Development of the nervous system

858: 823: 406: 1352:) 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). 1035:(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. 1088:

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.

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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.

1271:, 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. 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. In the 5252: 216: 4798: 1097:

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

649:, 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 1323: 1111:

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

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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. This is an activity-dependent event. Partial blockage of the receptor

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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

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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. They have distinct spatial and temporal patterns during development and their

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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

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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. It is estimated that glial guided migration represents 90% of migrating neurons in human and about

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in mediating some aspects of these processes such as the rate of neuronal migration, aspects of neuronal differentiation and axon pathfinding. Activity-dependent mechanisms influence neural circuit development and are crucial for laying out early connectivity maps and the continued refinement of

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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

980:, which make a long journey from their birthplace in the nose, through the forebrain, and into the hypothalamus. Many of the mechanisms of this migration have been worked out, starting with the extracellular guidance cues that trigger intracellular signaling. These intracellular signals, such as 1038:

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

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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

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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

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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

454:. 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 1050:

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

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and cortical slices. 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

923:, whose fibers serve as a scaffolding for migrating cells and a means of radial communication mediated by calcium dynamic activity, act as the main excitatory neuronal stem cell of the cerebral cortex or translocate to the cortical plate and differentiate either into 361:

differs from that formed in later developmental stages, and from adult CSF; it influences the behavior of neural precursors. 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

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along the dorsal side of the embryo. This is a part of the early patterning of the embryo (including the invertebrate embryo) that also establishes an anterior-posterior axis. The neural plate is the source of the majority of neurons and glial cells of the CNS. The

1221:, 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. 1248:. 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. In the motor system, periodic bursts of spontaneous activity are driven by excitatory 1359:: 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. 1328: 1324: 1326: 517:

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

1015:. 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. 1377:

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.

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synapses which occurs during development. 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

1382:, 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. 1327: 621:

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

1043: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. 947:

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

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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

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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

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Meirovitch Y, Kang K, Draft RW, Pavarino EC, Henao Echeverri MF, Yang F, et al. (September 2021). "Neuromuscular connectomes across development reveal synaptic ordering rules".

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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

265:. When the tube is closed at both ends it is filled with embryonic cerebrospinal fluid. As the embryo develops, the anterior part of the neural tube expands and forms three 1325: 737:(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 1080:. 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 5290: 386:

to different parts of the developing brain to self-organize into different brain structures. Once the neurons have reached their regional positions, they extend

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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

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Nadarajah B, Alifragis P, Wong RO, Parnavelas JG (June 2003). "Neuronal migration in the developing cerebral cortex: observations based on real-time imaging".

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Noctor SC, Flint AC, Weissman TA, Dammerman RS, Kriegstein AR (February 2001). "Neurons derived from radial glial cells establish radial units in neocortex".

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Nadarajah B, Brunstrom JE, Grutzendler J, Wong RO, Pearlman AL (February 2001). "Two modes of radial migration in early development of the cerebral cortex".

1084:. Fischbach and colleagues showed that receptor subunits are selectively transcribed from nuclei next to the synaptic site. This is mediated by neuregulins. 2801:"Suppression of β1-integrin in gonadotropin-releasing hormone cells disrupts migration and axonal extension resulting in severe reproductive alterations" 115: 2380:

Tamamaki N, Nakamura K, Okamoto K, Kaneko T (September 2001). "Radial glia is a progenitor of neocortical neurons in the developing cerebral cortex".

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The survival of neurons is regulated by survival factors, called trophic factors. The neurotrophic hypothesis was formulated by Victor Hamburger and

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Meister M, Wong RO, Baylor DA, Shatz CJ (May 1991). "Synchronous bursts of action potentials in ganglion cells of the developing mammalian retina".

2750:"Capture of microtubule plus-ends at the actin cortex promotes axophilic neuronal migration by enhancing microtubule tension in the leading process" 1476: 695:

The dorsal neural tube is patterned by BMPs from the epidermal ectoderm flanking the neural plate. These induce sensory interneurons by activating

2940:"Reversing the outcome of synapse elimination at developing neuromuscular junctions in vivo: evidence for synaptic competition and its mechanism" 906: 531:

differentiate into neural cells. Inhibition of TGF-β and BMP (bone morphogenetic protein) signaling can efficiently induce neural tissue from

442:. At the onset of gastrulation presumptive mesodermal cells move through the dorsal blastopore lip and form a layer of mesoderm in between the 32:

This article is about neural development in all types of animals, including humans. For information specific to the human nervous system, see

4372:"How to Direct the Edges of the Connectomes: Dynamics of the Consensus Connectomes and the Development of the Connections in the Human Brain" 5160: 1456: 5283: 224: 33: 4439: 3639: 1393:

can show how an animal's brain changes throughout its lifetime. As of 2021, scientists mapped and compared the whole brains of eight

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worms across their development on the neuronal level and the complete wiring of a single mammalian muscle from birth to adulthood.

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development, and at highest concentrations it induces floor plate differentiation. Failure of Shh-modulated differentiation causes

2652:"Hepatocyte growth factor acts as a motogen and guidance signal for gonadotropin hormone-releasing hormone-1 neuronal migration" 5276: 4632: 438:

which gives rise to the nervous system. The conversion of undifferentiated ectoderm to neuroectoderm requires signals from the

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and there are good markers to distinguish between neural and non-neural tissue. Examples of neural inducers are the molecules

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ablation during development has been known to result in deficits in network refinement in the visual system. In the immature

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O'Donovan MJ (February 1999). "The origin of spontaneous activity in developing networks of the vertebrate nervous system".

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and are critical for cell fate determination in the developing and adult mammalian brain. Epigenetic modifications include

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to the cerebral cortex. One example of ongoing tangential migration in a mature organism, observed in some animals, is the

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to describe and provide insight into the cellular and molecular mechanisms by which complex nervous systems develop, from

5575: 78: 63: 4313:"Parameterizable consensus connectomes from the Human Connectome Project: the Budapest Reference Connectome Server v3.0" 374:, which drive brain growth as they divide many times. Gradually some of the cells stop dividing and differentiate into 3710:

Torborg CL, Feller MB (July 2005). "Spontaneous patterned retinal activity and the refinement of retinal projections".

731: 724: 522:(which allow direct cell-cell interactions) the same cells differentiate into epidermis. This is due to the action of 17: 101: 73: 723:, which are expressed in overlapping domains along the anteroposterior axis under the control of retinoic acid. The 543:

In a later stage of development the superior part of the neural tube flexes at the level of the future midbrain—the

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O'Donovan MJ, Chub N, Wenner P (October 1998). "Mechanisms of spontaneous activity in developing spinal networks".

1207: 2274:"Bidirectional radial Ca(2+) activity regulates neurogenesis and migration during early cortical column formation" 1299:

data: the vertices of the graph correspond to anatomically labelled gray matter areas, and two such vertices, say

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map and eye-specific segregation. Retinotopic map refinement occurs in downstream visual targets in the brain-the

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Galli L, Maffei L (October 1988). "Spontaneous impulse activity of rat retinal ganglion cells in prenatal life".

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Several motorneurons compete for each neuromuscular junction, but only one survives until adulthood. Competition

696: 3663:"Spatial-temporal patterns of retinal waves underlying activity-dependent refinement of retinofugal projections" 3121:"Spontaneous rhythmic activity in early chick spinal cord influences distinct motor axon pathfinding decisions" 1356: 4565: 5256: 4981: 1609: 1345: 1139: 1121: 1076:. They also showed that the synaptogenic signal is produced by the nerve, and they identified the factor as 5393: 5135: 4956: 889:

cells must leave the stem cell niche and migrate outward to form the preplate, which is destined to become

764:. Neurons are 'post-mitotic', meaning that they will never divide again for the lifetime of the organism. 582:(which eventually become the optic nerve, retina and iris) forms at the basal plate of the prosencephalon. 5175: 2464:

Nadarajah B, Parnavelas JG (June 2002). "Modes of neuronal migration in the developing cerebral cortex".

1446: 680:- it induces cell differentiation dependent on its concentration. At low concentrations it forms ventral 83: 4841: 4258:

Szalkai B, Kerepesi C, Varga B, Grolmusz V (May 2015). "The Budapest Reference Connectome Server v2.0".

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tracts to migrate along; this is called axophilic migration. An example of this mode of migration is in

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receptors at the synapse. McMahan and Sanes showed that the synaptogenic signal is concentrated at the

1012: 623: 5452: 5170: 5145: 5014: 3489:"Traveling waves in developing cerebellar cortex mediated by asymmetrical Purkinje cell connectivity" 2853:"Multipolar migration: the third mode of radial neuronal migration in the developing cerebral cortex" 1147:

development even when sensory input is absent and is observed in many systems such as the developing

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Cserép C, Schwarcz AD, Pósfai B, László ZI, Kellermayer A, Környei Z, et al. (September 2022).

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and sweep across the retinal surface in the first few postnatal weeks. These waves are mediated by

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occurs due to specific environmental conditions - different concentrations of signaling molecules

2507:

Rakic P (May 1972). "Mode of cell migration to the superficial layers of fetal monkey neocortex".

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Rakic P (May 1972). "Mode of cell migration to the superficial layers of fetal monkey neocortex".

730:(3 prime end) genes in the Hox cluster are induced by retinoic acid in the hindbrain, whereas the 645:(Shh) from the notochord, which acts as the inducing tissue. Notochord-derived Shh signals to the 4946: 4657: 4652: 1311:

phase of the data processing finds an axonal fiber that connects the two areas, corresponding to

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dynamics, which produce cellular forces that interact with the extracellular environment through

807: 204: 161: 5504: 4941: 4931: 4722: 4677: 4605: 4600: 4065:"Calcium action potentials in hair cells pattern auditory neuron activity before hearing onset" 2550:

Letinic K, Zoncu R, Rakic P (June 2002). "Origin of GABAergic neurons in the human neocortex".

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Witvliet D, Mulcahy B, Mitchell JK, Meirovitch Y, Berger DR, Wu Y, et al. (August 2021).

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forms along the long axis of the neural plate, and the neural plate folds to give rise to the

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Tritsch NX, Rodríguez-Contreras A, Crins TT, Wang HC, Borst JG, Bergles DE (September 2010).

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Gato A, Alonso MI, Martín C, Carnicero E, Moro JA, De la Mano A, et al. (August 2014).

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Parkash J, Cimino I, Ferraris N, Casoni F, Wray S, Cappy H, et al. (November 2012).

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provide a synaptogenic factor. The identity of the astrocytic factors is not yet known.

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and the ectoderm. Mesodermal cells migrate along the dorsal midline to give rise to the

5370: 5323: 5195: 5089: 5029: 4752: 4732: 4526: 4513: 4470: 4416: 4383: 4371: 4347: 4324: 4312: 4293: 4267: 4197: 4164: 4140: 4113: 4089: 4064: 4040: 4015: 3993: 3980: 3955: 3931: 3898: 3874: 3849: 3822: 3797: 3735: 3687: 3662: 3562: 3537: 3513: 3488: 3428: 3380: 3371: 3355: 3331: 3321: 3304: 3242: 3194: 3169: 3145: 3120: 3096: 3086: 3069: 3009: 2966: 2939: 2877: 2868: 2852: 2825: 2800: 2776: 2749: 2725: 2700: 2676: 2651: 2627: 2602: 2583: 2532: 2489: 2405: 2362: 2306: 2273: 2254: 2211: 2168: 2117: 2092: 2068: 2043: 2016: 1991: 1938: 1913: 1912:

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Schoenwolf GC, Smith JL (2000). "Mechanisms of Neurulation". In Tuan RS, Lo CW (eds.).

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Huberman AD (February 2007). "Mechanisms of eye-specific visual circuit development".

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is a cell adhesion molecule that is present in both pre- and post-synaptic membranes.

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of the embryo. A part of the dorsal ectoderm becomes specified to neural ectoderm –

5509: 5437: 5432: 5155: 5039: 4966: 4881: 4762: 4712: 4508: 4500: 4411: 4401: 4342: 4334: 4297: 4277: 4192: 4184: 4135: 4125: 4084: 4076: 4035: 4027: 3975: 3967: 3926: 3918: 3869: 3861: 3817: 3809: 3770: 3719: 3682: 3674: 3635: 3598: 3557: 3549: 3508: 3500: 3459: 3412: 3375: 3367: 3326: 3316: 3277: 3226: 3189: 3181: 3140: 3132: 3091: 3081: 3040: 3001: 2961: 2951: 2909: 2872: 2864: 2820: 2816: 2812: 2771: 2761: 2720: 2716: 2712: 2671: 2667: 2663: 2622: 2614: 2587: 2567: 2516: 2473: 2436: 2409: 2389: 2366: 2346: 2301: 2293: 2238: 2195: 2172: 2152: 2112: 2104: 2063: 2055: 2011: 2003: 1960: 1933: 1925: 1819: 1811: 1745: 1737: 1685: 1580: 1572: 1441: 1368: 1281: 1272: 1245: 1233: 1184: 874: 803: 783: 761: 742: 631: 548: 507: 451: 3538:"Spontaneous, synchronous electrical activity in neonatal mouse cortical neurones" 2091:

Noack F, Pataskar A, Schneider M, Buchholz F, Tiwari VK, Calegari F (April 2019).

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Many neurons migrating along the anterior-posterior axis of the body use existing

626:, which are blocked by the organiser. The organiser may produce molecules such as 398:. Synaptic communication between neurons leads to the establishment of functional 5481: 5467: 5442: 5215: 5210: 5205: 5200: 5094: 5034: 4971: 4886: 4846: 4831: 4782: 4772: 4727: 4406: 4281: 3971: 3865: 3678: 3553: 3464: 3447: 3185: 2956: 1963: 1237: 1225: 1206:(LGN). Pharmacological disruption and mouse models lacking the β2 subunit of the 1152: 791: 779: 771: 579: 560: 519: 493: 382:, which are the main cellular components of the CNS. The newly generated neurons 298: 290: 4495: 1689: 496:

without inducing mesodermal genes as well. Neural induction is often studied in

405: 5300: 5180: 5150: 5079: 5054: 5044: 4891: 4871: 4861: 4747: 4667: 4504: 3850:"A role for correlated spontaneous activity in the assembly of neural circuits" 3230: 3170:"A role for correlated spontaneous activity in the assembly of neural circuits" 2603:"From nose to brain: development of gonadotrophin-releasing hormone-1 neurones" 2007: 1815: 1726:"Embryonic cerebrospinal fluid in brain development: neural progenitor control" 1576: 1436: 1291:

techniques may also uncover the macroscopic process of axonal development. The

1144: 961: 902: 826: 700: 642: 556: 399: 345:). The CSF-filled central chamber is continuous from the telencephalon to the 338: 274: 141: 4338: 4188: 4114:"Large-scale synchronized activity in the embryonic brainstem and spinal cord" 3813: 3446:

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of the vertebrate, the dorsal ectoderm becomes specified to give rise to the

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Processes which grow and shape an organism's nervous tissue over its lifetime

3774: 3281: 3070:"Spontaneous neuronal calcium spikes and waves during early differentiation" 1741: 798:. Methylcytosine demethylation is catalyzed in several sequential steps by 5542: 5532: 5514: 5380: 5365: 5348: 5308: 5190: 5185: 5069: 4901: 4826: 4777: 4767: 4742: 4737: 4707: 4641: 4522: 4425: 4356: 4289: 4206: 4149: 4098: 4049: 3989: 3940: 3883: 3831: 3731: 3696: 3612: 3571: 3522: 3473: 3424: 3389: 3238: 3203: 3154: 2975: 2921: 2886: 2834: 2785: 2734: 2685: 2636: 2579: 2485: 2450: 2401: 2358: 2315: 2297: 2207: 2164: 2126: 2108: 2077: 2025: 1947: 1833: 1759: 1707: 1594: 1413: 1308: 1210:

has shown that the lack of spontaneous activity leads to marked defects in

1156: 1073: 992: 910: 753: 738: 685: 607: 575: 486: 466: 435: 430:

and the nervous system; a part of the dorsal ectoderm becomes specified to

367: 314: 306: 249: 149: 48: 3782: 3647: 3340: 3289: 3105: 3054: 2528: 2520: 2250: 2242: 913:"cage" around the nucleus elongates and contracts in association with the 489:, and the open ends of the neural tube, called the neuropores, close off. 5499: 5489: 5410: 5353: 5328: 5059: 4906: 4165:"Emergence of patterned activity in the developing zebrafish spinal cord" 3640:

10.1002/(sici)1097-4695(199810)37:1<131::aid-neu10>3.0.co;2-h

3487:

Watt AJ, Cuntz H, Mori M, Nusser Z, Sjöström PJ, Häusser M (April 2009).

3029:"Intracellular Ca2+ fluctuations modulate the rate of neuronal migration" 1268: 1195: 1160: 989: 977: 935: 830: 799: 767: 681: 646: 627: 474: 470: 410: 379: 363: 350: 262: 258: 4440:"Why a tiny worm's brain development could shed light on human thinking" 3922: 2571: 2199: 5462: 5447: 5427: 5220: 4921: 3305:"Rhythmic spontaneous activity in the developing avian auditory system" 2059: 1292: 1276: 1211: 1164: 1108: 914: 654: 482: 334: 241: 230: 4471:"Connectomes across development reveal principles of brain maturation" 1194:. They are thought to instruct the formation of two sensory maps- the 614:, which acts as the 'organiser'. The dorsal half is controlled by the 5457: 5423: 5338: 5165: 4856: 2350: 1929: 1265: 1261: 1253: 1218: 1191: 1168: 924: 898: 878: 720: 677: 611: 591: 503: 447: 427: 391: 293:). These simple, early vesicles enlarge and further divide into the 286: 270: 184: 4576: 4080: 4031: 4016:"Tonotopic reorganization of developing auditory brainstem circuits" 3504: 1355:

It is hypothesized in that the growing structure copies the axonal

5537: 5074: 4388: 4329: 4062: 3356:"Primordial rhythmic bursting in embryonic cochlear ganglion cells" 2477: 2425:"Asymmetric inheritance of radial glial fibers by cortical neurons" 894: 615: 527: 443: 439: 395: 310: 278: 237: 157: 4584: 4272: 2186:

Samuels BA, Tsai LH (November 2004). "Nucleokinesis illuminated".

2156: 1992:"Retinoic acid synthesis and signaling during early organogenesis" 934:

Subsequent waves of neurons split the preplate by migrating along

594:, dorsal ectoderm forms all neural tissue and the nervous system. 402:

that mediate sensory and motor processing, and underlie behavior.

5333: 4468: 2142: 1911: 1349: 1229: 886: 650: 635: 511: 498: 461:

In the early embryo, the neural plate folds outwards to form the

375: 188: 180: 165: 140:), refers to the processes that generate, shape, and reshape the 2899: 931:. Somal translocation can occur at any time during development. 711:

Signals that control anteroposterior neural development include

473:. The formation of the neural tube from the ectoderm is called 171:

Defects in neural development can lead to malformations such as

4610: 3587:"Spontaneous correlated activity in developing neural circuits" 1845: 1843: 1176: 1081: 928: 838: 2649: 2090: 2379: 1077: 985: 387: 192: 148:

to adulthood. The field of neural development draws on both

4585:

Translating Neurodevelopmental Time Across Mammalian Species

4369: 4311:

Szalkai B, Kerepesi C, Varga B, Grolmusz V (February 2017).

4310: 4257: 2328: 1840: 1244:

release from supporting cells triggers action potentials in

215: 4370:

Kerepesi C, Szalkai B, Varga B, Grolmusz V (30 June 2016).

3798:"Assembly and disassembly of a retinal cholinergic network" 3661:

Stafford BK, Sher A, Litke AM, Feldheim DA (October 2009).

3445: 2995: 2798: 1886: 1249: 1047: 1046:

GDNF: Glial derived neurotrophic factor is a member of the

1040: 973: 670: 666: 662: 523: 330: 4797: 3953: 2423:

Miyata T, Kawaguchi A, Okano H, Ogawa M (September 2001).

370:. During this time, the walls of the neural tube contain 4162: 3660: 2422: 1723: 1636: 1288: 741:. Without this Hoxb-1 expression, a nerve similar to the 566:

The alar plate of the prosencephalon expands to form the

394:, which allow them to communicate with other neurons via 1534:(8th ed.). Sinauer Associates Publishers. pp. 1341: 1179:, waves of spontaneous action potentials arise from the 3848:

Kirkby LA, Sack GS, Firl A, Feller MB (December 2013).

3625: 3168:

Kirkby LA, Sack GS, Firl A, Feller MB (December 2013).

1657:

Anatomy & Physiology The Unity of Form and Function

1240:

neurons which relay auditory information to the brain.

3259: 3118: 1868: 706: 5298: 1232:

generate bursts of activity which spreads across the

618:

plate, which flanks either side of the neural plate.

3896: 3847: 3167: 3119:

Hanson MG, Milner LD, Landmesser LT (January 2008).

1275:, early waves of activity have been observed in the 1007:

There is also a method of neuronal migration called

477:. The ventral part of the neural tube is called the 3486: 2463: 756:is the process by which neurons are generated from 3897:Ackman JB, Burbridge TJ, Crair MC (October 2012). 2698: 1962: 1416:also occurs in specific parts of the adult brain. 1348:Server. The Budapest Reference Connectome Server ( 3353: 2991: 2989: 2987: 2985: 2549: 1854:(5th ed.). Appleton and Lange: McGraw Hill. 5557: 4013: 3535: 829:: younger neurons migrate past older ones using 4111: 3354:Jones TA, Jones SM, Paggett KC (October 2001). 2933: 2931: 2271: 1679: 917:to guide the nucleus to its final destination. 210: 4222: 3067: 2982: 2937: 2850: 1115: 881:, where the principal neural stem cell is the 5284: 4626: 4009: 4007: 3843: 3841: 3709: 3536:Corlew R, Bosma MM, Moody WJ (October 2004). 3402: 2272:Rash BG, Ackman JB, Rakic P (February 2016). 2041: 1905: 657:, leading to activation of the Gli family of 109: 5161:Intraoperative neurophysiological monitoring 4363: 3068:Gu X, Olson EC, Spitzer NC (November 1994). 2928: 2747: 2543: 2416: 2373: 1797: 1558: 1457:Role of cell adhesions in neural development 873:Neuronal precursor cells proliferate in the 465:. Beginning in the future neck region, the 4229:(Third ed.). Burlington MA: Elsevier. 4218: 4216: 3752: 3026: 2846: 2844: 2699:Hutchins BI, Klenke U, Wray S (July 2013). 2643: 2185: 2037: 2035: 1614:(Fifth ed.). Oxford University Press. 905:is then transported to the pial surface by 653:, relieving Patched-mediated inhibition of 225:Development of the nervous system in humans 34:Development of the nervous system in humans 5291: 5277: 4633: 4619: 4251: 4004: 3838: 3795: 2741: 2692: 2044:"DNA methylation dynamics in neurogenesis" 2042:Wang Z, Tang B, He Y, Jin P (March 2016). 1777:(Tenth ed.). Sinauer Associates Inc. 1682:Developmental Biology Protocols: Volume II 861:Tangential migration of interneurons from 116: 102: 4512: 4494: 4415: 4405: 4387: 4346: 4328: 4271: 4196: 4139: 4129: 4088: 4039: 3979: 3930: 3873: 3821: 3686: 3602: 3561: 3512: 3463: 3379: 3330: 3320: 3193: 3144: 3095: 3085: 3044: 2965: 2955: 2876: 2824: 2792: 2775: 2765: 2724: 2675: 2626: 2440: 2305: 2138: 2136: 2116: 2067: 2015: 1937: 1823: 1749: 1584: 1521: 1519: 1060: 802:that carry out oxidative reactions (e.g. 559:(future forebrain) and beneath it is the 4213: 4014:Kandler K, Clause A, Noh J (June 2009). 3216: 2841: 2032: 1798:Zhou Y, Song H, Ming GL (January 2024). 1559:Zhou Y, Song H, Ming GL (January 2024). 1336:Numerous braingraphs, computed from the 1321: 856: 821: 641:The ventral neural tube is patterned by 404: 219:Diagram of the vertebrate nervous system 214: 144:of animals, from the earliest stages of 4304: 2938:Turney SG, Lichtman JW (26 June 2012). 1989: 1983: 1969:(4th ed.). New York: McGraw-Hill. 1874: 1772: 1654: 1642: 1607: 1601: 1525: 1018: 42:This article is part of a series on the 14: 5558: 3584: 2851:Tabata H, Nakajima K (November 2003). 2594: 2133: 1889:Clinical Neuroanatomy and Neuroscience 1887:Estomih Mtui, Gregory Gruener (2006). 1849: 1719: 1717: 1659:. New York: McGraw Hill. p. 514. 1516: 1362: 999:to cause the movement of these cells. 684:, at higher concentrations it induces 574:, whilst its basal plate becomes the 5272: 4614: 3302: 2506: 2228: 1891:. Philadelphia: Saunders. p. 1. 1800:"Genetics of human brain development" 1561:"Genetics of human brain development" 1402: 1190:in the initial phase and later on by 817: 485:. The hollow interior is called the 5251: 4580:(peer-reviewed open access journal). 2600: 2509:The Journal of Comparative Neurology 2231:The Journal of Comparative Neurology 1055: 601: 187:, balance and vision disorders, and 3796:Ford KJ, Feller MB (January 2012). 1714: 1477:"Neurological Signs & Diseases" 1264:at later stages. In the developing 707:Rostrocaudal (Anteroposterior) axis 469:of this groove close to create the 24: 4601:How poverty might change the brain 4118:Frontiers in Cellular Neuroscience 3372:10.1523/JNEUROSCI.21-20-08129.2001 3322:10.1523/JNEUROSCI.14-03-01486.1994 3087:10.1523/JNEUROSCI.14-11-06325.1994 2869:10.1523/JNEUROSCI.23-31-09996.2003 2754:Frontiers in Cellular Neuroscience 1469: 1307:, are connected by an edge if the 547:, at the mesencephalic flexure or 538: 25: 5587: 5141:Development of the nervous system 4552:Development of the nervous system 4540: 4226:Development of the Nervous System 4223:Sanes D, Reh T, Harris W (2012). 3137:10.1016/j.brainresrev.2007.06.021 3027:Komuro H, Rakic P (August 1996). 1479:. 2 November 2016. Archived from 502:embryos since they have a simple 353:, and constitutes the developing 130:development of the nervous system 5250: 5239: 5238: 4796: 4640: 2619:10.1111/j.1365-2826.2010.02034.x 1287:Contemporary diffusion-weighted 1208:nicotinic acetylcholine receptor 481:; the dorsal part is called the 4462: 4432: 4156: 4105: 4056: 3947: 3890: 3789: 3746: 3724:10.1016/j.pneurobio.2005.09.002 3703: 3654: 3619: 3578: 3529: 3480: 3439: 3405:Current Opinion in Neurobiology 3396: 3347: 3296: 3253: 3219:Current Opinion in Neurobiology 3210: 3161: 3112: 3061: 3020: 2893: 2500: 2457: 2322: 2265: 2222: 2179: 2084: 1954: 1880: 1791: 1766: 1256:during the early stages and by 1091: 748: 4112:Momose-Sato Y, Sato K (2013). 2817:10.1523/JNEUROSCI.3057-12.2012 2717:10.1523/JNEUROSCI.3758-12.2013 2668:10.1523/JNEUROSCI.4979-06.2007 1673: 1648: 1552: 1495: 1357:development of the human brain 1350:http://connectome.pitgroup.org 1214:and eye-specific segregation. 796:DNA methyltransferases (DNMTs) 788:5-methylcytosine demethylation 770:play a key role in regulating 676:In this context Shh acts as a 366:or lifelong disabilities like 13: 1: 4982:Social cognitive neuroscience 3604:10.1016/s0896-6273(00)80724-2 3417:10.1016/s0959-4388(99)80012-9 3046:10.1016/s0896-6273(00)80159-2 2607:Journal of Neuroendocrinology 2442:10.1016/S0896-6273(01)00420-2 2394:10.1016/S0168-0102(01)00259-0 1463: 1346:Budapest Reference Connectome 1332:Consensus Connectome Dynamics 1140:activity-dependent mechanisms 1122:Activity-dependent plasticity 1002: 942: 703:transcription factor levels. 585: 5571:Embryology of nervous system 5566:Animal developmental biology 5394:Cardiac neural crest complex 4957:Molecular cellular cognition 4591:The Child's Developing Brain 4566:Resources in other libraries 4407:10.1371/journal.pone.0158680 4282:10.1016/j.neulet.2015.03.071 3972:10.1016/j.celrep.2022.111369 3866:10.1016/j.neuron.2013.10.030 3679:10.1016/j.neuron.2009.09.021 3554:10.1113/jphysiol.2004.071621 3465:10.1016/j.neuron.2007.03.007 3186:10.1016/j.neuron.2013.10.030 2957:10.1371/journal.pbio.1001352 2748:Hutchins BI, Wray S (2014). 2466:Nature Reviews. Neuroscience 1990:Duester G (September 2008). 1965:Principles of neural science 1852:Principles of neural science 1217:Recent studies confirm that 967: 417: 211:Vertebrate brain development 7: 5176:Neurodevelopmental disorder 5151:Neural network (biological) 5146:Neural network (artificial) 4496:10.1101/2020.04.30.066209v3 3360:The Journal of Neuroscience 3309:The Journal of Neuroscience 3074:The Journal of Neuroscience 2857:The Journal of Neuroscience 2805:The Journal of Neuroscience 2705:The Journal of Neuroscience 2656:The Journal of Neuroscience 1447:Brain development timelines 1419: 1340:can be downloaded from the 1116:Assembly of neural circuits 1101:can develop into synapses. 261:. This process is known as 10: 5592: 5576:Developmental neuroscience 4703:Computational neuroscience 4505:10.1038/s41586-021-03778-8 3231:10.1016/j.conb.2007.01.005 2008:10.1016/j.cell.2008.09.002 1816:10.1038/s41576-023-00626-5 1577:10.1038/s41576-023-00626-5 1406: 1385: 1366: 1204:lateral geniculate nucleus 1119: 1064: 1013:cortical intermediate zone 236:(CNS) is derived from the 222: 31: 5525: 5480: 5409: 5379: 5316: 5307: 5234: 5171:Neurodegenerative disease 5128: 5015:Evolutionary neuroscience 4990: 4930: 4805: 4794: 4666: 4648: 4561:Resources in your library 4339:10.1007/s11571-016-9407-z 4189:10.1016/j.cub.2011.12.002 3814:10.1017/S0952523811000216 3542:The Journal of Physiology 3006:10.1101/2021.09.20.460480 1690:10.1385/1-59259-065-9:125 1611:Principles of development 868: 810:) and enzymes of the DNA 5136:Brain–computer interface 5085:Neuromorphic engineering 5010:Educational neuroscience 4917:Nutritional neuroscience 4822:Clinical neurophysiology 4718:Integrative neuroscience 4131:10.3389/fncel.2013.00036 3712:Progress in Neurobiology 3585:Feller MB (April 1999). 2767:10.3389/fncel.2014.00400 1804:Nature Reviews. Genetics 1730:Croatian Medical Journal 1565:Nature Reviews. Genetics 1338:Human Connectome Project 1295:can be constructed from 954:rostral migratory stream 792:DNA cytosine methylation 780:DNA cytosine methylation 768:Epigenetic modifications 606:The ventral half of the 570:which gives rise to the 195:other disorders such as 175:, and a wide variety of 4947:Behavioral neuroscience 4317:Cognitive Neurodynamics 3775:10.1126/science.3175637 3628:Journal of Neurobiology 3303:Lippe WR (March 1994). 3282:10.1126/science.2035024 2914:10.1093/cercor/13.6.607 1742:10.3325/cmj.2014.55.299 978:GnRH-expressing neurons 909:, a process by which a 808:5-hydroxymethylcytosine 450:that develops into the 205:intellectual disability 4942:Affective neuroscience 4723:Molecular neuroscience 4678:Behavioral epigenetics 3125:Brain Research Reviews 3080:(11 Pt 1): 6325–6335. 2298:10.1126/sciadv.1501733 2109:10.26508/lsa.201900331 1452:Malleable intelligence 1333: 1181:retinal ganglion cells 1067:Neuromuscular junction 1061:Neuromuscular junction 997:cell adhesion proteins 865: 842: 533:pluripotent stem cells 414: 357:of the CNS. Embryonic 267:primary brain vesicles 234:central nervous system 220: 177:neurological disorders 5005:Cultural neuroscience 5000:Consumer neuroscience 4842:Neurogastroenterology 4698:Cellular neuroscience 3315:(3 Pt 2): 1486–1495. 2521:10.1002/cne.901450105 2382:Neuroscience Research 2243:10.1002/cne.901450105 2097:Life Science Alliance 1775:Developmental Biology 1530:Developmental biology 1503:"Neural Tube Defects" 1342:http://braingraph.org 1331: 1120:Further information: 860: 852:time-lapse microscopy 825: 659:transcription factors 610:is controlled by the 424:embryonic development 408: 223:Further information: 218: 154:developmental biology 146:embryonic development 5401:Truncal neural crest 5389:Cranial neural crest 4977:Sensory neuroscience 4817:Behavioral neurology 4788:Systems neuroscience 4260:Neuroscience Letters 2601:Wray S (July 2010). 1918:Nature Biotechnology 1025:Rita Levi Montalcini 1019:Neurotrophic factors 1009:multipolar migration 812:base excision repair 572:cerebral 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359:cerebrospinal fluid 269:, which become the 69:Reproductive system 5371:Adult neurogenesis 5324:Neural development 5196:Neuroimmune system 5090:Neurophenomenology 5030:Neural engineering 4753:Neuroendocrinology 4733:Neural engineering 4577:Neural Development 2863:(31): 9996–10001. 2060:10.2217/epi.15.119 1850:Kandel ER (2006). 1773:Gilbert S (2013). 1655:Saladin K (2011). 1608:Wolpert L (2015). 1526:Gilbert S (2006). 1409:Adult neurogenesis 1403:Adult neurogenesis 1334: 1224:In the developing 1128:neuronal migration 921:Radial glial cells 877:of the developing 866: 843: 833:as a scaffolding. 818:Neuronal migration 415: 355:ventricular system 221: 134:neural development 89:Circulatory system 18:Neural development 5553: 5552: 5476: 5475: 5419:Rostral neuropore 5266: 5265: 5115:Paleoneurobiology 5050:Neuroepistemology 5025:Neuroanthropology 4991:Interdisciplinary 4877:Neuropharmacology 4837:Neuroepidemiology 4606:The Teenage Brain 4596:Brain Development 4547:Library resources 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549:cephalic flexure 520:explant cultures 456:neural induction 452:vertebral column 138:neurodevelopment 118: 111: 104: 79:Endocrine system 64:Digestive system 39: 38: 21: 5591: 5590: 5586: 5585: 5584: 5582: 5581: 5580: 5556: 5555: 5554: 5549: 5521: 5472: 5468:Germinal matrix 5443:Pontine flexure 5405: 5375: 5303: 5297: 5267: 5262: 5230: 5216:Neurotechnology 5211:Neuroplasticity 5206:Neuromodulation 5201:Neuromanagement 5124: 5095:Neurophilosophy 4992: 4986: 4972:Neuropsychology 4933: 4926: 4887:Neuropsychiatry 4847:Neuroimmunology 4832:Neurocardiology 4808: 4801: 4792: 4783:Neurophysiology 4773:Neuromorphology 4728:Neural decoding 4669: 4662: 4644: 4639: 4572: 4571: 4570: 4555: 4554: 4550: 4543: 4538: 4467: 4463: 4453: 4451: 4450:on 20 June 2022 4438: 4437: 4433: 4382:(6): e0158680. 4368: 4364: 4309: 4305: 4256: 4252: 4237: 4221: 4214: 4169:Current Biology 4161: 4157: 4110: 4106: 4081:10.1038/nn.2604 4061: 4057: 4032:10.1038/nn.2332 4012: 4005: 3952: 3948: 3895: 3891: 3846: 3839: 3794: 3790: 3761:(4875): 90–91. 3751: 3747: 3708: 3704: 3659: 3655: 3624: 3620: 3583: 3579: 3534: 3530: 3505:10.1038/nn.2285 3485: 3481: 3444: 3440: 3401: 3397: 3352: 3348: 3301: 3297: 3258: 3254: 3215: 3211: 3166: 3162: 3117: 3113: 3066: 3062: 3025: 3021: 2994: 2983: 2950:(6): e1001352. 2936: 2929: 2902:Cerebral Cortex 2898: 2894: 2849: 2842: 2797: 2793: 2746: 2742: 2697: 2693: 2648: 2644: 2599: 2595: 2548: 2544: 2505: 2501: 2462: 2458: 2421: 2417: 2378: 2374: 2327: 2323: 2284:(2): e1501733. 2270: 2266: 2227: 2223: 2184: 2180: 2141: 2134: 2089: 2085: 2040: 2033: 1988: 1984: 1977: 1959: 1955: 1910: 1906: 1899: 1885: 1881: 1877:, pp. 163. 1873: 1869: 1862: 1848: 1841: 1796: 1792: 1785: 1771: 1767: 1722: 1715: 1700: 1678: 1674: 1667: 1653: 1649: 1641: 1637: 1622: 1606: 1602: 1557: 1553: 1546: 1524: 1517: 1507: 1505: 1501: 1500: 1496: 1486: 1484: 1475: 1474: 1470: 1466: 1461: 1422: 1411: 1405: 1388: 1371: 1365: 1322: 1238:spiral ganglion 1226:auditory system 1153:auditory system 1132:differentiation 1124: 1118: 1094: 1069: 1063: 1058: 1021: 1005: 970: 945: 871: 820: 814:(BER) pathway. 772:gene expression 751: 733: 726: 709: 604: 588: 580:optical vesicle 561:rhombencephalon 541: 539:Regionalization 432:neural ectoderm 420: 400:neural circuits 299:cerebral cortex 291:rhombencephalon 248:that forms the 240:—the outermost 227: 213: 122: 93: 47:Development of 37: 28: 23: 22: 15: 12: 11: 5: 5589: 5579: 5578: 5573: 5568: 5551: 5550: 5548: 5547: 5546: 5545: 5540: 5529: 5527: 5523: 5522: 5520: 5519: 5518: 5517: 5507: 5502: 5497: 5492: 5486: 5484: 5478: 5477: 5474: 5473: 5471: 5470: 5465: 5460: 5455: 5450: 5445: 5440: 5435: 5430: 5421: 5415: 5413: 5407: 5406: 5404: 5403: 5398: 5397: 5396: 5385: 5383: 5377: 5376: 5374: 5373: 5368: 5363: 5362: 5361: 5356: 5346: 5341: 5336: 5331: 5326: 5320: 5318: 5311: 5305: 5304: 5301:nervous system 5296: 5295: 5288: 5281: 5273: 5264: 5263: 5261: 5260: 5248: 5235: 5232: 5231: 5229: 5228: 5226:Self-awareness 5223: 5218: 5213: 5208: 5203: 5198: 5193: 5188: 5183: 5181:Neurodiversity 5178: 5173: 5168: 5163: 5158: 5153: 5148: 5143: 5138: 5132: 5130: 5126: 5125: 5123: 5122: 5117: 5112: 5107: 5102: 5097: 5092: 5087: 5082: 5080:Neuromarketing 5077: 5072: 5067: 5062: 5057: 5055:Neuroesthetics 5052: 5047: 5045:Neuroeconomics 5042: 5037: 5032: 5027: 5022: 5017: 5012: 5007: 5002: 4996: 4994: 4988: 4987: 4985: 4984: 4979: 4974: 4969: 4964: 4959: 4954: 4949: 4944: 4938: 4936: 4928: 4927: 4925: 4924: 4919: 4914: 4909: 4904: 4899: 4894: 4892:Neuroradiology 4889: 4884: 4879: 4874: 4872:Neuropathology 4869: 4864: 4862:Neuro-oncology 4859: 4854: 4849: 4844: 4839: 4834: 4829: 4824: 4819: 4813: 4811: 4803: 4802: 4795: 4793: 4791: 4790: 4785: 4780: 4775: 4770: 4765: 4760: 4755: 4750: 4748:Neurochemistry 4745: 4740: 4735: 4730: 4725: 4720: 4715: 4710: 4705: 4700: 4695: 4690: 4685: 4680: 4674: 4672: 4664: 4663: 4661: 4660: 4655: 4649: 4646: 4645: 4638: 4637: 4630: 4623: 4615: 4609: 4608: 4603: 4598: 4593: 4588: 4581: 4569: 4568: 4563: 4557: 4556: 4545: 4544: 4542: 4541:External links 4539: 4537: 4536: 4461: 4431: 4362: 4323:(1): 113–116. 4303: 4250: 4235: 4212: 4155: 4104: 4055: 4026:(6): 711–717. 4003: 3966:(12): 111369. 3946: 3889: 3837: 3788: 3745: 3718:(4): 213–235. 3702: 3673:(2): 200–212. 3653: 3634:(1): 131–145. 3618: 3597:(4): 653–656. 3577: 3528: 3499:(4): 463–473. 3479: 3458:(1): 105–120. 3438: 3395: 3346: 3295: 3252: 3209: 3160: 3111: 3060: 3039:(2): 275–285. 3019: 2981: 2927: 2908:(6): 607–611. 2892: 2840: 2791: 2740: 2691: 2662:(2): 431–445. 2642: 2613:(7): 743–753. 2593: 2542: 2499: 2478:10.1038/nrn845 2472:(6): 423–432. 2456: 2435:(5): 727–741. 2415: 2372: 2321: 2264: 2221: 2178: 2151:(2): 143–150. 2132: 2083: 2054:(3): 401–414. 2031: 2002:(6): 921–931. 1982: 1975: 1953: 1924:(3): 275–280. 1904: 1897: 1879: 1867: 1860: 1839: 1790: 1783: 1765: 1736:(4): 299–305. 1713: 1698: 1672: 1665: 1647: 1635: 1620: 1600: 1551: 1544: 1515: 1494: 1467: 1465: 1462: 1460: 1459: 1454: 1449: 1444: 1439: 1437:Pioneer neuron 1434: 1429: 1423: 1421: 1418: 1407:Main article: 1404: 1401: 1387: 1384: 1367:Main article: 1364: 1361: 1145:neural circuit 1117: 1114: 1093: 1090: 1065:Main article: 1062: 1059: 1057: 1054: 1053: 1052: 1044: 1036: 1020: 1017: 1004: 1001: 969: 966: 962:olfactory bulb 944: 941: 870: 867: 837:(red) release 827:Corticogenesis 819: 816: 750: 747: 708: 705: 697:Sr/Thr kinases 643:sonic hedgehog 603: 600: 587: 584: 557:prosencephalon 540: 537: 419: 416: 339:myelencephalon 275:prosencephalon 212: 209: 142:nervous system 124: 123: 121: 120: 113: 106: 98: 95: 94: 92: 91: 86: 81: 76: 74:Urinary system 71: 66: 61: 59:Nervous system 55: 52: 51: 44: 43: 26: 9: 6: 4: 3: 2: 5588: 5577: 5574: 5572: 5569: 5567: 5564: 5563: 5561: 5544: 5541: 5539: 5536: 5535: 5534: 5531: 5530: 5528: 5524: 5516: 5513: 5512: 5511: 5508: 5506: 5503: 5501: 5498: 5496: 5495:Optic vesicle 5493: 5491: 5488: 5487: 5485: 5483: 5479: 5469: 5466: 5464: 5461: 5459: 5456: 5454: 5451: 5449: 5446: 5444: 5441: 5439: 5436: 5434: 5431: 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5016: 5013: 5011: 5008: 5006: 5003: 5001: 4998: 4997: 4995: 4989: 4983: 4980: 4978: 4975: 4973: 4970: 4968: 4965: 4963: 4962:Motor control 4960: 4958: 4955: 4953: 4952:Chronobiology 4950: 4948: 4945: 4943: 4940: 4939: 4937: 4935: 4929: 4923: 4920: 4918: 4915: 4913: 4912:Neurovirology 4910: 4908: 4905: 4903: 4900: 4898: 4895: 4893: 4890: 4888: 4885: 4883: 4880: 4878: 4875: 4873: 4870: 4868: 4865: 4863: 4860: 4858: 4855: 4853: 4850: 4848: 4845: 4843: 4840: 4838: 4835: 4833: 4830: 4828: 4825: 4823: 4820: 4818: 4815: 4814: 4812: 4810: 4804: 4799: 4789: 4786: 4784: 4781: 4779: 4776: 4774: 4771: 4769: 4766: 4764: 4761: 4759: 4758:Neurogenetics 4756: 4754: 4751: 4749: 4746: 4744: 4741: 4739: 4736: 4734: 4731: 4729: 4726: 4724: 4721: 4719: 4716: 4714: 4711: 4709: 4706: 4704: 4701: 4699: 4696: 4694: 4693:Brain-reading 4691: 4689: 4688:Brain mapping 4686: 4684: 4681: 4679: 4676: 4675: 4673: 4671: 4665: 4659: 4656: 4654: 4651: 4650: 4647: 4643: 4636: 4631: 4629: 4624: 4622: 4617: 4616: 4613: 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2244: 2240: 2236: 2232: 2225: 2217: 2213: 2209: 2205: 2201: 2197: 2193: 2189: 2182: 2174: 2170: 2166: 2162: 2158: 2157:10.1038/83967 2154: 2150: 2146: 2139: 2137: 2128: 2124: 2119: 2114: 2110: 2106: 2102: 2098: 2094: 2087: 2079: 2075: 2070: 2065: 2061: 2057: 2053: 2049: 2045: 2038: 2036: 2027: 2023: 2018: 2013: 2009: 2005: 2001: 1997: 1993: 1986: 1978: 1972: 1967: 1966: 1957: 1949: 1945: 1940: 1935: 1931: 1927: 1923: 1919: 1915: 1908: 1900: 1894: 1890: 1883: 1876: 1871: 1863: 1857: 1853: 1846: 1844: 1835: 1831: 1826: 1821: 1817: 1813: 1809: 1805: 1801: 1794: 1786: 1780: 1776: 1769: 1761: 1757: 1752: 1747: 1743: 1739: 1735: 1731: 1727: 1720: 1718: 1709: 1705: 1701: 1695: 1691: 1687: 1683: 1676: 1668: 1662: 1658: 1651: 1644: 1639: 1631: 1627: 1623: 1617: 1613: 1612: 1604: 1596: 1592: 1587: 1582: 1578: 1574: 1570: 1566: 1562: 1555: 1547: 1541: 1537: 1532: 1531: 1522: 1520: 1504: 1498: 1483:on 2016-11-02 1482: 1478: 1472: 1468: 1458: 1455: 1453: 1450: 1448: 1445: 1443: 1440: 1438: 1435: 1433: 1430: 1428: 1427:Axon guidance 1425: 1424: 1417: 1415: 1410: 1400: 1398: 1397: 1392: 1391:Brain mapping 1383: 1381: 1376: 1370: 1360: 1358: 1353: 1351: 1347: 1343: 1339: 1320: 1318: 1314: 1310: 1306: 1302: 1298: 1297:diffusion MRI 1294: 1290: 1285: 1283: 1278: 1274: 1270: 1267: 1263: 1259: 1258:acetylcholine 1255: 1251: 1247: 1243: 1239: 1235: 1231: 1228:, developing 1227: 1222: 1220: 1215: 1213: 1209: 1205: 1201: 1197: 1193: 1189: 1188:acetylcholine 1186: 1182: 1178: 1172: 1170: 1166: 1162: 1158: 1154: 1150: 1149:visual system 1146: 1141: 1137: 1136:axon guidance 1133: 1129: 1123: 1113: 1110: 1106: 1102: 1100: 1089: 1085: 1083: 1079: 1075: 1068: 1049: 1045: 1042: 1037: 1034: 1031: 1030: 1029: 1026: 1016: 1014: 1010: 1000: 998: 994: 991: 987: 983: 979: 975: 965: 963: 959: 955: 951: 940: 937: 932: 930: 926: 922: 918: 916: 912: 908: 907:nucleokinesis 904: 900: 896: 892: 888: 885:. The first 884: 880: 876: 864: 859: 855: 853: 848: 840: 836: 832: 828: 824: 815: 813: 809: 805: 801: 797: 793: 789: 785: 781: 777: 773: 769: 765: 763: 759: 755: 746: 744: 740: 736: 729: 722: 718: 717:retinoic acid 714: 704: 702: 699:and altering 698: 693: 691: 687: 683: 679: 674: 672: 668: 664: 660: 656: 652: 648: 644: 639: 637: 633: 629: 625: 619: 617: 613: 609: 599: 597: 593: 583: 581: 577: 573: 569: 568:telencephalon 564: 562: 558: 554: 553:mesencephalon 550: 546: 545:mesencephalon 536: 534: 529: 525: 521: 515: 513: 509: 505: 501: 500: 495: 490: 488: 484: 480: 476: 472: 468: 464: 463:neural groove 459: 457: 453: 449: 445: 441: 437: 433: 429: 425: 422:During early 412: 409:Flowchart of 407: 403: 401: 397: 393: 389: 385: 381: 377: 373: 369: 365: 360: 356: 352: 348: 347:central canal 344: 340: 336: 332: 328: 327:metencephalon 324: 320: 319:mesencephalon 316: 312: 308: 304: 303:basal ganglia 300: 296: 295:telencephalon 292: 288: 284: 283:mesencephalon 280: 276: 272: 268: 264: 260: 256: 255:neural groove 251: 247: 246:neuroectoderm 243: 239: 235: 232: 226: 217: 208: 206: 202: 201:Down syndrome 198: 197:Rett syndrome 194: 190: 186: 182: 178: 174: 169: 167: 163: 159: 155: 151: 147: 143: 139: 135: 131: 119: 114: 112: 107: 105: 100: 99: 97: 96: 90: 87: 85: 82: 80: 77: 75: 72: 70: 67: 65: 62: 60: 57: 56: 54: 53: 50: 49:organ systems 46: 45: 41: 40: 35: 30: 19: 5543:Otic vesicle 5533:Otic placode 5515:Lens placode 5381:Neural crest 5366:Neuropoiesis 5349:Neural plate 5309:Neurogenesis 5255: 5243: 5191:Neuroimaging 5186:Neurogenesis 5140: 5070:Neurohistory 5035:Neurobiotics 4934:neuroscience 4902:Neurosurgery 4827:Epileptology 4809:neuroscience 4778:Neurophysics 4768:Neurometrics 4743:Neurobiology 4738:Neuroanatomy 4708:Connectomics 4642:Neuroscience 4583: 4575: 4551: 4478: 4474: 4464: 4454:21 September 4452:. 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Development of the nervous system

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