"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Motor units within a motor pool are <a href="/wiki/Motor_unit_recruitment" title="Motor unit recruitment">recruited in a stereotypical order</a>, from motor units that produce small amounts of force per spike, to those producing the largest force per spike. The gradient of motor unit force is correlated with a gradient in motor neuron soma size and motor neuron electrical excitability. This relationship was described by <a href="/wiki/Elwood_Henneman" title="Elwood Henneman">Elwood Henneman</a> and is known as <a href="/wiki/Henneman%27s_size_principle" title="Henneman's size principle">Henneman's size principle</a>, a fundamental discovery of neuroscience and an organizing principle of motor control.<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>For tasks requiring small forces, such as continual adjustment of posture, motor units with fewer muscle fibers that are slowly-contracting, but less fatigueable, are used. As more force is required, motor units with fast twitch, fast-fatigeable muscle fibers are recruited. </p> <pre> High| | _________________ Force required | / | | | | | _____________|_________________ | __________|_______________________________ Low|__________|__________________________________________ ↑ ↑ ↑ Time Type I Recruit first Type II A Type IIB </pre> <div class="mw-heading mw-heading2"><h2 id="Computational_issues_of_motor_control">Computational issues of motor control</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=4" title="Edit section: Computational issues of motor control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The nervous system produces movement by selecting which motor neurons are activated, and when. The finding that a recruitment order exists within a motor pool is thought to reflect a simplification of the problem: if a particular muscle should produce a particular force, then activate the motor pool along its recruitment hierarchy until that force is produced. </p><p>But then how to choose what force to produce in each muscle? The nervous system faces the following issues in solving this problem.<sup id="cite_ref-Franklin_425–442_4-1" class="reference"><a href="#cite_note-Franklin_425–442-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <ol><li>Redundancy. Infinite trajectories of movements can accomplish a goal (e.g. touch my nose). How is a trajectory chosen? Which trajectory is best?</li> <li>Noise. Noise is defined as small fluctuations that are unrelated to a signal, which can occur in neurons and synaptic connections at any point from sensation to muscle contraction.</li> <li>Delays. Motor neuron activity precedes muscle contraction, which precedes the movement. Sensory signals also reflect events that have already occurred. Such delays affect the choice of motor program.</li> <li>Uncertainty. Uncertainty arises because of neural noise, but also because inferences about the state of the world may not be correct (e.g. speed of on coming ball).</li> <li>Nonstationarity. Even as a movement is being executed, the state of the world changes, even through such simple effects as reactive forces on the rest of the body, <a href="/wiki/Neural_control_of_limb_stiffness" title="Neural control of limb stiffness">causing translation of a joint while it is actuated</a>.</li> <li>Nonlinearity. The effects of neural activity and muscle contraction are highly non-linear, which the nervous system must account for when predicting the consequences of a pattern of motor neuron activity.</li></ol> <p>Much ongoing research is dedicated to investigating how the nervous system deals with these issues, both at the behavioral level, as well as how neural circuits in the brain and spinal cord represent and deal with these factors to produce the fluid movements we witness in animals. </p><p>"Optimal feedback control" is an influential theoretical framing of these computation issues.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Model_systems_for_motor_control">Model systems for motor control</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=5" title="Edit section: Model systems for motor control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>All organisms face the computational challenges above, so neural circuits for motor control have been studied in <a href="/wiki/Kinesiology" title="Kinesiology">humans</a>, monkeys,<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> horses, cats,<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> mice,<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> fish<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> lamprey,<sup id="cite_ref-Grillner_1995_14-0" class="reference"><a href="#cite_note-Grillner_1995-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> flies,<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> locusts,<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> and nematodes,<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> among many others. Mammalian model systems like mice and monkeys offer the most straightforward comparative models for human health and disease. They are widely used to study the role of higher brain regions common to vertebrates, including the cerebral cortex, thalamus, basal ganglia and deep brain medullary and reticular circuits for motor control.<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> The <a href="/wiki/Hox_gene" title="Hox gene">genetics and neurophysiology of motor circuits</a> in the spine have also been studied in mammalian model organisms, but protective vertebrae make it difficult to study the functional role of spinal circuits in behaving animals. Here, larval and adult fish have been useful in discovering the functional logic of the local spinal circuits that coordinate motor neuron activity. Invertebrate model organisms do not have the same brain regions as vertebrates, but their brains must solve similar computational issues and thus are thought to have brain regions homologous to those involved in motor control in the vertebrate nervous system,<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> The organization of <a href="/wiki/Arthropod" title="Arthropod">arthropod</a> nervous systems into ganglia that control each leg as allowed researchers to record from neurons dedicated to moving a specific leg during behavior. </p><p>Model systems have also demonstrated the role of <a href="/wiki/Central_pattern_generator" title="Central pattern generator">central pattern generators</a> in driving rhythmic movements.<sup id="cite_ref-Grillner_1995_14-1" class="reference"><a href="#cite_note-Grillner_1995-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> A central pattern generator is a neural network that can generate rhythmic activity in the absence of an external control signal, such as a signal descending from the brain or feedback signals from sensors in the limbs (e.g. <a href="/wiki/Proprioception" title="Proprioception">proprioceptors</a>). Evidence suggests that real CPGs exist in several key motor control regions, such as the <a href="/wiki/Stomatogastric_ganglion" title="Stomatogastric ganglion">stomachs of arthropods</a> or the <a href="/wiki/Pre-B%C3%B6tzinger_complex" title="Pre-Bötzinger complex">pre-Boetzinger complex</a> that control breathing in humans. Furthermore, as a theoretical concept, CPGs have been useful to frame the possible role of sensory feedback in motor control. </p> <div class="mw-heading mw-heading2"><h2 id="Sensorimotor_feedback">Sensorimotor feedback</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=6" title="Edit section: Sensorimotor feedback"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Response_to_stimuli">Response to stimuli</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=7" title="Edit section: Response to stimuli"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The process of becoming aware of a sensory stimulus and using that information to influence an action occurs in stages. <a href="/wiki/Reaction_time" class="mw-redirect" title="Reaction time">Reaction time</a> of simple tasks can be used to reveal information about these stages. <i>Reaction time</i> refers to the period of time between when the stimulus is presented, and the end of the response. <i>Movement time</i> is the time it takes to complete the movement. Some of the first reaction time experiments were carried out by <a href="/wiki/Franciscus_Donders" title="Franciscus Donders">Franciscus Donders</a>, who used the difference in response times to a choice task to determine the length of time needed to process the stimuli and choose the correct response.<sup id="cite_ref-Donders_1969_20-0" class="reference"><a href="#cite_note-Donders_1969-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> While this approach is ultimately flawed, it gave rise to the idea that reaction time was made up of a stimulus identification, followed by a response selection, and ultimately culminates in carrying out the correct movement. Further research has provided evidence that these stages do exist, but that the response selection period of any reaction time increases as the number of available choices grows, a relationship known as <a href="/wiki/Hick%27s_law" title="Hick's law">Hick's law</a>.<sup id="cite_ref-Hick_1952_21-0" class="reference"><a href="#cite_note-Hick_1952-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Closed_loop_control">Closed loop control</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=8" title="Edit section: Closed loop control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The classical definition of a closed loop system for human movement comes from Jack A. Adams (1971).<sup id="cite_ref-Adams_1971_22-0" class="reference"><a href="#cite_note-Adams_1971-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Adams_1976_23-0" class="reference"><a href="#cite_note-Adams_1976-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> A reference of the desired output is compared to the actual output via error detection mechanisms; using feedback, the error is corrected for. Most movements that are carried out during day-to-day activity are formed using a continual process of accessing sensory information and using it to more accurately continue the motion. This type of motor control is called <a href="/wiki/Feedback_control" class="mw-redirect" title="Feedback control">feedback control</a>, as it relies on sensory feedback to control movements. Feedback control is a situated form of motor control, relying on sensory information about performance and specific sensory input from the environment in which the movement is carried out. This sensory input, while processed, does not necessarily cause conscious awareness of the action. <i>Closed loop control</i><sup id="cite_ref-Schmidt_1982_24-0" class="reference"><a href="#cite_note-Schmidt_1982-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 186">: 186 </span></sup> is a feedback based mechanism of motor control, where any act on the environment creates some sort of change that affects future performance through feedback. Closed loop motor control is best suited to continuously controlled actions, but does not work quickly enough for ballistic actions. Ballistic actions are actions that continue to the end without thinking about it, even when they no longer are appropriate.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2013)">citation needed</span></a></i>]</sup> Because feedback control relies on sensory information, it is as slow as sensory processing. These movements are subject to a speed-accuracy trade-off, because sensory processing is being used to control the movement, the faster the movement is carried out, the less accurate it becomes. </p> <div class="mw-heading mw-heading3"><h3 id="Open_loop_control">Open loop control</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=9" title="Edit section: Open loop control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The classical definition from Jack A. Adams is:<sup id="cite_ref-Adams_1971_22-1" class="reference"><a href="#cite_note-Adams_1971-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Adams_1976_23-1" class="reference"><a href="#cite_note-Adams_1976-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> “An open loop system has no feedback or mechanisms for error regulation. The input events for a system exert their influence, the system effects its transformation on the input and the system has an output...... A traffic light with fixed timing snarls traffic when the load is heavy and impedes the flow when the traffic is light. The system has no compensatory capability.” </p><p>Some movements, however, occur too quickly to integrate sensory information, and instead must rely on <a href="/wiki/Feed_forward_(control)" title="Feed forward (control)">feed forward control</a>. <i>Open loop control</i> is a feed forward form of motor control, and is used to control rapid, ballistic movements that end before any sensory information can be processed. To best study this type of control, most research focuses on deafferentation studies, often involving cats or monkeys whose sensory nerves have been disconnected from their spinal cords. Monkeys who lost all sensory information from their arms resumed normal behavior after recovering from the deafferentation procedure. Most skills were relearned, but fine motor control became very difficult.<sup id="cite_ref-Taub1966_25-0" class="reference"><a href="#cite_note-Taub1966-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> It has been shown that the open loop control can be adapted to different disease conditions and can therefore be used to extract signatures of different motor disorders by varying the cost functional governing the system.<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-heading2"><h2 id="Coordination">Coordination</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=10" title="Edit section: Coordination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A core motor control issue is coordinating the various components of the <a href="/wiki/Motor_system" title="Motor system">motor system</a> to act in unison to produce movement. </p><p>Peripheral neurons receive input from the <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> and innervate the muscles. In turn, muscles generate forces which actuate joints. Getting the pieces to work together is a challenging problem for the motor system and how this problem is resolved is an active area of study in motor control research. </p> <div class="mw-heading mw-heading3"><h3 id="Reflexes">Reflexes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=11" title="Edit section: Reflexes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In some cases the coordination of motor components is hard-wired, consisting of fixed neuromuscular pathways that are called <a href="/wiki/Reflex" title="Reflex"><i>reflexes</i></a>. Reflexes are typically characterized as automatic and fixed motor responses, and they occur on a much faster time scale than what is possible for reactions that depend on perceptual processing.<sup id="cite_ref-Dewhurst_1967_27-0" class="reference"><a href="#cite_note-Dewhurst_1967-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Reflexes play a fundamental role in stabilizing the motor system, providing almost immediate compensation for small perturbations and maintaining fixed execution patterns. Some reflex loops are routed solely through the spinal cord without receiving input from the brain, and thus do not require attention or conscious control. Others involve lower brain areas and can be influenced by prior instructions or intentions, but they remain independent of perceptual processing and online control. </p><p>The simplest reflex is the <i><a href="/wiki/Reflex_arc#Monosynaptic_vs._polysynaptic" title="Reflex arc">monosynaptic reflex</a></i> or short-loop reflex, such as the monosynaptic stretch response. In this example, <a href="/wiki/Ia_afferent" class="mw-redirect" title="Ia afferent">Ia afferent</a> neurons are activated by <a href="/wiki/Muscle_spindle" title="Muscle spindle">muscle spindles</a> when they deform due to the stretching of the muscle. In the spinal cord, these afferent neurons synapse directly onto <a href="/wiki/Alpha_motor_neurons" class="mw-redirect" title="Alpha motor neurons">alpha motor neurons</a> that regulate the contraction of the same muscle.<sup id="cite_ref-Pearson2000_28-0" class="reference"><a href="#cite_note-Pearson2000-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Thus, any stretching of a muscle automatically signals a reflexive contraction of that muscle, without any central control. As the name and the description implies, monosynaptic reflexes depend on a single synaptic connection between an afferent sensory neuron and efferent motor neuron. In general the actions of monosynaptic reflexes are fixed and cannot be controlled or influenced by intention or instruction. However, there is some evidence to suggest that the <a href="/wiki/Gain_(electronics)" title="Gain (electronics)">gain</a> or magnitude of these reflexes can be adjusted by context and experience.<sup id="cite_ref-Matthews_1986_29-0" class="reference"><a href="#cite_note-Matthews_1986-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p><p><i>Polysynaptic reflexes</i> or long-loop reflexes are reflex arcs which involve more than a single synaptic connection in the spinal cord. These loops may include cortical regions of the brain as well, and are thus slower than their monosynaptic counterparts due to the greater travel time. However, actions controlled by polysynaptic reflex loops are still faster than actions which require perceptual processing.<sup id="cite_ref-Schmidt_1988_30-0" class="reference"><a href="#cite_note-Schmidt_1988-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 171, 578">: 171, 578 </span></sup> While the actions of short-loop reflexes are fixed, polysynaptic reflexes can often be regulated by instruction or prior experience.<sup id="cite_ref-Evarts_1973_31-0" class="reference"><a href="#cite_note-Evarts_1973-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> A common example of a long loop reflex is the <a href="/wiki/Asymmetrical_tonic_neck_reflex" title="Asymmetrical tonic neck reflex">asymmetrical tonic neck reflex</a> observed in infants. </p> <div class="mw-heading mw-heading3"><h3 id="Synergies">Synergies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=12" title="Edit section: Synergies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A motor <i><a href="/wiki/Motor_coordination#Muscle_synergies" title="Motor coordination">synergy</a></i> is a neural organization of a multi-element system that (1) organizes sharing of a task among a set of elemental variables; and (2) ensures co-variation among elemental variables with the purpose to stabilize performance variables.<sup id="cite_ref-Latash2007_32-0" class="reference"><a href="#cite_note-Latash2007-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Alnajjar_2013_33-0" class="reference"><a href="#cite_note-Alnajjar_2013-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> The components of a synergy need not be physically connected, but instead are connected by their response to perceptual information about the particular motor task being executed. Synergies are learned, rather than being hardwired like reflexes, and are organized in a task-dependent manner; a synergy is structured for a particular action and not determined generally for the components themselves. <a href="/wiki/Nikolai_Bernstein" title="Nikolai Bernstein">Nikolai Bernstein</a> famously demonstrated synergies at work in the hammering actions of professional blacksmiths. The muscles of the arm controlling the movement of the hammer are informationally linked in such a way that errors and variability in one muscle are automatically compensated for by the actions of the other muscles. These compensatory actions are reflex-like in that they occur faster than perceptual processing would seem to allow, yet they are only present in expert performance, not in novices. In the case of blacksmiths, the synergy in question is organized specifically for hammering actions and is not a general purpose organization of the muscles of the arm. Synergies have two defining characteristics in addition to being task dependent; sharing and flexibility/stability.<sup id="cite_ref-Latash2008_34-0" class="reference"><a href="#cite_note-Latash2008-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p><p>"Sharing" requires that the execution of a particular motor task depends on the combined actions of all the components that make up the synergy. Often, there are more components involved than are strictly needed for the particular task (<a href="#Redundancy">see "Redundancy" below</a>), but the control of that motor task is distributed across all components nonetheless. A simple demonstration comes from a two-finger force production task, where participants are required to generate a fixed amount of force by pushing down on two force plates with two different fingers.<sup id="cite_ref-Scholz2002_35-0" class="reference"><a href="#cite_note-Scholz2002-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> In this task, participants generated a particular force output by combining the contributions of independent fingers. While the force produced by any single finger can vary, this variation is constrained by the action of the other such that the desired force is always generated. </p><p>Co-variation also provides "flexibility and stability" to motor tasks. Considering again the force production task, if one finger did not produce enough force, it could be compensated for by the other.<sup id="cite_ref-Scholz2002_35-1" class="reference"><a href="#cite_note-Scholz2002-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> The components of a motor synergy are expected to change their action to compensate for the errors and variability in other components that could affect the outcome of the motor task. This provides flexibility because it allows for multiple motor solutions to particular tasks, and it provides motor stability by preventing errors in individual motor components from affecting the task itself. </p><p>Synergies simplify the computational difficulty of motor control. Coordinating the numerous <a href="/wiki/Degrees_of_freedom_(mechanics)" title="Degrees of freedom (mechanics)">degrees of freedom</a> in the body is a challenging problem, both because of the tremendous complexity of the motor system, as well as the different levels at which this organization can occur (neural, muscular, kinematic, spatial, etc.). Because the components of a synergy are functionally coupled for a specific task, execution of motor tasks can be accomplished by activating the relevant synergy with a single neural signal.<sup id="cite_ref-Bernstein1967_36-0" class="reference"><a href="#cite_note-Bernstein1967-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The need to control all of the relevant components independently is removed because organization emerges automatically as a consequence of the systematic covariation of components. Similar to how reflexes are physically connected and thus do not require control of individual components by the central nervous system, actions can be executed through synergies with minimal executive control because they are functionally connected. Beside motor synergies, the term of sensory synergies has recently been introduced.<sup id="cite_ref-Alnajjar_2015_37-0" class="reference"><a href="#cite_note-Alnajjar_2015-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Sensory synergy are believed to play an important role in integrating the mixture of environmental inputs to provide low-dimensional information to the CNS thus guiding the recruitment of motor synergies. </p><p>Synergies are fundamental for controlling complex movements, such as the ones of the hand during grasping. Their importance has been demonstrated for both muscle control and in the kinematic domain in several studies, lately on studies including large cohorts of subjects.<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><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> The relevance of synergies for hand grasps is also enforced by studies on hand grasp taxonomies, showing muscular and kinematic similarities among specific groups of grasps, leading to specific clusters of movements.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Motor_Programs">Motor Programs</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=13" title="Edit section: Motor Programs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While synergies represent coordination derived from peripheral interactions of motor components, <a href="/wiki/Motor_program" title="Motor program">motor programs</a> are specific, pre-structured motor activation patterns that are generated and executed by a central controller (in the case of a biological organism, the brain).<sup id="cite_ref-Schmidt_1988_30-1" class="reference"><a href="#cite_note-Schmidt_1988-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 227">: 227 </span></sup> They represent at top-down approach to motor coordination, rather than the bottom-up approach offered by synergies. Motor programs are executed in an open-loop manner, although sensory information is most likely used to sense the current state of the organism and determine the appropriate goals. However, similar to <a href="/wiki/Central_pattern_generator" title="Central pattern generator">central pattern generators</a>, once the program has been executed, it cannot be altered online by additional sensory information. </p><p>Evidence for the existence of motor programs comes from studies of rapid movement execution and the difficulty associated with changing those movements once they have been initiated. For example, people who are asked to make fast arm swings have extreme difficulty in halting that movement when provided with a "STOP" signal after the movement has been initiated.<sup id="cite_ref-Henry1961_42-0" class="reference"><a href="#cite_note-Henry1961-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> This reversal difficulty persists even if the stop signal is presented after the initial "GO" signal but <i>before</i> the movement actually begins. This research suggests that once selection and execution of a motor program begins, it must run to completion before another action can be taken. This effect has been found even when the movement that is being executed by a particular motor program is prevented from occurring at all. People who attempt to execute particular movements (such as pushing with the arm), but unknowingly have the action of their body arrested before any movement can actually take place, show the same muscle activation patterns (including stabilizing and support activation that does not actually generate the movement) as when they are allowed to complete their intended action.<sup id="cite_ref-Wadman1979_43-0" class="reference"><a href="#cite_note-Wadman1979-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>Although the evidence for motor programs seems persuasive, there have been several important criticisms of the theory. The first is the problem of storage. If each movement an organism could generate requires its own motor program, it would seem necessary for that organism to possess an unlimited repository of such programs and where these would be kept is not clear. Aside from the enormous memory requirements such a facility would take, no motor program storage area in the brain has yet been identified. The second problem is concerned with novelty in movement. If a specific motor program is required for any particular movement, it is not clear how one would ever produce a novel movement. At best, an individual would have to practice any new movement before executing it with any success, and at worst, would be incapable of new movements because no motor program would exist for new movements. These difficulties have led to a more nuanced notion of motor programs known as <b>generalized motor programs</b>.<sup id="cite_ref-Schmidt_1988_30-2" class="reference"><a href="#cite_note-Schmidt_1988-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 240–257">: 240–257 </span></sup> A generalized motor program is a program for a particular <i>class</i> of action, rather than a specific movement. This program is parameterized by the context of the environment and the current state of the organism. </p> <div class="mw-heading mw-heading3"><h3 id="Redundancy">Redundancy</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=14" title="Edit section: Redundancy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An important issue for coordinating the motor system is the problem of the <b>redundancy</b> of motor degrees of freedom. As detailed in the "<a href="#Synergies">Synergies</a>" section, many actions and movements can be executed in multiple ways because functional synergies controlling those actions are able to co-vary without changing the outcome of the action. This is possible because there are more motor components involved in the production of actions than are generally required by the physical constraints on that action. For example, the human arm has seven joints which determine the position of the hand in the world. However, only three spatial dimensions are needed to specify any location the hand could be placed in. This excess of kinematic degrees of freedom means that there are multiple arm configurations that correspond to any particular location of the hand. </p><p>Some of the earliest and most influential work on the study of motor redundancy came from the Russian physiologist <a href="/wiki/Nikolai_Bernstein" title="Nikolai Bernstein">Nikolai Bernstein</a>. Bernstein's research was primarily concerned with understanding how coordination was developed for skilled actions. He observed that the redundancy of the motor system made it possible to execute actions and movements in a multitude of different ways while achieving equivalent outcomes.<sup id="cite_ref-Bernstein1967_36-1" class="reference"><a href="#cite_note-Bernstein1967-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> This equivalency in motor action means that there is no one-to-one correspondence between the desired movements and the coordination of the motor system needed to execute those movements. Any desired movement or action does not have a particular coordination of neurons, muscles, and kinematics that make it possible. This motor equivalency problem became known as the <b><a href="/wiki/Degrees_of_freedom_problem" title="Degrees of freedom problem">degrees of freedom problem</a></b> because it is a product of having redundant degrees of freedom available in the motor system. </p> <div class="mw-heading mw-heading2"><h2 id="Perception_in_motor_control">Perception in motor control</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=15" title="Edit section: Perception in motor control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Related, yet distinct from the issue of how the <i>processing</i> of sensory information affects the control of movements and actions is the question of how the perception of the world structures action. <a href="/wiki/Perception" title="Perception">Perception</a> is extremely important in motor control because it carries the relevant information about objects, environments and bodies which is used in organizing and executing actions and movements. What is perceived and how the subsequent information is used to organize the motor system is an ongoing area of research. </p> <div class="mw-heading mw-heading3"><h3 id="Model_based_control_strategies">Model based control strategies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=16" title="Edit section: Model based control strategies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most model based strategies of motor control rely on perceptual information, but assume that this information is not always useful, veridical or constant. Optical information is interrupted by eye blinks, motion is obstructed by objects in the environment, distortions can change the appearance of object shape. Model based and representational control strategies are those that rely on accurate <a href="/wiki/Internal_models" class="mw-redirect" title="Internal models">internal models</a> of the environment, constructed from a combination of perceptual information and prior knowledge, as the primary source information for planning and executing actions, even in the absence of perceptual information.<sup id="cite_ref-Kawato1999_44-0" class="reference"><a href="#cite_note-Kawato1999-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Inference_and_indirect_perception">Inference and indirect perception</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=17" title="Edit section: Inference and indirect perception"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many models of the perceptual system assume <a href="/wiki/Indirect_perception" class="mw-redirect" title="Indirect perception">indirect perception</a>, or the notion that the world that gets perceived is not identical to the actual environment. Environmental information must go through several stages before being perceived, and the transitions between these stages introduce ambiguity. What actually gets perceived is the mind's best guess about what is occurring in the environment based on previous experience. Support for this idea comes from the <a href="/wiki/Ames_room" title="Ames room">Ames room</a> illusion, where a distorted room causes the viewer to see objects known to be a constant size as growing or shrinking as they move around the room. The room itself is seen as being square, or at least consisting of right angles, as all previous rooms the perceiver has encountered have had those properties. Another example of this ambiguity comes from the <a href="/wiki/Doctrine_of_specific_nerve_energies" class="mw-redirect" title="Doctrine of specific nerve energies">doctrine of specific nerve energies</a>. The doctrine presents the finding that there are distinct nerve types for different types of sensory input, and these nerves respond in a characteristic way regardless of the method of stimulation. That is to say, the color red causes optical nerves to fire in a specific pattern that is processed by the brain as experiencing the color red. However, if that same nerve is electrically stimulated in an identical pattern, the brain could perceive the color red when no corresponding stimuli is present. </p> <div class="mw-heading mw-heading4"><h4 id="Forward_models">Forward models</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=18" title="Edit section: Forward models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Internal_model_(motor_control)#Forward_models" title="Internal model (motor control)">Forward models</a> are a predictive internal model of motor control that takes the available perceptual information, combined with a particular motor program, and tries to predict the outcome of the planned motor movement. Forward models structure action by determining how the forces, velocities, and positions of motor components affect changes in the environment and in the individual. It is proposed that forward models help with the <a href="/wiki/Neural_control_of_limb_stiffness" title="Neural control of limb stiffness">Neural control of limb stiffness</a> when individuals interact with their environment. Forward models are thought to use motor programs as input to predict the outcome of an action. An error signal is generated when the predictions made by a forward model do not match the actual outcome of the movement, prompting an update of an existing model and providing a mechanism for learning. These models explain why it is impossible to tickle yourself. A sensation is experienced as ticklish when it is unpredictable. However, forward models predict the outcome of your motor movements, meaning the motion is predictable, and therefore not ticklish.<sup id="cite_ref-Blakemore2000_45-0" class="reference"><a href="#cite_note-Blakemore2000-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> </p><p>Evidence for forward models comes from studies of motor adaptation. When a person's goal-directed reaching movements are perturbed by a force field, they gradually, but steadily, adapt the movement of their arm to allow them to again reach their goal. However, they do so in such a way that preserves some high level movement characteristics; bell-shaped velocity profiles, straight line translation of the hand, and smooth, continuous movements.<sup id="cite_ref-Shadmehr1994_46-0" class="reference"><a href="#cite_note-Shadmehr1994-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> These movement features are recovered, despite the fact that they require startlingly different arm dynamics (i.e. torques and forces). This recovery provides evidence that what is motivating movement is a particular motor plan, and the individual is using a forward model to predict how arm dynamics change the movement of the arm to achieve particular task level characteristics. Differences between the expected arm movement and the observed arm movement produces an error signal which is used as the basis for learning. Additional evidence for forward models comes from experiments which require subjects to determine the location of an effector following an unvisualized movement<sup id="cite_ref-Wolpert1995_47-0" class="reference"><a href="#cite_note-Wolpert1995-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Inverse_models">Inverse models</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=19" title="Edit section: Inverse models"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Internal_model_(motor_control)#Inverse_models" title="Internal model (motor control)">Inverse models</a> predict the necessary movements of motor components to achieve a desired perceptual outcome. They can also take the outcome of a motion and attempt to determine the sequence of motor commands that resulted in that state. These types of models are particularly useful for open loop control, and allow for specific types of movements, such as fixating on a stationary object while the head is moving. Complementary to forward models, inverse models attempt to estimate how to achieve a particular perceptual outcome in order to generate the appropriate motor plan. Because inverse models and forward model are so closely associated, studies of internal models are often used as evidence for the roles of both model types in action. </p><p>Motor adaptation studies, therefore, also make a case for inverse models. Motor movements seem to follow predefined "plans" that preserve certain invariant features of the movement. In the reaching task mentioned above, the persistence of bell-shaped velocity profiles and smooth, straight hand trajectories provides evidence for the existence of such plans.<sup id="cite_ref-Shadmehr1994_46-1" class="reference"><a href="#cite_note-Shadmehr1994-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Movements that achieve these desired task-level outcomes are estimated by an inverse model. Adaptation therefore proceeds as a process of estimating the necessary movements with an inverse model, simulating with a forward model the outcome of those movement plans, observing the difference between the desired outcome and the actual outcome, and updating the models for a future attempt. </p> <div class="mw-heading mw-heading3"><h3 id="Information_based_control">Information based control</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=20" title="Edit section: Information based control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An alternative to model based control is <b>information based control</b>. Informational control strategies organize movements and actions based on perceptual information about the environment, rather than on <a href="/wiki/Cognitive_model" title="Cognitive model">cognitive models</a> or representations of the world. The actions of the motor system are organized by information about the environment and information about the current state of the agent.<sup id="cite_ref-Warren2006_48-0" class="reference"><a href="#cite_note-Warren2006-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Information based control strategies often treat the environment and the organism as a single system, with action proceeding as a natural consequence of the interactions of this system. A core assumption of information based control strategies is that perceptions of the environment are rich in information and veridical for the purposes of producing actions. This runs counter to the assumptions of indirect perception made by model based control strategies. </p> <div class="mw-heading mw-heading4"><h4 id="Direct_perception">Direct perception</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=21" title="Edit section: Direct perception"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><b>Direct perception</b> in the cognitive sense is related to the philosophical notion of <a href="/wiki/Na%C3%AFve_realism" title="Naïve realism">naïve or direct realism</a> in that it is predicated on the assumption that what we perceive is what is actually in the world. James J. Gibson is credited with recasting direct perception as <a href="/wiki/Ecological_psychology#Gibson" title="Ecological psychology">ecological perception</a>.<sup id="cite_ref-Gibson1986_49-0" class="reference"><a href="#cite_note-Gibson1986-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> While the problem of indirect perception proposes that physical information about object in our environment is not available due to the ambiguity of sensory information, proponents of direct perception (like Gibson) suggest that the relevant information specified in <a href="/wiki/Ambient_optic_array" title="Ambient optic array">ambient optic array</a> is the distal physical properties of objects. This specifying information reveals the action opportunities the environment affords. These <a href="/wiki/Affordance" title="Affordance">affordances</a> are directly perceivable without ambiguity, and thus preclude the need for internal models or representations of the world. Affordances exist only as a byproduct of the interactions between an agent and its environment, and thus perception is an "<a href="/wiki/Ecological" class="mw-redirect" title="Ecological">ecological</a>" endeavor, depending on the whole agent/environment system rather than on the agent in isolation. </p><p>Because affordances are action possibilities, perception is directly connected to the production of actions and movements. The role of perception is to provide information that specifies how actions should be organized and controlled,<sup id="cite_ref-Michaels1981_50-0" class="reference"><a href="#cite_note-Michaels1981-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> and the motor system is "tuned" to respond to specific type of information in particular ways. Through this relationship, control of the motor system and the execution of actions is dictated by the information of the environment. As an example, a doorway "affords" passing through, but a wall does not. How one might pass through a doorway is specified by the visual information received from the environment, as well as the information perceived about one's own body. Together, this information determines the pass-ability of a doorway, but not a wall. In addition, the act of moving towards and passing through the doorway generates more information and this in turn specifies further action. The conclusion of direct perception is that actions and perceptions are critically linked and one cannot be fully understood without the other. </p> <div class="mw-heading mw-heading4"><h4 id="Behavioral_dynamics">Behavioral dynamics</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=22" title="Edit section: Behavioral dynamics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Building on the assumptions of direct perception behavioral dynamics is a behavioral control theory that treats perceptual organisms as dynamic systems that respond to informational variables with actions, in a functional manner.<sup id="cite_ref-Warren2006_48-1" class="reference"><a href="#cite_note-Warren2006-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Under this understanding of behavior, actions unfold as the natural consequence of the interaction between the organisms and the available information about the environment, which specified in body-relevant variables. Much of the research in behavioral dynamics has focused on locomotion, where visually specified information (such as optic flow, time-to-contact, optical expansion, etc.) is used to determine how to navigate the environment<sup id="cite_ref-Fajen2003_51-0" class="reference"><a href="#cite_note-Fajen2003-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fajen2011_52-0" class="reference"><a href="#cite_note-Fajen2011-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Interaction forces between the human and the environment also affect behavioral dynamics as seen in by the <a href="/wiki/Neural_control_of_limb_stiffness" title="Neural control of limb stiffness">Neural control of limb stiffness</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Planning_in_motor_control">Planning in motor control</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=23" title="Edit section: Planning in motor control"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Individual_movement_optimization">Individual movement optimization</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=24" title="Edit section: Individual movement optimization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are several mathematical models that describe how the central nervous system (CNS) derives reaching movements of limbs and eyes. The minimum jerk model states that the CNS minimizes <a href="/wiki/Jerk_(physics)" title="Jerk (physics)">jerk</a> of a limb endpoint trajectory over the time of reaching, which results in a smooth trajectory.<sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> However, this model is based solely on the kinematics of movement and does not consider the underlying dynamics of the musculoskeletal system. Hence, the minimum torque-change model was introduced as an alternative, where the CNS minimizes the joint <a href="/wiki/Torque" title="Torque">torque</a> change over the time of reaching.<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> </p><p>Later it was argued that there is no clear explanation about how could the CNS actually estimate complex quantities such as jerk or torque change and then integrate them over the duration of a trajectory. In response, model based on signal-dependent noise was proposed instead, which states that the CNS selects a trajectory by minimizing the variance of the final position of the limb endpoint. Since there is a motor noise in the neural system that is proportional to the activation of the muscles, the faster movements induce more motor noise and are thus less precise.<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> This is also in line with the <a href="/wiki/Fitts%27_Law" class="mw-redirect" title="Fitts' Law">Fitts' Law</a> and speed-accuracy trade-off.<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> <a href="/wiki/Optimal_control" title="Optimal control">Optimal control</a> theory was used to further extend the model based on signal-dependent noise, where the CNS optimizes an objective function that consists of a term related to accuracy and additionally a term related to metabolic cost of movement.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>Another type of models is based on cost-benefit trade-off, where the objective function includes metabolic cost of movement and a subjective reward related to reaching the target accurately. In this case the reward for a successful reach within the desired target is discounted by the duration of reaching, since the gained reward is perceived less valuable when spending more time on it.<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> However, these models were deterministic and did not account for motor noise, which is an essential property of stochastic motor control that results in speed-accuracy trade-off. To address that, a new model was later proposed to incorporate the motor noise and to unify cost-benefit and speed-accuracy trade-offs.<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> <div class="mw-heading mw-heading3"><h3 id="Multi-component_movements">Multi-component movements</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=25" title="Edit section: Multi-component movements"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Some studies observed that the CNS can split a complex movement into sub-movements. The initial sub-movement tends to be fast and imprecise in order to bring the limb endpoint into vicinity of the target as soon as possible. Then, the final sub-movement tends to be slow and precise in order to correct for accumulated error by the first initial sub-movement and to successfully reach the target.<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><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> A later study further explored how the CNS selects a temporary target of the initial sub-movement in different conditions. For example, when the actual target size decreases and thus complexity increases, the temporary target of the initial sub-movement moves away from the actual target in order to give more space for the final corrective action. Longer reaching distances have a similar effect, since more error is accumulated in the initial sub-movement and thus requiring more complex final correction. In less complex conditions, when the final actual target is large and the movement is short, the CNS tends to use a single movement, without splitting it into multiple competents.<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> </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=Motor_control&action=edit&section=26" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Motor_learning" title="Motor learning">Motor learning</a></li> <li><a href="/wiki/Motor_skill" title="Motor skill">Motor skill</a></li> <li><a href="/wiki/Motor_coordination" title="Motor coordination">Motor coordination</a></li> <li><a href="/wiki/Motor_cortex" title="Motor cortex">Motor cortex</a></li> <li><a href="/wiki/Multisensory_integration" title="Multisensory integration">Multisensory integration</a></li> <li><a href="/wiki/Proprioception" title="Proprioception">Proprioception</a></li> <li><a href="/wiki/Sensory_processing" title="Sensory processing">Sensory processing</a></li> <li><a href="/wiki/Sensory-motor_coupling" title="Sensory-motor coupling">Sensory-motor coupling</a></li> <li><a href="/wiki/Two-alternative_forced_choice" title="Two-alternative forced choice">Two-alternative forced choice</a></li> <li><a href="/wiki/Psychomotor_learning" title="Psychomotor learning">Psychomotor learning</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=Motor_control&action=edit&section=27" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFSibson1850" class="citation journal cs1 cs1-prop-long-vol">Sibson F (1850). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=U58EAAAAQAAJ&pg=PA181">"On The Causes Which Excite And Influence Respiration In Health And Disease"</a>. <i>The Transactions of the Provincial Medical and Surgical Association</i>. 5 - New Series: 181–350. <q>In all these instances the act of inspiration is excited through the reflex function of the nervous system -- the sudden impression made on the skin stimulates the extremities of the incident nerves; the stimulus is conveyed by the incident nerves to the spinal nervous centre, and is thence transmitted back over the motor nerves of inspiration. That these respiratory movements are purely excito-motor, and performed without the intervention of sensation, in many of those instances in which the excited movements are most energetic, is proved by the case with which remarkable movements of respiration were occasioned by stimulating the surface in cases of syncope, hysteria, and epilepsy, cases in which sensation was altogether absent, and was only restored after repeatedly stimulating the surface, and so inducing deep reflex inspirations again and again by exciting the incident nerves. [Page 206]</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Transactions+of+the+Provincial+Medical+and+Surgical+Association&rft.atitle=On+The+Causes+Which+Excite+And+Influence+Respiration+In+Health+And+Disease.&rft.volume=5+-+New+Series&rft.pages=181-350&rft.date=1850&rft.aulast=Sibson&rft.aufirst=F&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DU58EAAAAQAAJ%26pg%3DPA181&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Rosenbaum1991-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-Rosenbaum1991_2-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRosenbaum1991" class="citation book cs1">Rosenbaum DA (1991). <i>Human motor control</i>. San Diego, CA: Academic Press. p. 411. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-597300-7" title="Special:BookSources/978-0-12-597300-7"><bdi>978-0-12-597300-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Human+motor+control&rft.place=San+Diego%2C+CA&rft.pages=411&rft.pub=Academic+Press&rft.date=1991&rft.isbn=978-0-12-597300-7&rft.aulast=Rosenbaum&rft.aufirst=DA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Wise2002-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Wise2002_3-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWiseShadmehr2002" class="citation book cs1">Wise SP, Shadmehr R (July 10, 2002). "Motor Control". <i>Encyclopedia of the Human Brain</i>. Academic Press. pp. 137–157. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-227210-3" title="Special:BookSources/978-0-12-227210-3"><bdi>978-0-12-227210-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Motor+Control&rft.btitle=Encyclopedia+of+the+Human+Brain&rft.pages=137-157&rft.pub=Academic+Press&rft.date=2002-07-10&rft.isbn=978-0-12-227210-3&rft.aulast=Wise&rft.aufirst=SP&rft.au=Shadmehr%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Franklin_425–442-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Franklin_425–442_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Franklin_425–442_4-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFranklinWolpert2011" class="citation journal cs1">Franklin DW, Wolpert DM (November 2011). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuron.2011.10.006">"Computational mechanisms of sensorimotor control"</a>. <i>Neuron</i>. <b>72</b> (3): 425–442. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.neuron.2011.10.006">10.1016/j.neuron.2011.10.006</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/22078503">22078503</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuron&rft.atitle=Computational+mechanisms+of+sensorimotor+control&rft.volume=72&rft.issue=3&rft.pages=425-442&rft.date=2011-11&rft_id=info%3Adoi%2F10.1016%2Fj.neuron.2011.10.006&rft_id=info%3Apmid%2F22078503&rft.aulast=Franklin&rft.aufirst=DW&rft.au=Wolpert%2C+DM&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.neuron.2011.10.006&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWolpert2011" class="citation web cs1">Wolpert D (3 November 2011). <a rel="nofollow" class="external text" href="https://www.ted.com/talks/daniel_wolpert_the_real_reason_for_brains">"The real reason for brains"</a>. TED Conferences, LLC<span class="reference-accessdate">. Retrieved <span class="nowrap">2020-03-27</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=The+real+reason+for+brains&rft.pub=TED+Conferences%2C+LLC&rft.date=2011-11-03&rft.aulast=Wolpert&rft.aufirst=D&rft_id=https%3A%2F%2Fwww.ted.com%2Ftalks%2Fdaniel_wolpert_the_real_reason_for_brains&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Kernell,_Daniel._2006-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kernell,_Daniel._2006_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kernell,_Daniel._2006_6-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKernell2006" class="citation book cs1">Kernell D (2006). <i>The motoneurone and its muscle fibres</i>. 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"Deep homology of arthropod central complex and vertebrate basal ganglia". <i>Science</i>. <b>340</b> (6129): 157–161. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2013Sci...340..157S">2013Sci...340..157S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1231828">10.1126/science.1231828</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23580521">23580521</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:206546163">206546163</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Deep+homology+of+arthropod+central+complex+and+vertebrate+basal+ganglia&rft.volume=340&rft.issue=6129&rft.pages=157-161&rft.date=2013-04&rft_id=info%3Adoi%2F10.1126%2Fscience.1231828&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A206546163%23id-name%3DS2CID&rft_id=info%3Apmid%2F23580521&rft_id=info%3Abibcode%2F2013Sci...340..157S&rft.aulast=Strausfeld&rft.aufirst=NJ&rft.au=Hirth%2C+F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Donders_1969-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-Donders_1969_20-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDonders1969" class="citation journal cs1">Donders FC (1969). "On the speed of mental processes". <i>Acta Psychologica</i>. <b>30</b>: 412–431. <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%2F0001-6918%2869%2990065-1">10.1016/0001-6918(69)90065-1</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/5811531">5811531</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Acta+Psychologica&rft.atitle=On+the+speed+of+mental+processes&rft.volume=30&rft.pages=412-431&rft.date=1969&rft_id=info%3Adoi%2F10.1016%2F0001-6918%2869%2990065-1&rft_id=info%3Apmid%2F5811531&rft.aulast=Donders&rft.aufirst=FC&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Hick_1952-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-Hick_1952_21-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHick1952" class="citation journal cs1">Hick WE (1952). <a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F17470215208416600">"On the rate of gain of information"</a>. <i>Quarterly Journal of Experimental Psychology</i>. <b>4</b> (1): 11–26. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F17470215208416600">10.1080/17470215208416600</a></span>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:39060506">39060506</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Quarterly+Journal+of+Experimental+Psychology&rft.atitle=On+the+rate+of+gain+of+information&rft.volume=4&rft.issue=1&rft.pages=11-26&rft.date=1952&rft_id=info%3Adoi%2F10.1080%2F17470215208416600&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A39060506%23id-name%3DS2CID&rft.aulast=Hick&rft.aufirst=WE&rft_id=https%3A%2F%2Fdoi.org%2F10.1080%252F17470215208416600&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Adams_1971-22"><span class="mw-cite-backlink">^ <a href="#cite_ref-Adams_1971_22-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Adams_1971_22-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAdams1971" class="citation journal cs1">Adams JA (1971). "A closed loop theory of motor learning". <i>Journal of Motor Behavior</i>. <b>3</b> (2): 111-150 (116). <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222895.1971.10734898">10.1080/00222895.1971.10734898</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15155169">15155169</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Motor+Behavior&rft.atitle=A+closed+loop+theory+of+motor+learning&rft.volume=3&rft.issue=2&rft.pages=111-150+%28116%29&rft.date=1971&rft_id=info%3Adoi%2F10.1080%2F00222895.1971.10734898&rft_id=info%3Apmid%2F15155169&rft.aulast=Adams&rft.aufirst=JA&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-Adams_1976-23"><span class="mw-cite-backlink">^ <a href="#cite_ref-Adams_1976_23-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Adams_1976_23-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAdams1976" class="citation book cs1">Adams JA (1976). 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"A century later: Woodworth's (1899) two-component model of goal-directed aiming". <i>Psychological Bulletin</i>. <b>127</b> (3): 342–357. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1037%2F0033-2909.127.3.342">10.1037/0033-2909.127.3.342</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11393300">11393300</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Psychological+Bulletin&rft.atitle=A+century+later%3A+Woodworth%27s+%281899%29+two-component+model+of+goal-directed+aiming&rft.volume=127&rft.issue=3&rft.pages=342-357&rft.date=2001-05&rft_id=info%3Adoi%2F10.1037%2F0033-2909.127.3.342&rft_id=info%3Apmid%2F11393300&rft.aulast=Elliott&rft.aufirst=D&rft.au=Helsen%2C+WF&rft.au=Chua%2C+R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> <li id="cite_note-63"><span class="mw-cite-backlink"><b><a href="#cite_ref-63">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPeternelBabič2019" class="citation journal cs1">Peternel L, Babič J (December 2019). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6934494">"Target of initial sub-movement in multi-component arm-reaching strategy"</a>. <i>Scientific Reports</i>. <b>9</b> (1): 20101. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2019NatSR...920101P">2019NatSR...920101P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fs41598-019-56430-x">10.1038/s41598-019-56430-x</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/PMC6934494">6934494</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/31882708">31882708</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Scientific+Reports&rft.atitle=Target+of+initial+sub-movement+in+multi-component+arm-reaching+strategy&rft.volume=9&rft.issue=1&rft.pages=20101&rft.date=2019-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6934494%23id-name%3DPMC&rft_id=info%3Apmid%2F31882708&rft_id=info%3Adoi%2F10.1038%2Fs41598-019-56430-x&rft_id=info%3Abibcode%2F2019NatSR...920101P&rft.aulast=Peternel&rft.aufirst=L&rft.au=Babi%C4%8D%2C+J&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6934494&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=28" 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="CITEREFSchmidtLee2011" class="citation book cs1">Schmidt RA, Lee TD (2011). <i>Motor control and learning : a behavioral emphasis</i>. Champaign, IL: Human Kinetics. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7360-7961-7" title="Special:BookSources/978-0-7360-7961-7"><bdi>978-0-7360-7961-7</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/814261802">814261802</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Motor+control+and+learning+%3A+a+behavioral+emphasis&rft.place=Champaign%2C+IL&rft.pub=Human+Kinetics&rft.date=2011&rft_id=info%3Aoclcnum%2F814261802&rft.isbn=978-0-7360-7961-7&rft.aulast=Schmidt&rft.aufirst=RA&rft.au=Lee%2C+TD&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFShadmehrWise2005" class="citation book cs1">Shadmehr R, Wise SP (2005). <i>The computational neurobiology of reaching and pointing : a foundation for motor learning</i>. Cambridge, Mass.: MIT Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-262-19508-9" title="Special:BookSources/978-0-262-19508-9"><bdi>978-0-262-19508-9</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/54529569">54529569</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+computational+neurobiology+of+reaching+and+pointing+%3A+a+foundation+for+motor+learning&rft.place=Cambridge%2C+Mass.&rft.pub=MIT+Press&rft.date=2005&rft_id=info%3Aoclcnum%2F54529569&rft.isbn=978-0-262-19508-9&rft.aulast=Shadmehr&rft.aufirst=R&rft.au=Wise%2C+SP&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBlagouchineMoreau2009" class="citation journal cs1">Blagouchine IV, Moreau E (November 2009). "Control of a speech robot via an optimum neural-network-based internal model with constraints". <i>IEEE Transactions on Robotics</i>. <b>26</b> (1): 142–159. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1109%2FTRO.2009.2033331">10.1109/TRO.2009.2033331</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:8415982">8415982</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=IEEE+Transactions+on+Robotics&rft.atitle=Control+of+a+speech+robot+via+an+optimum+neural-network-based+internal+model+with+constraints.&rft.volume=26&rft.issue=1&rft.pages=142-159&rft.date=2009-11&rft_id=info%3Adoi%2F10.1109%2FTRO.2009.2033331&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A8415982%23id-name%3DS2CID&rft.aulast=Blagouchine&rft.aufirst=IV&rft.au=Moreau%2C+E&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></li></ul> </div> <div class="mw-heading mw-heading3"><h3 id="Research_in_athletes">Research in athletes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Motor_control&action=edit&section=29" title="Edit section: Research in athletes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239549316"><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGray2011" class="citation journal cs1">Gray R (2011). "Links Between Attention, Performance Pressure, and Movement in Skilled Motor Action". <i>Current Directions in Psychological Science</i>. <b>20</b> (5): 301–306. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1177%2F0963721411416572">10.1177/0963721411416572</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:15905566">15905566</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Directions+in+Psychological+Science&rft.atitle=Links+Between+Attention%2C+Performance+Pressure%2C+and+Movement+in+Skilled+Motor+Action&rft.volume=20&rft.issue=5&rft.pages=301-306&rft.date=2011&rft_id=info%3Adoi%2F10.1177%2F0963721411416572&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A15905566%23id-name%3DS2CID&rft.aulast=Gray&rft.aufirst=R&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMikheevMohrAfanasievLandis2002" class="citation journal cs1">Mikheev M, Mohr C, Afanasiev S, Landis T, Thut G (2002). "Motor control and cerebral hemispheric specialization in highly qualified judo wrestlers". <i>Neuropsychologia</i>. <b>40</b> (8): 1209–1219. <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%2Fs0028-3932%2801%2900227-5">10.1016/s0028-3932(01)00227-5</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11931924">11931924</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:7825661">7825661</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neuropsychologia&rft.atitle=Motor+control+and+cerebral+hemispheric+specialization+in+highly+qualified+judo+wrestlers&rft.volume=40&rft.issue=8&rft.pages=1209-1219&rft.date=2002&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A7825661%23id-name%3DS2CID&rft_id=info%3Apmid%2F11931924&rft_id=info%3Adoi%2F10.1016%2Fs0028-3932%2801%2900227-5&rft.aulast=Mikheev&rft.aufirst=M&rft.au=Mohr%2C+C&rft.au=Afanasiev%2C+S&rft.au=Landis%2C+T&rft.au=Thut%2C+G&rfr_id=info%3Asid%2Fen.wikipedia.org%3AMotor+control" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPaulGanesanSandhuSimon2012" class="citation journal cs1">Paul M, Ganesan S, Sandhu J, Simon J (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.4103%2F1947-489X.210753">"Effect of Sensory Motor Rhythm Neurofeedback on Psycho-physiological, Electroencephalographic Measures and Performance of Archery Players"</a>. <i>Ibnosina Journal of Medicine and Biomedical Sciences</i>. <b>4</b> (2): 32–39. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.4103%2F1947-489X.210753">10.4103/1947-489X.210753</a></span>.</cite><span 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href="/wiki/Template:Neuroscience" title="Template:Neuroscience"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Neuroscience" title="Template talk:Neuroscience"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Neuroscience" title="Special:EditPage/Template:Neuroscience"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Neuroscience" style="font-size:114%;margin:0 4em"><a href="/wiki/Neuroscience" title="Neuroscience">Neuroscience</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="3"><div> <ul><li><a href="/wiki/Outline_of_neuroscience" title="Outline of neuroscience">Outline</a></li> <li><a href="/wiki/History_of_neuroscience" title="History of neuroscience">History</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Basic_research" title="Basic research">Basic<br />science</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/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a></li> <li><a href="/wiki/Behavioural_genetics" title="Behavioural genetics">Behavioral genetics</a></li> <li><a href="/wiki/Brain_mapping" title="Brain mapping">Brain mapping</a></li> <li><a href="/wiki/Brain-reading" title="Brain-reading">Brain-reading</a></li> <li><a href="/wiki/Cellular_neuroscience" title="Cellular neuroscience">Cellular neuroscience</a></li> <li><a href="/wiki/Computational_neuroscience" title="Computational neuroscience">Computational neuroscience</a></li> <li><a href="/wiki/Connectomics" title="Connectomics">Connectomics</a></li> <li><a href="/wiki/Imaging_genetics" title="Imaging genetics">Imaging genetics</a></li> <li><a href="/wiki/Integrative_neuroscience" title="Integrative neuroscience">Integrative neuroscience</a></li> <li><a href="/wiki/Molecular_neuroscience" title="Molecular neuroscience">Molecular neuroscience</a></li> <li><a href="/wiki/Neural_decoding" title="Neural decoding">Neural decoding</a></li> <li><a href="/wiki/Neural_engineering" title="Neural engineering">Neural engineering</a></li> <li><a href="/wiki/Neuroanatomy" title="Neuroanatomy">Neuroanatomy</a></li> <li><a href="/wiki/Neurobiology" class="mw-redirect" title="Neurobiology">Neurobiology</a></li> <li><a href="/wiki/Neurochemistry" title="Neurochemistry">Neurochemistry</a></li> <li><a href="/wiki/Neuroendocrinology" title="Neuroendocrinology">Neuroendocrinology</a></li> <li><a href="/wiki/Neurogenetics" title="Neurogenetics">Neurogenetics</a></li> <li><a href="/wiki/Neuroinformatics" title="Neuroinformatics">Neuroinformatics</a></li> <li><a href="/wiki/Neurometrics" title="Neurometrics">Neurometrics</a></li> <li><a href="/wiki/Neuromorphology" title="Neuromorphology">Neuromorphology</a></li> <li><a href="/wiki/Neurophysics" title="Neurophysics">Neurophysics</a></li> <li><a href="/wiki/Neurophysiology" title="Neurophysiology">Neurophysiology</a></li> <li><a href="/wiki/Systems_neuroscience" title="Systems neuroscience">Systems neuroscience</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Gray739.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/130px-Gray739.png" decoding="async" width="130" height="99" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/195px-Gray739.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/96/Gray739.png/260px-Gray739.png 2x" data-file-width="500" data-file-height="379" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Clinical_neuroscience" title="Clinical neuroscience">Clinical<br />neuroscience</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioral_neurology" title="Behavioral neurology">Behavioral neurology</a></li> <li><a href="/wiki/Clinical_neurophysiology" title="Clinical neurophysiology">Clinical neurophysiology</a></li> <li><a href="/wiki/Epileptology" class="mw-redirect" title="Epileptology">Epileptology</a></li> <li><a href="/wiki/Neurocardiology" title="Neurocardiology">Neurocardiology</a></li> <li><a href="/wiki/Neuroepidemiology" title="Neuroepidemiology">Neuroepidemiology</a></li> <li><a href="/wiki/Enteric_nervous_system#Function" title="Enteric nervous system">Neurogastroenterology</a></li> <li><a href="/wiki/Neuroimmunology" title="Neuroimmunology">Neuroimmunology</a></li> <li><a href="/wiki/Neurointensive_care" title="Neurointensive care">Neurointensive care</a></li> <li><a href="/wiki/Neurology" title="Neurology">Neurology</a></li> <li><a href="/wiki/Neuro-oncology" title="Neuro-oncology">Neuro-oncology</a></li> <li><a href="/wiki/Neuro-ophthalmology" title="Neuro-ophthalmology">Neuro-ophthalmology</a></li> <li><a href="/wiki/Neuropathology" title="Neuropathology">Neuropathology</a></li> <li><a href="/wiki/Neuropharmacology" title="Neuropharmacology">Neuropharmacology</a></li> <li><a href="/wiki/Neuroprosthetics" title="Neuroprosthetics">Neuroprosthetics</a></li> <li><a href="/wiki/Neuropsychiatry" title="Neuropsychiatry">Neuropsychiatry</a></li> <li><a href="/wiki/Neuroradiology" title="Neuroradiology">Neuroradiology</a></li> <li><a href="/wiki/Neurorehabilitation" title="Neurorehabilitation">Neurorehabilitation</a></li> <li><a href="/wiki/Neurosurgery" title="Neurosurgery">Neurosurgery</a></li> <li><a href="/wiki/Neurotology" title="Neurotology">Neurotology</a></li> <li><a href="/wiki/Neurovirology" title="Neurovirology">Neurovirology</a></li> <li><a href="/wiki/Nutritional_neuroscience" title="Nutritional neuroscience">Nutritional neuroscience</a></li> <li><a href="/wiki/Psychiatry" title="Psychiatry">Psychiatry</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">Cognitive<br />neuroscience</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective neuroscience</a></li> <li><a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">Behavioral neuroscience</a></li> <li><a href="/wiki/Chronobiology" title="Chronobiology">Chronobiology</a></li> <li><a href="/wiki/Molecular_cellular_cognition" title="Molecular cellular cognition">Molecular cellular cognition</a></li> <li><a class="mw-selflink selflink">Motor control</a></li> <li><a href="/wiki/Neurolinguistics" title="Neurolinguistics">Neurolinguistics</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Sensory_neuroscience" title="Sensory neuroscience">Sensory neuroscience</a></li> <li><a href="/wiki/Social_cognitive_neuroscience" title="Social cognitive neuroscience">Social cognitive neuroscience</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Interdisciplinary<br />fields</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Consumer_neuroscience" title="Consumer neuroscience">Consumer neuroscience</a></li> <li><a href="/wiki/Cultural_neuroscience" title="Cultural neuroscience">Cultural neuroscience</a></li> <li><a href="/wiki/Educational_neuroscience" title="Educational neuroscience">Educational neuroscience</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Global_neurosurgery" title="Global neurosurgery">Global neurosurgery</a></li> <li><a href="/wiki/Neuroanthropology" title="Neuroanthropology">Neuroanthropology</a></li> <li><a href="/wiki/Neural_engineering" title="Neural engineering">Neural engineering</a></li> <li><a href="/wiki/Neurotechnology" title="Neurotechnology">Neurobiotics</a></li> <li><a href="/wiki/Neurocriminology" title="Neurocriminology">Neurocriminology</a></li> <li><a href="/wiki/Neuroeconomics" title="Neuroeconomics">Neuroeconomics</a></li> <li><a href="/wiki/Neuroepistemology" title="Neuroepistemology">Neuroepistemology</a></li> <li><a href="/wiki/Neuroesthetics" title="Neuroesthetics">Neuroesthetics</a></li> <li><a href="/wiki/Neuroethics" title="Neuroethics">Neuroethics</a></li> <li><a href="/wiki/Neuroethology" title="Neuroethology">Neuroethology</a></li> <li><a href="/wiki/Neurohistory" title="Neurohistory">Neurohistory</a></li> <li><a href="/wiki/Neurolaw" title="Neurolaw">Neurolaw</a></li> <li><a href="/wiki/Neuromarketing" title="Neuromarketing">Neuromarketing</a></li> <li><a href="/wiki/Neuromorphic_engineering" class="mw-redirect" title="Neuromorphic engineering">Neuromorphic engineering</a></li> <li><a href="/wiki/Neurophenomenology" title="Neurophenomenology">Neurophenomenology</a></li> <li><a href="/wiki/Neurophilosophy" title="Neurophilosophy">Neurophilosophy</a></li> <li><a href="/wiki/Neuropolitics" title="Neuropolitics">Neuropolitics</a></li> <li><a href="/wiki/Neurorobotics" title="Neurorobotics">Neurorobotics</a></li> <li><a href="/wiki/Neuroscience_of_religion" title="Neuroscience of religion">Neurotheology</a></li> <li><a href="/wiki/Paleoneurobiology" title="Paleoneurobiology">Paleoneurobiology</a></li> <li><a href="/wiki/Social_neuroscience" title="Social neuroscience">Social neuroscience</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Concepts</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Brain%E2%80%93computer_interface" title="Brain–computer interface">Brain–computer interface</a></li> <li><a href="/wiki/Development_of_the_nervous_system" title="Development of the nervous system">Development of the nervous system</a></li> <li><a href="/wiki/Artificial_neural_network" class="mw-redirect" title="Artificial neural network">Neural network (artificial)</a></li> <li><a href="/wiki/Neural_circuit" title="Neural circuit">Neural network (biological)</a></li> <li><a href="/wiki/Detection_theory" title="Detection theory">Detection theory</a></li> <li><a href="/wiki/Intraoperative_neurophysiological_monitoring" title="Intraoperative neurophysiological monitoring">Intraoperative neurophysiological monitoring</a></li> <li><a href="/wiki/Neurochip" title="Neurochip">Neurochip</a></li> <li><a href="/wiki/Neurodegenerative_disease" title="Neurodegenerative disease">Neurodegenerative disease</a></li> <li><a href="/wiki/Neurodevelopmental_disorder" title="Neurodevelopmental disorder">Neurodevelopmental disorder</a></li> <li><a href="/wiki/Neurodiversity" title="Neurodiversity">Neurodiversity</a></li> <li><a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a></li> <li><a href="/wiki/Neuroimaging" title="Neuroimaging">Neuroimaging</a></li> <li><a href="/wiki/Neuroimmune_system" title="Neuroimmune system">Neuroimmune system</a></li> <li><a href="/wiki/Neuromanagement" title="Neuromanagement">Neuromanagement</a></li> <li><a href="/wiki/Neuromodulation" title="Neuromodulation">Neuromodulation</a></li> <li><a href="/wiki/Neuroplasticity" title="Neuroplasticity">Neuroplasticity</a></li> <li><a href="/wiki/Neurotechnology" title="Neurotechnology">Neurotechnology</a></li> <li><a href="/wiki/Neurotoxin" title="Neurotoxin">Neurotoxin</a></li> <li><a href="/wiki/Neural_basis_of_self" title="Neural basis of self">Self-awareness</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="3"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Neuroscience" title="Category:Neuroscience">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Neuroscience" class="extiw" title="commons:Category:Neuroscience">Commons</a></b></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_psychology" 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_psychology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/History_of_evolutionary_psychology" title="History of evolutionary psychology">History</a> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li></ul></li> <li><a href="/wiki/Theoretical_foundations_of_evolutionary_psychology" title="Theoretical foundations of evolutionary psychology">Theoretical foundations</a> <ul><li><a href="/wiki/Adaptationism" title="Adaptationism">Adaptationism</a></li> <li><a href="/wiki/Cognitive_revolution" title="Cognitive revolution">Cognitive revolution</a></li> <li><a href="/wiki/Cognitivism_(psychology)" title="Cognitivism (psychology)">Cognitivism</a></li> <li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene selection theory</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li></ul></li> <li><a href="/wiki/Criticism_of_evolutionary_psychology" title="Criticism of evolutionary psychology">Criticism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;"><a href="/wiki/Human_evolution" title="Human evolution">Evolutionary<br />processes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Psychological_adaptation" title="Psychological adaptation">Adaptations</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Hamiltonian_spite" title="Hamiltonian spite">Hamiltonian spite</a></li> <li><a href="/wiki/Reciprocal_altruism_in_humans" title="Reciprocal altruism in humans">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">By-products</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness_in_humans" title="Inclusive fitness in humans">Inclusive</a></li></ul></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a> <ul><li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Sexual_selection_in_humans" title="Sexual selection in humans">Sexual selection</a> <ul><li><a href="/wiki/Costly_signaling_theory_in_evolutionary_psychology" title="Costly signaling theory in evolutionary psychology">Costly signaling</a></li> <li><a href="/wiki/Male_intrasexual_competition" title="Male intrasexual competition">Male</a>/<a href="/wiki/Female_intrasexual_competition" title="Female intrasexual competition">female intrasexual competition</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Areas</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution_of_cognition" title="Evolution of cognition">Cognition</a> /<br /><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Affect_(psychology)" title="Affect (psychology)">Affect</a> <ul><li><a href="/wiki/Affect_display" title="Affect display">Display</a></li> <li><a href="/wiki/Display_rules" title="Display rules">Display rules</a></li> <li><a href="/wiki/Facial_expression" title="Facial expression">Facial expression</a></li></ul></li> <li><a href="/wiki/Behavioral_modernity" title="Behavioral modernity">Behavioral modernity</a></li> <li><a href="/wiki/Cognitive_module" title="Cognitive module">Cognitive module</a>/<a href="/wiki/Modularity_of_mind" title="Modularity of mind">modularity of mind</a> <ul><li><a href="/wiki/Automatic_and_controlled_processes" title="Automatic and controlled processes">Automatic and controlled processes</a></li> <li><a href="/wiki/Computational_theory_of_mind" title="Computational theory of mind">Computational theory of mind</a></li> <li><a href="/wiki/Domain-general_learning" title="Domain-general learning">Domain generality</a></li> <li><a href="/wiki/Domain_specificity" title="Domain specificity">Domain specificity</a></li> <li><a href="/wiki/Dual_process_theory" title="Dual process theory">Dual process theory</a></li></ul></li> <li><a href="/wiki/Cognitive_tradeoff_hypothesis" title="Cognitive tradeoff hypothesis">Cognitive tradeoff hypothesis</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">Evolution of the brain</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">Evolution of nervous systems</a></li> <li><a href="/wiki/Fight-or-flight_response" title="Fight-or-flight response">Fight-or-flight response</a> <ul><li><a href="/wiki/Arachnophobia" title="Arachnophobia">Arachnophobia</a></li> <li><a href="/wiki/Fear_of_falling" title="Fear of falling">Basophobia</a></li> <li><a href="/wiki/Ophidiophobia" title="Ophidiophobia">Ophidiophobia</a></li></ul></li> <li><a href="/wiki/Folk_biology" title="Folk biology">Folk biology</a>/<a href="/wiki/Folk_taxonomy" title="Folk taxonomy">taxonomy</a></li> <li><a href="/wiki/Folk_psychology" title="Folk psychology">Folk psychology</a>/<a href="/wiki/Theory_of_mind" title="Theory of mind">theory of mind</a></li> <li><a href="/wiki/Evolution_of_human_intelligence" title="Evolution of human intelligence">Intelligence</a> <ul><li><a href="/wiki/Flynn_effect" title="Flynn effect">Flynn effect</a></li> <li><a href="/wiki/Wason_selection_task" title="Wason selection task">Wason selection task</a></li></ul></li> <li><a class="mw-selflink selflink">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_impacts_of_cars" title="Societal impacts of cars">Societal impacts of cars</a> <ul><li><a href="/wiki/Distracted_driving" title="Distracted driving">Distracted driving</a></li> <li><a href="/wiki/Lead%E2%80%93crime_hypothesis" title="Lead–crime hypothesis">Lead–crime hypothesis</a></li> <li><a href="/wiki/Mobile_phones_and_driving_safety" title="Mobile phones and driving safety">Mobile phones and driving safety</a></li> <li><a href="/wiki/Texting_while_driving" title="Texting while driving">Texting while driving</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sexology" title="Sexology">Sex</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Human_sexual_activity" title="Human sexual activity">Activity</a></li> <li><a href="/wiki/Attachment_in_adults" title="Attachment in adults">Adult attachment</a></li> <li><a href="/wiki/Age_disparity_in_sexual_relationships" title="Age disparity in sexual relationships">Age disparity</a></li> <li><a href="/wiki/Sexual_arousal" title="Sexual arousal">Arousal</a></li> <li><a href="/wiki/Concealed_ovulation" title="Concealed ovulation">Concealed ovulation</a></li> <li><a href="/wiki/Coolidge_effect" title="Coolidge effect">Coolidge effect</a></li> <li><a href="/wiki/Sexual_desire" title="Sexual desire">Desire</a></li> <li><a href="/wiki/Sexual_fantasy" title="Sexual fantasy">Fantasy</a></li> <li><a href="/wiki/Effects_of_hormones_on_sexual_motivation" title="Effects of hormones on sexual motivation">Hormonal motivation</a></li> <li><a href="/wiki/Sexual_jealousy" title="Sexual jealousy">Jealousy</a></li> <li><a href="/wiki/Mate_guarding_in_humans" title="Mate guarding in humans">Mate guarding</a></li> <li><a href="/wiki/Mating_preferences" title="Mating preferences">Mating preferences</a></li> <li><a href="/wiki/Human_mating_strategies" title="Human mating strategies">Mating strategies</a></li> <li><a href="/wiki/Biology_and_sexual_orientation" title="Biology and sexual orientation">Orientation</a></li> <li><a href="/wiki/Ovulatory_shift_hypothesis" title="Ovulatory shift hypothesis">Ovulatory shift hypothesis</a></li> <li><a href="/wiki/Pair_bond" title="Pair bond">Pair bond</a></li> <li><a href="/wiki/Physical_attractiveness" title="Physical attractiveness">Physical</a>/<a href="/wiki/Sexual_attraction" title="Sexual attraction">Sexual attraction</a></li> <li><a href="/wiki/Human_sexuality" title="Human sexuality">Sexuality</a>/<a href="/wiki/Human_male_sexuality" title="Human male sexuality">male</a>/<a href="/wiki/Human_female_sexuality" title="Human female sexuality">female</a></li> <li><a href="/wiki/Sexy_son_hypothesis" title="Sexy son hypothesis">Sexy son hypothesis</a></li> <li><a href="/wiki/Westermarck_effect" title="Westermarck effect">Westermarck effect</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Sex_differences_in_psychology" title="Sex differences in psychology">Sex differences</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aggression" title="Aggression">Aggression</a></li> <li><a href="/wiki/Sex_and_gender_differences_in_autism" title="Sex and gender differences in autism">Autism</a></li> <li><a href="/wiki/Sex_differences_in_cognition" title="Sex differences in cognition">Cognition</a></li> <li><a href="/wiki/Sex_differences_in_crime" title="Sex differences in crime">Crime</a></li> <li><a href="/wiki/Sexual_division_of_labour" title="Sexual division of labour">Division of labour</a></li> <li><a href="/wiki/Sex_differences_in_emotional_intelligence" title="Sex differences in emotional intelligence">Emotional intelligence</a></li> <li><a href="/wiki/Empathising%E2%80%93systemising_theory" title="Empathising–systemising theory">Empathising–systemising theory</a></li> <li><a href="/wiki/Gender_role" title="Gender role">Gender role</a></li> <li><a href="/wiki/Sex_differences_in_intelligence" title="Sex differences in intelligence">Intelligence</a></li> <li><a href="/wiki/Sex_differences_in_memory" title="Sex differences in memory">Memory</a></li> <li><a href="/wiki/Mental_disorders_and_gender" title="Mental disorders and gender">Mental health</a></li> <li><a href="/wiki/Sex_differences_in_narcissism" title="Sex differences in narcissism">Narcissism</a></li> <li><a href="/wiki/Neuroscience_of_sex_differences" title="Neuroscience of sex differences">Neuroscience</a></li> <li><a href="/wiki/Sex_differences_in_schizophrenia" title="Sex differences in schizophrenia">Schizophrenia</a></li> <li><a href="/wiki/Substance_abuse" title="Substance abuse">Substance abuse</a></li> <li><a href="/wiki/Gender_differences_in_suicide" title="Gender differences in suicide">Suicide</a></li> <li><a href="/wiki/Variability_hypothesis" title="Variability hypothesis">Variability hypothesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align:left;">Related subjects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Academic disciplines</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioral_economics" title="Behavioral economics">Behavioral</a>/<a href="/wiki/Evolutionary_economics" title="Evolutionary economics">evolutionary economics</a></li> <li><a href="/wiki/Behavioral_epigenetics" title="Behavioral epigenetics">Behavioral epigenetics</a>/<a href="/wiki/Behavioural_genetics" title="Behavioural genetics">genetics</a></li> <li><a href="/wiki/Affective_neuroscience" title="Affective neuroscience">Affective</a>/<a href="/wiki/Behavioral_neuroscience" title="Behavioral neuroscience">behavioral</a>/<a href="/wiki/Cognitive_neuroscience" title="Cognitive neuroscience">cognitive</a>/<a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">evolutionary neuroscience</a></li> <li><a href="/wiki/Biocultural_anthropology" title="Biocultural anthropology">Biocultural anthropology</a></li> <li><a href="/wiki/Biological_psychiatry" title="Biological psychiatry">Biological psychiatry</a></li> <li><a href="/wiki/Cognitive_psychology" title="Cognitive psychology">Cognitive psychology</a></li> <li><a href="/wiki/Cognitive_science" title="Cognitive science">Cognitive science</a></li> <li><a href="/wiki/Cross-cultural_psychology" title="Cross-cultural psychology">Cross-cultural psychology</a></li> <li><a href="/wiki/Ethology" title="Ethology">Ethology</a></li> <li><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Functional_psychology" title="Functional psychology">Functional psychology</a></li> <li><a href="/wiki/Neuropsychology" title="Neuropsychology">Neuropsychology</a></li> <li><a href="/wiki/Philosophy_of_mind" title="Philosophy of mind">Philosophy of mind</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Primatology" title="Primatology">Primatology</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Research topics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Great_ape_language" title="Great ape language">Great ape language</a></li> <li><a href="/wiki/Human%E2%80%93animal_communication" title="Human–animal communication">Human–animal communication</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Primate_cognition" title="Primate cognition">Primate cognition</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a></li> <li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li> <li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theoretical positions</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cultural_selection_theory" title="Cultural selection theory">Cultural selection theory</a></li> <li><a href="/wiki/Determinism" title="Determinism">Determinism</a>/<a href="/wiki/Indeterminism" title="Indeterminism">indeterminism</a> <ul><li><a href="/wiki/Biological_determinism" title="Biological determinism">Biological determinism</a></li> <li><a href="/wiki/Connectionism" title="Connectionism">Connectionism</a></li> <li><a href="/wiki/Cultural_determinism" title="Cultural determinism">Cultural determinism</a></li> <li><a href="/wiki/Environmental_determinism" title="Environmental determinism">Environmental determinism</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature versus nurture</a></li> <li><a href="/wiki/Psychological_nativism" title="Psychological nativism">Psychological nativism</a></li> <li><a href="/wiki/Social_constructionism" title="Social constructionism">Social constructionism</a></li> <li><a href="/wiki/Social_determinism" title="Social determinism">Social determinism</a></li> <li><a href="/wiki/Standard_social_science_model" title="Standard social science model">Standard social science model</a></li></ul></li> <li><a href="/wiki/Functionalism_(philosophy_of_mind)" title="Functionalism (philosophy of mind)">Functionalism</a></li> <li><a href="/wiki/Memetics" title="Memetics">Memetics</a></li> <li><a href="/wiki/Multilineal_evolution" title="Multilineal evolution">Multilineal evolution</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Neoevolutionism" title="Neoevolutionism">Neoevolutionism</a></li> <li><a href="/wiki/Sociocultural_evolution" title="Sociocultural evolution">Sociocultural evolution</a></li> <li><a href="/wiki/Unilineal_evolution" title="Unilineal evolution">Unilineal evolution</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><b><a href="/wiki/List_of_evolutionary_psychologists" title="List of evolutionary psychologists">Evolutionary psychologists</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" 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