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Patterning subsection</span> </button> <ul id="toc-Patterning-sublist" class="vector-toc-list"> <li id="toc-Dorsoventral_axis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dorsoventral_axis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Dorsoventral axis</span> </div> </a> <ul id="toc-Dorsoventral_axis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Rostrocaudal_(Anteroposterior)_axis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Rostrocaudal_(Anteroposterior)_axis"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Rostrocaudal (Anteroposterior) axis</span> </div> </a> <ul id="toc-Rostrocaudal_(Anteroposterior)_axis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Neurogenesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Neurogenesis"> <div 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id="toc-Radial-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tangential" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tangential"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.2</span> <span>Tangential</span> </div> </a> <ul id="toc-Tangential-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Axophilic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Axophilic"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.3</span> <span>Axophilic</span> </div> </a> <ul id="toc-Axophilic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Multipolar" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Multipolar"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.4</span> <span>Multipolar</span> </div> </a> <ul id="toc-Multipolar-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Neurotrophic_factors" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Neurotrophic_factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Neurotrophic factors</span> </div> </a> <ul id="toc-Neurotrophic_factors-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Synapse_formation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Synapse_formation"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Synapse formation</span> </div> </a> <button aria-controls="toc-Synapse_formation-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 Synapse formation subsection</span> </button> <ul id="toc-Synapse_formation-sublist" class="vector-toc-list"> <li id="toc-Neuromuscular_junction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Neuromuscular_junction"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Neuromuscular junction</span> </div> </a> <ul id="toc-Neuromuscular_junction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-CNS_synapses" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#CNS_synapses"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>CNS synapses</span> </div> </a> <ul id="toc-CNS_synapses-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Assembly_of_neural_circuits" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Assembly_of_neural_circuits"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.3</span> <span>Assembly of neural circuits</span> </div> </a> <ul id="toc-Assembly_of_neural_circuits-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Synapse_elimination" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Synapse_elimination"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Synapse elimination</span> </div> </a> <ul id="toc-Synapse_elimination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mapping" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Mapping"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</span> <span>Mapping</span> </div> </a> <ul id="toc-Mapping-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Adult_neurogenesis" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Adult_neurogenesis"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Adult neurogenesis</span> </div> </a> <ul id="toc-Adult_neurogenesis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">14</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> 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href="https://da.wikipedia.org/wiki/Neural_udvikling" title="Neural udvikling – Danish" lang="da" hreflang="da" data-title="Neural udvikling" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Desarrollo_neural" title="Desarrollo neural – Spanish" lang="es" hreflang="es" data-title="Desarrollo neural" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AA%DA%A9%D9%88%DB%8C%D9%86_%D8%AF%D8%B3%D8%AA%DA%AF%D8%A7%D9%87_%D8%B9%D8%B5%D8%A8%DB%8C" title="تکوین دستگاه عصبی – Persian" lang="fa" hreflang="fa" data-title="تکوین دستگاه عصبی" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Neurod%C3%A9veloppement" title="Neurodéveloppement – French" lang="fr" hreflang="fr" data-title="Neurodéveloppement" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Forbairt_inchinne_sa_leanbh" title="Forbairt inchinne sa leanbh – Irish" lang="ga" hreflang="ga" data-title="Forbairt inchinne sa leanbh" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Neurosviluppo" title="Neurosviluppo – Italian" lang="it" hreflang="it" data-title="Neurosviluppo" data-language-autonym="Italiano" 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a{color:var(--color-progressive)!important}}@media print{body.ns-0 .mw-parser-output .sidebar{display:none!important}}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><table class="sidebar nomobile nowraplinks" style="border:1px solid blue;background:misty-rose;color:inherit;"><tbody><tr><td class="sidebar-pretitle">This article is part of a series on the</td></tr><tr><th class="sidebar-title-with-pretitle">Development of <a href="/wiki/Organ_system" title="Organ system">organ systems</a></th></tr><tr><td class="sidebar-content"> <div class="hlist"><ul><li><a class="mw-selflink selflink">Nervous system</a></li><li><a href="/wiki/Development_of_the_digestive_system" title="Development of the digestive system">Digestive system</a></li><li><a href="/wiki/Development_of_the_reproductive_system" title="Development of the reproductive system">Reproductive system</a></li><li><a href="/wiki/Development_of_the_urinary_system" title="Development of the urinary system">Urinary system</a></li><li><a href="/wiki/Development_of_the_endocrine_system" title="Development of the endocrine system">Endocrine system</a></li><li><a href="/wiki/Development_of_the_human_body" title="Development of the human body">Human development</a></li><li><a href="/wiki/Fetal_circulation" title="Fetal circulation">Circulatory system</a></li></ul></div></td> </tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Organ_system_development" title="Template:Organ system development"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Organ_system_development" title="Template talk:Organ system development"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Organ_system_development" title="Special:EditPage/Template:Organ system development"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p>The <b>development of the nervous system</b>, or <b>neural development</b> (<b>neurodevelopment</b>), refers to the processes that generate, shape, and reshape the <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a> of animals, from the earliest stages of <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a> to adulthood. The field of neural development draws on both <a href="/wiki/Neuroscience" title="Neuroscience">neuroscience</a> and <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a> to describe and provide insight into the cellular and molecular mechanisms by which complex nervous systems develop, from <a href="/wiki/Nematode" title="Nematode">nematodes</a> and <a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">fruit flies</a> to <a href="/wiki/Mammals" class="mw-redirect" title="Mammals">mammals</a>. </p><p>Defects in neural development can lead to malformations such as <a href="/wiki/Holoprosencephaly" title="Holoprosencephaly">holoprosencephaly</a>, and a wide variety of <a href="/wiki/Neurological_disorder" title="Neurological disorder">neurological disorders</a> including <a href="/wiki/Paresis" title="Paresis">limb paresis</a> and <a href="/wiki/Paralysis" title="Paralysis">paralysis</a>, balance and vision disorders, and <a href="/wiki/Seizure" title="Seizure">seizures</a>,<sup id="cite_ref-Vet_1-0" class="reference"><a href="#cite_note-Vet-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> and in <a href="/wiki/Human" title="Human">humans</a> other disorders such as <a href="/wiki/Rett_syndrome" title="Rett syndrome">Rett syndrome</a>, <a href="/wiki/Down_syndrome" title="Down syndrome">Down syndrome</a> and <a href="/wiki/Intellectual_disability" title="Intellectual disability">intellectual disability</a>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Vertebrate_brain_development">Vertebrate brain development</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=1" title="Edit section: Vertebrate brain development"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:NSdiagram.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/NSdiagram.svg/480px-NSdiagram.svg.png" decoding="async" width="480" height="383" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/55/NSdiagram.svg/720px-NSdiagram.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/55/NSdiagram.svg/960px-NSdiagram.svg.png 2x" data-file-width="512" data-file-height="408" /></a><figcaption>Diagram of the vertebrate nervous system</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Development_of_the_nervous_system_in_humans" title="Development of the nervous system in humans">Development of the nervous system in humans</a></div> <p>The <a href="/wiki/Vertebrate" title="Vertebrate">vertebrate</a> <a href="/wiki/Central_nervous_system" title="Central nervous system">central nervous system</a> (CNS) is derived from the <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a>—the outermost <a href="/wiki/Germ_layer" title="Germ layer">germ layer</a> of the embryo. A part of the dorsal ectoderm becomes specified to neural ectoderm – <a href="/wiki/Neuroectoderm" title="Neuroectoderm">neuroectoderm</a> that forms the <a href="/wiki/Neural_plate" title="Neural plate">neural plate</a> along the dorsal side of the embryo.<sup id="cite_ref-Gilbert_3-0" class="reference"><a href="#cite_note-Gilbert-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> This is a part of the early patterning of the embryo (including the invertebrate embryo) that also establishes an anterior-posterior axis.<sup id="cite_ref-Wolpert1_5-0" class="reference"><a href="#cite_note-Wolpert1-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEWolpert2015522–526_6-0" class="reference"><a href="#cite_note-FOOTNOTEWolpert2015522–526-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The neural plate is the source of the majority of neurons and glial cells of the CNS. The <a href="/wiki/Neural_groove" title="Neural groove">neural groove</a> forms along the long axis of the neural plate, and the neural plate folds to give rise to the <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>.<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> This process is known as <a href="/wiki/Neurulation" title="Neurulation">neurulation</a>.<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> When the tube is closed at both ends it is filled with embryonic cerebrospinal fluid.<sup id="cite_ref-Gato_9-0" class="reference"><a href="#cite_note-Gato-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> As the embryo develops, the anterior part of the neural tube expands and forms three <a href="/wiki/Primary_brain_vesicles" class="mw-redirect" title="Primary brain vesicles">primary brain vesicles</a>, which become the <a href="/wiki/Forebrain" title="Forebrain">forebrain</a> (<a href="/wiki/Prosencephalon" class="mw-redirect" title="Prosencephalon">prosencephalon</a>), <a href="/wiki/Midbrain" title="Midbrain">midbrain</a> (<a href="/wiki/Mesencephalon" class="mw-redirect" title="Mesencephalon">mesencephalon</a>), and <a href="/wiki/Hindbrain" title="Hindbrain">hindbrain</a> (<a href="/wiki/Rhombencephalon" class="mw-redirect" title="Rhombencephalon">rhombencephalon</a>). These simple, early vesicles enlarge and further divide into the <a href="/wiki/Telencephalon" class="mw-redirect" title="Telencephalon">telencephalon</a> (future <a href="/wiki/Cerebral_cortex" title="Cerebral cortex">cerebral cortex</a> and <a href="/wiki/Basal_ganglia" title="Basal ganglia">basal ganglia</a>), <a href="/wiki/Diencephalon" title="Diencephalon">diencephalon</a> (future <a href="/wiki/Thalamus" title="Thalamus">thalamus</a> and <a href="/wiki/Hypothalamus" title="Hypothalamus">hypothalamus</a>), <a href="/wiki/Mesencephalon" class="mw-redirect" title="Mesencephalon">mesencephalon</a> (future <a href="/wiki/Inferior_colliculus" title="Inferior colliculus">colliculi</a>), <a href="/wiki/Metencephalon" title="Metencephalon">metencephalon</a> (future <a href="/wiki/Pons" title="Pons">pons</a> and <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a>), and <a href="/wiki/Myelencephalon" title="Myelencephalon">myelencephalon</a> (future <a href="/wiki/Medulla_oblongata" title="Medulla oblongata">medulla</a>).<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The CSF-filled central chamber is continuous from the telencephalon to the <a href="/wiki/Central_canal" title="Central canal">central canal</a> of the <a href="/wiki/Spinal_cord" title="Spinal cord">spinal cord</a>, and constitutes the developing <a href="/wiki/Ventricular_system" title="Ventricular system">ventricular system</a> of the CNS. Embryonic <a href="/wiki/Cerebrospinal_fluid" title="Cerebrospinal fluid">cerebrospinal fluid</a> differs from that formed in later developmental stages, and from adult CSF; it influences the behavior of neural precursors.<sup id="cite_ref-Gato_9-1" class="reference"><a href="#cite_note-Gato-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Because the neural tube gives rise to the brain and spinal cord any mutations at this stage in development can lead to fatal deformities like <a href="/wiki/Anencephaly" title="Anencephaly">anencephaly</a> or lifelong disabilities like <a href="/wiki/Spina_bifida" title="Spina bifida">spina bifida</a>. During this time, the walls of the neural tube contain <a href="/wiki/Neural_stem_cells" class="mw-redirect" title="Neural stem cells">neural stem cells</a>, which drive brain growth as they divide many times. Gradually some of the cells stop dividing and differentiate into <a href="/wiki/Neurons" class="mw-redirect" title="Neurons">neurons</a> and <a href="/wiki/Glial_cells" class="mw-redirect" title="Glial cells">glial cells</a>, which are the main cellular components of the CNS.<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> The newly generated neurons <a href="/wiki/Cellular_migration" class="mw-redirect" title="Cellular migration">migrate</a> to different parts of the developing brain to self-organize into different brain structures. Once the neurons have reached their regional positions, they extend <a href="/wiki/Axons" class="mw-redirect" title="Axons">axons</a> and <a href="/wiki/Dendrites" class="mw-redirect" title="Dendrites">dendrites</a>, which allow them to communicate with other neurons via <a href="/wiki/Synapses" class="mw-redirect" title="Synapses">synapses</a>. Synaptic communication between neurons leads to the establishment of functional <a href="/wiki/Neural_circuit" title="Neural circuit">neural circuits</a> that mediate sensory and motor processing, and underlie behavior.<sup id="cite_ref-Principles_of_neural_science_12-0" class="reference"><a href="#cite_note-Principles_of_neural_science-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/wiki/File:Development_of_nervous_system.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Development_of_nervous_system.svg/750px-Development_of_nervous_system.svg.png" decoding="async" width="750" height="304" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/67/Development_of_nervous_system.svg/1125px-Development_of_nervous_system.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/67/Development_of_nervous_system.svg/1500px-Development_of_nervous_system.svg.png 2x" data-file-width="1850" data-file-height="750" /></a><figcaption>Flowchart of <a href="/wiki/Human_brain" title="Human brain">human brain</a> development</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Induction">Induction</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=2" title="Edit section: Induction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>During early <a href="/wiki/Embryonic_development" class="mw-redirect" title="Embryonic development">embryonic development</a> of the vertebrate, the dorsal ectoderm becomes specified to give rise to the <a href="/wiki/Epidermis" title="Epidermis">epidermis</a> and the nervous system; a part of the dorsal ectoderm becomes specified to <a href="/wiki/Neuroectoderm" title="Neuroectoderm">neural ectoderm</a> to form the <a href="/wiki/Neural_plate" title="Neural plate">neural plate</a> which gives rise to the nervous system.<sup id="cite_ref-Gilbert_3-1" class="reference"><a href="#cite_note-Gilbert-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEWolpert2015163_13-0" class="reference"><a href="#cite_note-FOOTNOTEWolpert2015163-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The conversion of undifferentiated ectoderm to neuroectoderm requires signals from the <a href="/wiki/Mesoderm" title="Mesoderm">mesoderm</a>. At the onset of gastrulation presumptive mesodermal cells move through the dorsal blastopore lip and form a layer of mesoderm in between the <a href="/wiki/Endoderm" title="Endoderm">endoderm</a> and the ectoderm. Mesodermal cells migrate along the dorsal midline to give rise to the <a href="/wiki/Notochord" title="Notochord">notochord</a> that develops into the <a href="/wiki/Vertebral_column" title="Vertebral column">vertebral column</a>. Neuroectoderm overlying the notochord develops into the neural plate in response to a diffusible signal produced by the notochord. The remainder of the ectoderm gives rise to the epidermis. The ability of the mesoderm to convert the overlying ectoderm into neural tissue is called <b>neural induction</b>. </p><p>In the early embryo, the neural plate folds outwards to form the <a href="/wiki/Neural_groove" title="Neural groove">neural groove</a>. Beginning in the future neck region, the <a href="/wiki/Neural_folds" class="mw-redirect" title="Neural folds">neural folds</a> of this groove close to create the <a href="/wiki/Neural_tube" title="Neural tube">neural tube</a>. The formation of the neural tube from the ectoderm is called <a href="/wiki/Neurulation" title="Neurulation">neurulation</a>. The ventral part of the neural tube is called the <a href="/wiki/Basal_plate_(neural_tube)" title="Basal plate (neural tube)">basal plate</a>; the dorsal part is called the <a href="/wiki/Alar_plate" title="Alar plate">alar plate</a>. The hollow interior is called the <a href="/wiki/Neural_canal" class="mw-redirect" title="Neural canal">neural canal</a>, and the open ends of the neural tube, called the neuropores, close off.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p><p>A transplanted blastopore lip can convert ectoderm into neural tissue and is said to have an inductive effect. Neural inducers are molecules that can induce the expression of neural genes in ectoderm <a href="/wiki/Explant_culture" title="Explant culture">explants</a> without inducing mesodermal genes as well. Neural induction is often studied in <i><a href="/wiki/Xenopus" title="Xenopus">Xenopus</a></i> embryos since they have a simple <a href="/wiki/Body_plan" title="Body plan">body plan</a> and there are good markers to distinguish between neural and non-neural tissue. Examples of neural inducers are the molecules <a href="/wiki/Noggin_(protein)" title="Noggin (protein)">noggin</a> and <a href="/wiki/Chordin" title="Chordin">chordin</a>. </p><p>When embryonic ectodermal cells are cultured at low density in the absence of mesodermal cells they undergo neural differentiation (express neural genes), suggesting that neural differentiation is the default fate of ectodermal cells. In <a href="/wiki/Explant_culture" title="Explant culture">explant cultures</a> (which allow direct cell-cell interactions) the same cells differentiate into epidermis. This is due to the action of <a href="/wiki/BMP4" class="mw-redirect" title="BMP4">BMP4</a> (a <a href="/wiki/TGF_beta" class="mw-redirect" title="TGF beta">TGF-β</a> family protein) that induces ectodermal cultures to differentiate into epidermis. During neural induction, noggin and chordin are produced by the dorsal mesoderm (notochord) and diffuse into the overlying ectoderm to inhibit the activity of BMP4. This inhibition of BMP4 causes the cells to differentiate into neural cells. Inhibition of TGF-β and BMP (bone morphogenetic protein) signaling can efficiently induce neural tissue from <a href="/wiki/Pluripotent_stem_cells" class="mw-redirect" title="Pluripotent stem cells">pluripotent stem cells</a>.<sup id="cite_ref-Chambers_2009_15-0" class="reference"><a href="#cite_note-Chambers_2009-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Regionalization">Regionalization</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=3" title="Edit section: Regionalization"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In a later stage of development the superior part of the neural tube flexes at the level of the future midbrain—the <a href="/wiki/Mesencephalon" class="mw-redirect" title="Mesencephalon">mesencephalon</a>, at the mesencephalic flexure or <a href="/wiki/Cephalic_flexure" class="mw-redirect" title="Cephalic flexure">cephalic flexure</a>. Above the <a href="/wiki/Mesencephalon" class="mw-redirect" title="Mesencephalon">mesencephalon</a> is the <a href="/wiki/Prosencephalon" class="mw-redirect" title="Prosencephalon">prosencephalon</a> (future forebrain) and beneath it is the <a href="/wiki/Rhombencephalon" class="mw-redirect" title="Rhombencephalon">rhombencephalon</a> (future hindbrain). </p><p>The alar plate of the prosencephalon expands to form the <a href="/wiki/Telencephalon" class="mw-redirect" title="Telencephalon">telencephalon</a> which gives rise to the <a href="/wiki/Cerebral_hemisphere" title="Cerebral hemisphere">cerebral hemispheres</a>, whilst its basal plate becomes the <a href="/wiki/Diencephalon" title="Diencephalon">diencephalon</a>. The <a href="/wiki/Optical_vesicle" class="mw-redirect" title="Optical vesicle">optical vesicle</a> (which eventually become the optic nerve, retina and iris) forms at the basal plate of the prosencephalon. </p> <div class="mw-heading mw-heading2"><h2 id="Patterning">Patterning</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=4" title="Edit section: Patterning"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In <a href="/wiki/Chordates" class="mw-redirect" title="Chordates">chordates</a>, dorsal ectoderm forms all neural tissue and the nervous system. <a href="/wiki/Pattern_formation#Biology" title="Pattern formation">Patterning</a> occurs due to specific environmental conditions - different concentrations of signaling molecules </p> <div class="mw-heading mw-heading3"><h3 id="Dorsoventral_axis">Dorsoventral axis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=5" title="Edit section: Dorsoventral axis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ventral half of the <a href="/wiki/Neural_plate" title="Neural plate">neural plate</a> is controlled by the <a href="/wiki/Notochord" title="Notochord">notochord</a>, which acts as the 'organiser'. The dorsal half is controlled by the <a href="/wiki/Ectoderm" title="Ectoderm">ectoderm</a> plate, which flanks either side of the neural plate.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p><p>Ectoderm follows a default pathway to become neural tissue. Evidence for this comes from single, cultured cells of ectoderm, which go on to form neural tissue. This is postulated to be because of a lack of <a href="/wiki/Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMPs</a>, which are blocked by the organiser. The organiser may produce molecules such as <a href="/wiki/Follistatin" title="Follistatin">follistatin</a>, <a href="/wiki/Noggin_(protein)" title="Noggin (protein)">noggin</a> and <a href="/wiki/Chordin" title="Chordin">chordin</a> that inhibit BMPs. </p><p>The ventral neural tube is patterned by <a href="/wiki/Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">sonic hedgehog</a> (Shh) from the notochord, which acts as the inducing tissue. Notochord-derived Shh signals to the <a href="/wiki/Floor_plate_(biology)" title="Floor plate (biology)">floor plate</a>, and induces Shh expression in the floor plate. Floor plate-derived Shh subsequently signals to other cells in the neural tube, and is essential for proper specification of ventral neuron progenitor domains. Loss of Shh from the notochord and/or floor plate prevents proper specification of these progenitor domains. Shh binds <a href="/wiki/Patched" title="Patched">Patched1</a>, relieving Patched-mediated inhibition of <a href="/wiki/Smoothened" title="Smoothened">Smoothened</a>, leading to activation of the Gli family of <a href="/wiki/Transcription_factor" title="Transcription factor">transcription factors</a> (<a href="/wiki/GLI1" title="GLI1">GLI1</a>, <a href="/wiki/GLI2" title="GLI2">GLI2</a>, and <a href="/wiki/GLI3" title="GLI3">GLI3</a>). </p><p>In this context Shh acts as a <a href="/wiki/Morphogen" title="Morphogen">morphogen</a> - it induces cell differentiation dependent on its concentration. At low concentrations it forms ventral <a href="/wiki/Interneuron" title="Interneuron">interneurons</a>, at higher concentrations it induces <a href="/wiki/Motor_neuron" title="Motor neuron">motor neuron</a> development, and at highest concentrations it induces floor plate differentiation. Failure of Shh-modulated differentiation causes <a href="/wiki/Holoprosencephaly" title="Holoprosencephaly">holoprosencephaly</a>. </p><p>The dorsal neural tube is patterned by BMPs from the epidermal ectoderm flanking the neural plate. These induce sensory interneurons by activating <a href="/wiki/Serine/threonine_protein_kinase" class="mw-redirect" title="Serine/threonine protein kinase">Sr/Thr kinases</a> and altering <a href="/wiki/SMAD_(protein)" title="SMAD (protein)">SMAD</a> transcription factor levels. </p> <div class="mw-heading mw-heading3"><h3 id="Rostrocaudal_(Anteroposterior)_axis"><span id="Rostrocaudal_.28Anteroposterior.29_axis"></span>Rostrocaudal (Anteroposterior) axis</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=6" title="Edit section: Rostrocaudal (Anteroposterior) axis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Signals that control anteroposterior neural development include <a href="/wiki/Fibroblast_growth_factor" title="Fibroblast growth factor">FGF</a> and <a href="/wiki/Retinoic_acid" title="Retinoic acid">retinoic acid</a>, which act in the hindbrain and spinal cord.<sup id="cite_ref-Duester_17-0" class="reference"><a href="#cite_note-Duester-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> The hindbrain, for example, is patterned by <a href="/wiki/Hox_genes" class="mw-redirect" title="Hox genes">Hox genes</a>, which are expressed in overlapping domains along the anteroposterior axis under the control of retinoic acid. The <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">3<span class="nowrap" style="padding-left:0.15em;">′</span></a> (3 prime end) genes in the Hox cluster are induced by retinoic acid in the hindbrain, whereas the <a href="/wiki/Directionality_(molecular_biology)" title="Directionality (molecular biology)">5<span class="nowrap" style="padding-left:0.15em;">′</span></a> (5 prime end) Hox genes are not induced by retinoic acid and are expressed more posteriorly in the spinal cord. Hoxb-1 is expressed in rhombomere 4 and gives rise to the <a href="/wiki/Facial_nerve" title="Facial nerve">facial nerve</a>. Without this Hoxb-1 expression, a nerve similar to the <a href="/wiki/Trigeminal_nerve" title="Trigeminal nerve">trigeminal nerve</a> arises. </p> <div class="mw-heading mw-heading2"><h2 id="Neurogenesis">Neurogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=7" title="Edit section: Neurogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a> is the process by which neurons are generated from <a href="/wiki/Neural_stem_cells" class="mw-redirect" title="Neural stem cells">neural stem cells</a> and <a href="/wiki/Progenitor_cells" class="mw-redirect" title="Progenitor cells">progenitor cells</a>. Neurons are 'post-mitotic', meaning that they will never divide again for the lifetime of the organism.<sup id="cite_ref-Principles_of_neural_science_12-1" class="reference"><a href="#cite_note-Principles_of_neural_science-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetic modifications</a> play a key role in regulating <a href="/wiki/Gene_expression" title="Gene expression">gene expression</a> in differentiating <a href="/wiki/Neural_stem_cells" class="mw-redirect" title="Neural stem cells">neural stem cells</a> and are critical for cell fate determination in the developing and adult mammalian brain. Epigenetic modifications include <a href="/wiki/DNA_methylation" title="DNA methylation">DNA cytosine methylation</a> to form <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a> and <a href="/wiki/DNA_demethylation" title="DNA demethylation">5-methylcytosine demethylation</a>.<sup id="cite_ref-Wang2016_18-0" class="reference"><a href="#cite_note-Wang2016-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><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> <a href="/wiki/DNA_methylation" title="DNA methylation">DNA cytosine methylation</a> is catalyzed by <a href="/wiki/DNA_methyltransferase" title="DNA methyltransferase">DNA methyltransferases (DNMTs)</a>. Methylcytosine demethylation is catalyzed in several sequential steps by <a href="/wiki/TET_enzymes" title="TET enzymes">TET enzymes</a> that carry out oxidative reactions (e.g. <a href="/wiki/5-methylcytosine" class="mw-redirect" title="5-methylcytosine">5-methylcytosine</a> to <a href="/wiki/5-hydroxymethylcytosine" class="mw-redirect" title="5-hydroxymethylcytosine">5-hydroxymethylcytosine</a>) and enzymes of the DNA <a href="/wiki/Base_excision_repair" title="Base excision repair">base excision repair</a> (BER) pathway.<sup id="cite_ref-Wang2016_18-1" class="reference"><a href="#cite_note-Wang2016-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Neuronal_migration">Neuronal migration</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=8" title="Edit section: Neuronal migration"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Corticogenesis_in_a_wild-type_mouse.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Corticogenesis_in_a_wild-type_mouse.png/220px-Corticogenesis_in_a_wild-type_mouse.png" decoding="async" width="220" height="333" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Corticogenesis_in_a_wild-type_mouse.png/330px-Corticogenesis_in_a_wild-type_mouse.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Corticogenesis_in_a_wild-type_mouse.png/440px-Corticogenesis_in_a_wild-type_mouse.png 2x" data-file-width="570" data-file-height="864" /></a><figcaption><a href="/wiki/Corticogenesis" class="mw-redirect" title="Corticogenesis">Corticogenesis</a>: younger neurons migrate past older ones using <a href="/wiki/Radial_glia" class="mw-redirect" title="Radial glia">radial glia</a> as a scaffolding. <a href="/wiki/Cajal%E2%80%93Retzius_cell" title="Cajal–Retzius cell">Cajal–Retzius cells</a> (red) release <a href="/wiki/Reelin" title="Reelin">reelin</a> (orange).</figcaption></figure> <p>Neuronal <a href="/wiki/Cellular_migration" class="mw-redirect" title="Cellular migration">migration</a> is the method by which neurons travel from their origin or birthplace to their final position in the brain. There are several ways they can do this, e.g. by radial migration or tangential migration. Sequences of radial migration (also known as glial guidance) and somal translocation have been captured by <a href="/wiki/Time-lapse_microscopy" title="Time-lapse microscopy">time-lapse microscopy</a>.<sup id="cite_ref-Nadar1_20-0" class="reference"><a href="#cite_note-Nadar1-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png/220px-Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png" decoding="async" width="220" height="311" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png/330px-Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png 1.5x, //upload.wikimedia.org/wikipedia/commons/7/70/Interneuron-radial_glial_interactions_in_the_developing_cerebral_cortex.png 2x" data-file-width="425" data-file-height="600" /></a><figcaption>Tangential migration of interneurons from <a href="/wiki/Ganglionic_eminence" title="Ganglionic eminence">ganglionic eminence</a></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Radial">Radial</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=9" title="Edit section: Radial"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Neuronal precursor cells proliferate in the <a href="/wiki/Ventricular_zone" title="Ventricular zone">ventricular zone</a> of the developing <a href="/wiki/Neocortex" title="Neocortex">neocortex</a>, where the principal neural stem cell is the <a href="/wiki/Radial_glial_cell" title="Radial glial cell">radial glial cell</a>. The first <a href="/wiki/Mitosis" title="Mitosis">postmitotic</a> cells must leave the stem cell niche and migrate outward to form the preplate, which is destined to become <a href="/wiki/Cajal%E2%80%93Retzius_cell" title="Cajal–Retzius cell">Cajal–Retzius cells</a> and <a href="/wiki/Subplate" title="Subplate">subplate</a> neurons. These cells do so by somal translocation. Neurons migrating with this mode of locomotion are bipolar and attach the leading edge of the process to the <a href="/wiki/Pia_mater" title="Pia mater">pia</a>. The <a href="/wiki/Soma_(biology)" title="Soma (biology)">soma</a> is then transported to the pial surface by <a href="/w/index.php?title=Nucleokinesis&action=edit&redlink=1" class="new" title="Nucleokinesis (page does not exist)">nucleokinesis</a>, a process by which a <a href="/wiki/Microtubules" class="mw-redirect" title="Microtubules">microtubule</a> "cage" around the nucleus elongates and contracts in association with the <a href="/wiki/Centrosome" title="Centrosome">centrosome</a> to guide the nucleus to its final destination.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Radial_glial_cell" title="Radial glial cell">Radial glial cells</a>, whose fibers serve as a scaffolding for migrating cells and a means of radial communication mediated by calcium dynamic activity,<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> act as the main excitatory neuronal stem cell of the cerebral cortex<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid11535293_25-0" class="reference"><a href="#cite_note-pmid11535293-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> or translocate to the cortical plate and differentiate either into <a href="/wiki/Astrocyte" title="Astrocyte">astrocytes</a> or <a href="/wiki/Neuron" title="Neuron">neurons</a>.<sup id="cite_ref-pmid11567613_26-0" class="reference"><a href="#cite_note-pmid11567613-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Somal translocation can occur at any time during development.<sup id="cite_ref-Nadar1_20-1" class="reference"><a href="#cite_note-Nadar1-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p><p>Subsequent waves of neurons split the preplate by migrating along <a href="/wiki/Radial_glia" class="mw-redirect" title="Radial glia">radial glial</a> fibres to form the cortical plate. Each wave of migrating cells travel past their predecessors forming layers in an inside-out manner, meaning that the youngest neurons are the closest to the surface.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> It is estimated that glial guided migration represents 90% of migrating neurons in human and about 75% in rodents.<sup id="cite_ref-pmid12050665_29-0" class="reference"><a href="#cite_note-pmid12050665-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tangential">Tangential</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=10" title="Edit section: Tangential"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most interneurons migrate tangentially through multiple modes of migration to reach their appropriate location in the cortex. An example of tangential migration is the movement of interneurons from the <a href="/wiki/Ganglionic_eminence" title="Ganglionic eminence">ganglionic eminence</a> to the cerebral cortex. One example of ongoing tangential migration in a mature organism, observed in some animals, is the <a href="/wiki/Rostral_migratory_stream" title="Rostral migratory stream">rostral migratory stream</a> connecting <a href="/wiki/Subventricular_zone" title="Subventricular zone">subventricular zone</a> and <a href="/wiki/Olfactory_bulb" title="Olfactory bulb">olfactory bulb</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Axophilic">Axophilic</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=11" title="Edit section: Axophilic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many neurons migrating along the anterior-posterior axis of the body use existing <a href="/wiki/Axon" title="Axon">axon</a> tracts to migrate along; this is called axophilic migration. An example of this mode of migration is in <a href="/wiki/GnRH_Neuron" class="mw-redirect" title="GnRH Neuron">GnRH-expressing neurons</a>, which make a long journey from their birthplace in the nose, through the forebrain, and into the hypothalamus.<sup id="cite_ref-Wray10_30-0" class="reference"><a href="#cite_note-Wray10-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> Many of the mechanisms of this migration have been worked out, starting with the extracellular guidance cues<sup id="cite_ref-Giacobini07_31-0" class="reference"><a href="#cite_note-Giacobini07-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> that trigger intracellular signaling. These intracellular signals, such as <a href="/wiki/Calcium_signaling" title="Calcium signaling">calcium signaling</a>, lead to <a href="/wiki/Actin" title="Actin">actin</a><sup id="cite_ref-Hutchins13_32-0" class="reference"><a href="#cite_note-Hutchins13-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Microtubule" title="Microtubule">microtubule</a><sup id="cite_ref-Hutchins14_33-0" class="reference"><a href="#cite_note-Hutchins14-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Cytoskeleton" title="Cytoskeleton">cytoskeletal</a> dynamics, which produce cellular forces that interact with the extracellular environment through <a href="/wiki/Cell_adhesion_protein" class="mw-redirect" title="Cell adhesion protein">cell adhesion proteins</a><sup id="cite_ref-Parkash12_34-0" class="reference"><a href="#cite_note-Parkash12-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> to cause the movement of these cells. </p> <div class="mw-heading mw-heading3"><h3 id="Multipolar">Multipolar</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=12" title="Edit section: Multipolar"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There is also a method of neuronal migration called <b>multipolar migration</b>.<sup id="cite_ref-Tabata03_35-0" class="reference"><a href="#cite_note-Tabata03-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> This is seen in multipolar cells, which in the human, are abundantly present in the <a href="/wiki/Cortical_intermediate_zone" class="mw-redirect" title="Cortical intermediate zone">cortical intermediate zone</a>. They do not resemble the cells migrating by locomotion or somal translocation. Instead these multipolar cells express neuronal markers and extend multiple thin processes in various directions independently of the radial glial fibers.<sup id="cite_ref-Tabata03_35-1" class="reference"><a href="#cite_note-Tabata03-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Neurotrophic_factors">Neurotrophic factors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=13" title="Edit section: Neurotrophic factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The survival of neurons is regulated by survival factors, called trophic factors. The neurotrophic hypothesis was formulated by Victor Hamburger and <a href="/wiki/Rita_Levi_Montalcini" class="mw-redirect" title="Rita Levi Montalcini">Rita Levi Montalcini</a> based on studies of the developing nervous system. Victor Hamburger discovered that implanting an extra limb in the developing chick led to an increase in the number of spinal motor neurons. Initially he thought that the extra limb was inducing proliferation of motor neurons, but he and his colleagues later showed that there was a great deal of motor neuron death during normal development, and the extra limb prevented this cell death. According to the neurotrophic hypothesis, growing axons compete for limiting amounts of target-derived trophic factors and axons that fail to receive sufficient trophic support die by apoptosis. It is now clear that factors produced by a number of sources contribute to neuronal survival. </p> <ul><li><a href="/wiki/Nerve_Growth_Factor" class="mw-redirect" title="Nerve Growth Factor">Nerve Growth Factor</a> (NGF): Rita Levi Montalcini and Stanley Cohen purified the first trophic factor, Nerve Growth Factor (NGF), for which they received the Nobel Prize. There are three NGF-related trophic factors: BDNF, NT3, and NT4, which regulate survival of various neuronal populations. The Trk proteins act as receptors for NGF and related factors. Trk is a receptor tyrosine kinase. Trk dimerization and phosphorylation leads to activation of various intracellular signaling pathways including the MAP kinase, Akt, and PKC pathways.</li> <li>CNTF: Ciliary neurotrophic factor is another protein that acts as a survival factor for motor neurons. CNTF acts via a receptor complex that includes CNTFRα, GP130, and LIFRβ. Activation of the receptor leads to phosphorylation and recruitment of the JAK kinase, which in turn phosphorylates <a href="/wiki/LIFR" class="mw-redirect" title="LIFR">LIFR</a>β. LIFRβ acts as a docking site for the STAT transcription factors. JAK kinase phosphorylates STAT proteins, which dissociate from the receptor and translocate to the nucleus to regulate gene expression.</li> <li>GDNF: Glial derived neurotrophic factor is a member of the <a href="/wiki/TGFb" class="mw-redirect" title="TGFb">TGFb</a> family of proteins, and is a potent trophic factor for striatal neurons. The functional receptor is a heterodimer, composed of type 1 and type 2 receptors. Activation of the type 1 receptor leads to phosphorylation of Smad proteins, which translocate to the nucleus to activate gene expression.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Synapse_formation">Synapse formation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=14" title="Edit section: Synapse formation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Neuromuscular_junction">Neuromuscular junction</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=15" title="Edit section: Neuromuscular junction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Neuromuscular_junction" title="Neuromuscular junction">Neuromuscular junction</a></div> <p>Much of our understanding of synapse formation comes from studies at the neuromuscular junction. The transmitter at this synapse is acetylcholine. The acetylcholine receptor (AchR) is present at the surface of muscle cells before synapse formation. The arrival of the nerve induces clustering of the receptors at the synapse. McMahan and Sanes showed that the synaptogenic signal is concentrated at the <a href="/wiki/Basal_lamina" title="Basal lamina">basal lamina</a>. They also showed that the synaptogenic signal is produced by the nerve, and they identified the factor as <a href="/wiki/Agrin" title="Agrin">Agrin</a>. Agrin induces clustering of AchRs on the muscle surface and synapse formation is disrupted in agrin knockout mice. Agrin transduces the signal via MuSK receptor to <a href="/wiki/Rapsyn" class="mw-redirect" title="Rapsyn">rapsyn</a>. Fischbach and colleagues showed that receptor subunits are selectively transcribed from nuclei next to the synaptic site. This is mediated by neuregulins. </p><p>In the mature synapse each muscle fiber is innervated by one motor neuron. However, during development, many of the fibers are innervated by multiple axons. Lichtman and colleagues have studied the process of synapses elimination.<sup id="cite_ref-pmid22745601_37-0" class="reference"><a href="#cite_note-pmid22745601-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> This is an activity-dependent event. Partial blockage of the receptor leads to retraction of corresponding presynaptic terminals. Later they used a connectomic approach, i.e., tracing out all the connections between motor neurons and muscle fibers, to characterize developmental synapse elimination on the level of a full circuit. Analysis confirmed the massive rewiring, 10-fold decrease in the number of synapses, that takes place as axons prune their motor units but add more synaptic areas at the NMJs with which they remain in contact.<sup id="cite_ref-Meirovitch2021_38-0" class="reference"><a href="#cite_note-Meirovitch2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="CNS_synapses">CNS synapses</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=16" title="Edit section: CNS synapses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Agrin appears not to be a central mediator of CNS synapse formation and there is active interest in identifying signals that mediate CNS synaptogenesis. Neurons in culture develop synapses that are similar to those that form in vivo, suggesting that synaptogenic signals can function properly in vitro. CNS synaptogenesis studies have focused mainly on glutamatergic synapses. Imaging experiments show that dendrites are highly dynamic during development and often initiate contact with axons. This is followed by recruitment of postsynaptic proteins to the site of contact. Stephen Smith and colleagues have shown that contact initiated by <a href="/wiki/Dendritic_Filopodia" class="mw-redirect" title="Dendritic Filopodia">dendritic filopodia</a> can develop into synapses. </p><p>Induction of synapse formation by glial factors: Barres and colleagues made the observation that factors in glial conditioned media induce synapse formation in retinal ganglion cell cultures. Synapse formation in the CNS is correlated with astrocyte differentiation suggesting that astrocytes might provide a synaptogenic factor. The identity of the astrocytic factors is not yet known. </p><p><a href="/wiki/Neuroligin" title="Neuroligin">Neuroligins</a> and SynCAM as synaptogenic signals: Sudhof, Serafini, Scheiffele and colleagues have shown that neuroligins and SynCAM can act as factors that induce presynaptic differentiation. Neuroligins are concentrated at the postsynaptic site and act via neurexins concentrated in the presynaptic axons. SynCAM is a cell adhesion molecule that is present in both pre- and post-synaptic membranes. </p> <div class="mw-heading mw-heading3"><h3 id="Assembly_of_neural_circuits">Assembly of neural circuits</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=17" title="Edit section: Assembly of neural circuits"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Activity-dependent_plasticity" title="Activity-dependent plasticity">Activity-dependent plasticity</a></div> <p>The processes of <a href="/wiki/Neuronal_migration" class="mw-redirect" title="Neuronal migration">neuronal migration</a>, <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">differentiation</a> and <a href="/wiki/Axon_guidance" title="Axon guidance">axon guidance</a> are generally believed to be activity-independent mechanisms and rely on hard-wired genetic programs in the neurons themselves. Research findings however have implicated a role for <a href="/wiki/Activity-dependent_plasticity" title="Activity-dependent plasticity">activity-dependent mechanisms</a> in mediating some aspects of these processes such as the rate of neuronal migration,<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> aspects of neuronal differentiation<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> and axon pathfinding.<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> Activity-dependent mechanisms influence neural circuit development and are crucial for laying out early connectivity maps and the continued refinement of synapses which occurs during development.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> There are two distinct types of neural activity we observe in developing circuits -early spontaneous activity and sensory-evoked activity. Spontaneous activity occurs early during <a href="/wiki/Neural_circuit" title="Neural circuit">neural circuit</a> development even when sensory input is absent and is observed in many systems such as the developing <a href="/wiki/Visual_system" title="Visual system">visual system</a>,<sup id="cite_ref-43" class="reference"><a href="#cite_note-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Auditory_system" title="Auditory system">auditory system</a>,<sup id="cite_ref-45" class="reference"><a href="#cite_note-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Motor_system" title="Motor system">motor system</a>,<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Hippocampus" title="Hippocampus">hippocampus</a>,<sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> and <a href="/wiki/Neocortex" title="Neocortex">neocortex</a>.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> </p><p>Experimental techniques such as direct electrophysiological recording, fluorescence imaging using calcium indicators and optogenetic techniques have shed light on the nature and function of these early bursts of activity.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> They have distinct spatial and temporal patterns during development<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> and their ablation during development has been known to result in deficits in network refinement in the visual system.<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> In the immature <a href="/wiki/Retina" title="Retina">retina</a>, waves of spontaneous action potentials arise from the <a href="/wiki/Retinal_ganglion_cells" class="mw-redirect" title="Retinal ganglion cells">retinal ganglion cells</a> and sweep across the retinal surface in the first few postnatal weeks.<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> These waves are mediated by <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a> <a href="/wiki/Acetylcholine" title="Acetylcholine">acetylcholine</a> in the initial phase and later on by <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a>.<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> They are thought to instruct the formation of two sensory maps- the <a href="/wiki/Retinotopic" class="mw-redirect" title="Retinotopic">retinotopic</a> map and eye-specific segregation.<sup id="cite_ref-ReferenceA_57-0" class="reference"><a href="#cite_note-ReferenceA-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Retinotopic map refinement occurs in downstream visual targets in the brain-the <a href="/wiki/Superior_colliculus" title="Superior colliculus">superior colliculus</a> (SC) and dorsal <a href="/wiki/Lateral_geniculate_nucleus" title="Lateral geniculate nucleus">lateral geniculate nucleus</a> (LGN).<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> Pharmacological disruption and mouse models lacking the β2 subunit of the <a href="/wiki/Nicotinic_acetylcholine_receptor" title="Nicotinic acetylcholine receptor">nicotinic acetylcholine receptor</a> has shown that the lack of spontaneous activity leads to marked defects in <a href="/wiki/Retinotopy" title="Retinotopy">retinotopy</a> and eye-specific segregation.<sup id="cite_ref-ReferenceA_57-1" class="reference"><a href="#cite_note-ReferenceA-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>Recent studies confirm that <a href="/wiki/Microglia" title="Microglia">microglia</a>, the resident immune cell of the brain, establish direct contacts with the cell bodies of developing neurons, and through these connections, regulate neurogenesis, migration, integration and the formation of neuronal networks in an activity-dependent manner.<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> </p><p>In the developing <a href="/wiki/Auditory_system" title="Auditory system">auditory system</a>, developing <a href="/wiki/Cochlea" title="Cochlea">cochlea</a> generate bursts of activity which spreads across the <a href="/wiki/Inner_hair_cells" class="mw-redirect" title="Inner hair cells">inner hair cells</a> and <a href="/wiki/Spiral_ganglion" title="Spiral ganglion">spiral ganglion</a> neurons which relay auditory information to the brain.<sup id="cite_ref-ReferenceB_60-0" class="reference"><a href="#cite_note-ReferenceB-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">ATP</a> release from supporting cells triggers action potentials in <a href="/wiki/Inner_hair_cells" class="mw-redirect" title="Inner hair cells">inner hair cells</a>.<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> In the auditory system, spontaneous activity is thought to be involved in tonotopic map formation by segregating cochlear neuron axons tuned to high and low frequencies.<sup id="cite_ref-ReferenceB_60-1" class="reference"><a href="#cite_note-ReferenceB-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> In the motor system, periodic bursts of spontaneous activity are driven by excitatory <a href="/wiki/GABA" title="GABA">GABA</a> and <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> during the early stages and by <a href="/wiki/Acetylcholine" title="Acetylcholine">acetylcholine</a> and <a href="/wiki/Glutamate" class="mw-redirect" title="Glutamate">glutamate</a> at later stages.<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> In the developing <a href="/wiki/Zebrafish" title="Zebrafish">zebrafish</a> <a href="/wiki/Spinal_cord" title="Spinal cord">spinal cord</a>, early spontaneous activity is required for the formation of increasingly synchronous alternating bursts between ipsilateral and contralateral regions of the spinal cord and for the integration of new cells into the circuit.<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> Motor neurons innervating the same twitch muscle fibers are thought to maintain synchronous activity which allows both neurons to remain in contact with the muscle fiber in adulthood.<sup id="cite_ref-Meirovitch2021_38-1" class="reference"><a href="#cite_note-Meirovitch2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> In the <a href="/wiki/Cortex_(anatomy)" title="Cortex (anatomy)">cortex</a>, early waves of activity have been observed in the <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a> and cortical slices.<sup id="cite_ref-Elsevier_64-0" class="reference"><a href="#cite_note-Elsevier-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> Once sensory stimulus becomes available, final fine-tuning of sensory-coding maps and circuit refinement begins to rely more and more on sensory-evoked activity as demonstrated by classic experiments about the effects of sensory deprivation during <a href="/wiki/Critical_periods" class="mw-redirect" title="Critical periods">critical periods</a>.<sup id="cite_ref-Elsevier_64-1" class="reference"><a href="#cite_note-Elsevier-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p><p> Contemporary diffusion-weighted <a href="/wiki/MRI" class="mw-redirect" title="MRI">MRI</a> techniques may also uncover the macroscopic process of axonal development. The <a href="/wiki/Connectome" title="Connectome">connectome</a> can be constructed from <a href="/wiki/Diffusion_MRI" class="mw-redirect" title="Diffusion MRI">diffusion MRI</a> data: the vertices of the graph correspond to anatomically labelled gray matter areas, and two such vertices, say <i>u</i> and <i>v</i>, are connected by an edge if the <a href="/wiki/Tractography" title="Tractography">tractography</a> phase of the data processing finds an axonal fiber that connects the two areas, corresponding to <i>u</i> and <i>v</i>. </p><figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><span><video id="mwe_player_0" poster="//upload.wikimedia.org/wikipedia/commons/thumb/f/f5/Ossz_forog.webm/220px--Ossz_forog.webm.jpg" controls="" preload="none" data-mw-tmh="" class="mw-file-element" width="220" height="165" data-durationhint="127" data-mwtitle="Ossz_forog.webm" data-mwprovider="wikimediacommons" resource="/wiki/File:Ossz_forog.webm"><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/f/f5/Ossz_forog.webm/Ossz_forog.webm.360p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="360p.vp9.webm" data-width="480" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/f/f5/Ossz_forog.webm" type="video/webm; codecs="vp8, vorbis"" data-width="480" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/f/f5/Ossz_forog.webm/Ossz_forog.webm.360p.webm" type="video/webm; codecs="vp8, vorbis"" data-transcodekey="360p.webm" data-width="480" data-height="360" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/f/f5/Ossz_forog.webm/Ossz_forog.webm.144p.mjpeg.mov" type="video/quicktime" data-transcodekey="144p.mjpeg.mov" data-width="192" data-height="144" /><source src="//upload.wikimedia.org/wikipedia/commons/transcoded/f/f5/Ossz_forog.webm/Ossz_forog.webm.240p.vp9.webm" type="video/webm; codecs="vp9, opus"" data-transcodekey="240p.vp9.webm" data-width="320" data-height="240" /></video></span><figcaption>Consensus Connectome Dynamics</figcaption></figure><p> Numerous braingraphs, computed from the <a href="/wiki/Human_Connectome_Project" title="Human Connectome Project">Human Connectome Project</a> can be downloaded from the <a rel="nofollow" class="external free" href="http://braingraph.org">http://braingraph.org</a> site. The Consensus Connectome Dynamics (CCD) is a remarkable phenomenon that was discovered by continuously decreasing the minimum confidence-parameter at the graphical interface of the <a href="/wiki/Budapest_Reference_Connectome" title="Budapest Reference Connectome">Budapest Reference Connectome</a> Server.<sup id="cite_ref-connectome_65-0" class="reference"><a href="#cite_note-connectome-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-bpconn3_66-0" class="reference"><a href="#cite_note-bpconn3-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> The Budapest Reference Connectome Server (<a rel="nofollow" class="external free" href="http://connectome.pitgroup.org">http://connectome.pitgroup.org</a>) depicts the cerebral connections of n=418 subjects with a frequency-parameter k: For any k=1,2,...,n one can view the graph of the edges that are present in at least k connectomes. If parameter k is decreased one-by-one from k=n through k=1 then more and more edges appear in the graph, since the inclusion condition is relaxed. The surprising observation is that the appearance of the edges is far from random: it resembles a growing, complex structure, like a tree or a shrub (visualized on the animation on the left). </p><p>It is hypothesized in <sup id="cite_ref-Kerepesi2016_67-0" class="reference"><a href="#cite_note-Kerepesi2016-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> that the growing structure copies the axonal <a href="/wiki/Development_of_the_human_brain" class="mw-redirect" title="Development of the human brain">development of the human brain</a>: the earliest developing connections (axonal fibers) are common at most of the subjects, and the subsequently developing connections have larger and larger variance, because their variances are accumulated in the process of axonal development. </p> <div class="mw-heading mw-heading2"><h2 id="Synapse_elimination">Synapse elimination</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=18" title="Edit section: Synapse elimination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Synaptic_pruning" title="Synaptic pruning">Synaptic pruning</a></div> <p>Several motorneurons compete for each neuromuscular junction, but only one survives until adulthood.<sup id="cite_ref-pmid22745601_37-1" class="reference"><a href="#cite_note-pmid22745601-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Competition <i>in vitro</i> has been shown to involve a limited neurotrophic substance that is released, or that neural activity infers advantage to strong post-synaptic connections by giving resistance to a toxin also released upon nerve stimulation. <i>In vivo</i>, it is suggested that muscle fibres select the strongest neuron through a retrograde signal or that activity-dependent synapse elimination mechanisms determine the identity of the "winning" axon at a motor endplate.<sup id="cite_ref-Meirovitch2021_38-2" class="reference"><a href="#cite_note-Meirovitch2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mapping">Mapping</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=19" title="Edit section: Mapping"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Brain_mapping" title="Brain mapping">Brain mapping</a> can show how an animal's brain changes throughout its lifetime. As of 2021, scientists mapped and compared the whole brains of eight <i><a href="/wiki/Caenorhabditis_elegans" title="Caenorhabditis elegans">C. elegans</a></i> worms across their development on the neuronal level<sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-69" class="reference"><a href="#cite_note-69"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> and the complete wiring of a single mammalian muscle from birth to adulthood.<sup id="cite_ref-Meirovitch2021_38-3" class="reference"><a href="#cite_note-Meirovitch2021-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Adult_neurogenesis">Adult neurogenesis</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=20" title="Edit section: Adult neurogenesis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Adult_neurogenesis" title="Adult neurogenesis">Adult neurogenesis</a></div> <p><a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a> also occurs in specific parts of the adult brain. </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=Development_of_the_nervous_system&action=edit&section=21" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col" style="column-width: 22em;"> <ul><li><a href="/wiki/Axon_guidance" title="Axon guidance">Axon guidance</a></li> <li><a href="/wiki/KCC2#Developmental_changes_in_expression" class="mw-redirect" title="KCC2">KCC2</a></li> <li><a href="/wiki/Pioneer_neuron" title="Pioneer neuron">Pioneer neuron</a></li> <li><a href="/wiki/Neural_Darwinism" title="Neural Darwinism">Neural Darwinism</a></li> <li><a href="/wiki/Brain_development_timelines" title="Brain development timelines">Brain development timelines</a></li> <li><a href="/wiki/Malleable_intelligence" class="mw-redirect" title="Malleable intelligence">Malleable intelligence</a></li> <li><a href="/wiki/Role_of_cell_adhesions_in_neural_development" title="Role of cell adhesions in neural development">Role of cell adhesions in neural development</a></li></ul></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=22" 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-Vet-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-Vet_1-0">^</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 class="citation web cs1"><a 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Retrieved <span class="nowrap">1 May</span> 2020</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Neurological+Signs+%26+Diseases&rft.date=2016-11-02&rft_id=http%3A%2F%2Fvetneuro.com%2FNeurologicalSignsDiseases%2Ftabid%2F4171%2FDefault.aspx&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADevelopment+of+the+nervous+system" class="Z3988"></span></span> </li> <li id="cite_note-2"><span class="mw-cite-backlink"><b><a href="#cite_ref-2">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.nlm.nih.gov/medlineplus/neuraltubedefects.html">"Neural Tube Defects"</a><span class="reference-accessdate">. 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Sinauer Associates Publishers. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/developmentalbio00gilb_292/page/n392">373</a>–379. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87893-250-4" title="Special:BookSources/978-0-87893-250-4"><bdi>978-0-87893-250-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Developmental+biology&rft.pages=373-379&rft.edition=8th&rft.pub=Sinauer+Associates+Publishers&rft.date=2006&rft.isbn=978-0-87893-250-4&rft.aulast=Gilbert&rft.aufirst=S&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdevelopmentalbio00gilb_292&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADevelopment+of+the+nervous+system" class="Z3988"></span></span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZhouSongMing2024" class="citation journal cs1">Zhou Y, Song H, Ming GL (January 2024). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10926850">"Genetics of human brain development"</a>. <i>Nature Reviews. Genetics</i>. <b>25</b> (1): 26–45. <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%2Fs41576-023-00626-5">10.1038/s41576-023-00626-5</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/PMC10926850">10926850</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/37507490">37507490</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature+Reviews.+Genetics&rft.atitle=Genetics+of+human+brain+development&rft.volume=25&rft.issue=1&rft.pages=26-45&rft.date=2024-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC10926850%23id-name%3DPMC&rft_id=info%3Apmid%2F37507490&rft_id=info%3Adoi%2F10.1038%2Fs41576-023-00626-5&rft.aulast=Zhou&rft.aufirst=Y&rft.au=Song%2C+H&rft.au=Ming%2C+GL&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC10926850&rfr_id=info%3Asid%2Fen.wikipedia.org%3ADevelopment+of+the+nervous+system" class="Z3988"></span></span> </li> <li id="cite_note-Wolpert1-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-Wolpert1_5-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWolpert2015" class="citation book cs1">Wolpert L (2015). <a rel="nofollow" class="external text" href="https://archive.org/details/principlesofdeve0000unse_j4n3/page/n5/mode/2up"><i>Principles of development</i></a> (Fifth ed.). 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(May 2020)">page needed</span></a></i>]</sup></span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZhouSongMing2024" class="citation journal cs1">Zhou Y, Song H, Ming GL (January 2024). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10926850">"Genetics of human brain development"</a>. <i>Nature Reviews. 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(May 2020)">page needed</span></a></i>]</sup></span> </li> <li id="cite_note-FOOTNOTEWolpert2015163-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEWolpert2015163_13-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFWolpert2015">Wolpert 2015</a>, pp. 163.</span> </li> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEstomih_MtuiGregory_Gruener2006" class="citation book cs1">Estomih Mtui, Gregory Gruener (2006). <i>Clinical Neuroanatomy and Neuroscience</i>. 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class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Development_of_the_nervous_system&action=edit&section=23" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output 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href="https://ftl.toolforge.org/cgi-bin/ftl?st=wp&su=neural+development">Resources in your library</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=wp&su=neural+development&library=0CHOOSE0">Resources in other libraries</a></li> </ul></div></div> </div> <ul><li><i><a rel="nofollow" class="external text" href="http://www.neuraldevelopment.com/">Neural Development</a></i> (peer-reviewed open access journal).</li> <li><i><a rel="nofollow" class="external text" href="http://www.translatingtime.net/">Translating Neurodevelopmental Time Across Mammalian Species</a></i></li> <li><a rel="nofollow" class="external text" href="https://www.nytimes.com/interactive/2008/09/15/health/20080915-brain-development.html?_r">The Child's Developing Brain</a></li> <li><a rel="nofollow" class="external text" href="http://www.breakingthecycles.com/blog/wp-content/uploads/BreakingTheCycles.com_.Thompsons.BrainDevelopmentTimeLapseStudy3.jpg">Brain Development</a></li> <li><a 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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="Development_of_the_nervous_system" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Development_of_nervous_system" title="Template:Development of nervous system"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Development_of_nervous_system" title="Template talk:Development of nervous system"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Development_of_nervous_system" title="Special:EditPage/Template:Development of nervous system"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Development_of_the_nervous_system" style="font-size:114%;margin:0 4em">Development of the <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neurogenesis" title="Neurogenesis">Neurogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">General</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/Neural_development" class="mw-redirect" title="Neural development">Neural development</a></li> <li><a href="/wiki/Neurulation" title="Neurulation">Neurulation</a></li> <li><a href="/wiki/Neurula" title="Neurula">Neurula</a></li> <li><a href="/wiki/Notochord" title="Notochord">Notochord</a></li> <li><a href="/wiki/Neuroectoderm" title="Neuroectoderm">Neuroectoderm</a></li> <li><a href="/wiki/Neural_plate" title="Neural plate">Neural plate</a> <ul><li><a href="/wiki/Neural_fold" title="Neural fold">Neural fold</a></li> <li><a href="/wiki/Neural_groove" title="Neural groove">Neural groove</a></li></ul></li> <li><a href="/wiki/Neuropoiesis" title="Neuropoiesis">Neuropoiesis</a></li> <li><a href="/wiki/Adult_neurogenesis" title="Adult neurogenesis">Adult neurogenesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neural_crest" title="Neural crest">Neural crest</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/Cranial_neural_crest" title="Cranial neural crest">Cranial neural crest</a> <ul><li><a href="/wiki/Cardiac_neural_crest_complex" class="mw-redirect" title="Cardiac neural crest complex">Cardiac neural crest complex</a></li></ul></li> <li><a href="/wiki/Truncal_neural_crest" class="mw-redirect" title="Truncal neural crest">Truncal neural crest</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Neural_tube" title="Neural tube">Neural tube</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/Rostral_neuropore" title="Rostral neuropore">Rostral neuropore</a></li> <li><a href="/wiki/Neuromere" title="Neuromere">Neuromere</a> / <a href="/wiki/Rhombomere" title="Rhombomere">Rhombomere</a></li> <li><a href="/wiki/Cephalic_flexure" class="mw-redirect" title="Cephalic flexure">Cephalic flexure</a></li> <li><a href="/wiki/Cervical_flexure" class="mw-redirect" title="Cervical flexure">Cervical flexure</a></li> <li><a href="/wiki/Pontine_flexure" class="mw-redirect" title="Pontine flexure">Pontine flexure</a></li> <li><a href="/wiki/Alar_plate" title="Alar plate">Alar plate</a></li> <li><a href="/wiki/Basal_plate_(neural_tube)" title="Basal plate (neural tube)">Basal plate</a></li> <li><a href="/wiki/Glioblast" title="Glioblast">Glioblast</a></li> <li><a href="/wiki/Neuroblast" title="Neuroblast">Neuroblast</a></li> <li><a href="/wiki/Germinal_matrix" title="Germinal matrix">Germinal matrix</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Eye_development" title="Eye development">Eye</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Neural_tube" title="Neural tube">Neural tube</a></li> <li><a href="/wiki/Optic_vesicle" title="Optic vesicle">Optic vesicle</a></li> <li><a href="/wiki/Optic_stalk" title="Optic stalk">Optic stalk</a></li> <li><a href="/wiki/Optic_cup_(embryology)" title="Optic cup (embryology)">Optic cup</a></li> <li><a href="/wiki/Surface_ectoderm" title="Surface ectoderm">Surface ectoderm</a> <ul><li><a href="/wiki/Lens_placode" title="Lens placode">Lens placode</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Ear</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Otic_placode" title="Otic placode">Otic placode</a> <ul><li><a href="/wiki/Otic_pit" title="Otic pit">Otic pit</a></li> <li><a href="/wiki/Otic_vesicle" title="Otic vesicle">Otic vesicle</a></li></ul></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 authority-control" aria-label="Navbox" style="padding:3px"><table class="nowraplinks hlist navbox-inner" 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