174:; the splitting of the vertical branch into two horizontal branches gives rise to a distinctive "T" shape. A parallel fiber runs for an average of 3 mm in each direction from the split, for a total length of about 6 mm (about 1/10 of the total width of the cortical layer). As they run along, the parallel fibers pass through the dendritic trees of
42:
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of mossy fiber inputs. The idea is that with each granule cell receiving input from only 4–5 mossy fibers, a granule cell would not respond if only a single one of its inputs was active, but would respond if more than one were active. This "combinatorial coding" scheme would potentially allow the
223:
Granule cells receive all of their input from mossy fibers, but outnumber them 200 to 1 (in humans). Thus, the information in the granule cell population activity state is the same as the information in the mossy fibers, but recoded in a much more expansive way. Because granule cells are so small
202:
and the early postnatal period, with CGNP proliferation in the EGL peaking during early development (P7, postnatal day 7, in the mouse). As CGNPs terminally differentiate into cerebellum granule cells (also called cerebellar granule neurons, CGNs), they migrate to the internal granule layer (IGL),
146:
is used for several unrelated types of small neurons in various parts of the brain.) Cerebellar granule cells are also the most numerous neurons in the brain: in humans, estimates of their total number average around 50 billion, which means that they constitute about 3/4 of the brain's neurons.
259:
All these features have been observed in murine and human cerebellar tissues, so the mouse model seems to be a good animal model to study the genome structure of cerebellar granule cells, despite the difference in lifespan between the two types of organisms.
283:
between granule cells and
Purkinje cells through chromatin remodelling (specifically, it suppresses genomic accessibility). Mutations in it lead to alterations in synaptogenesis, as a consequence of increased accessibility to genome-wide
224:
and so densely packed, it has been very difficult to record their spike activity in behaving animals, so there is little data to use as a basis of theorizing. The most popular concept of their function was proposed by
241:
Cerebellar granule cells acquire a characteristic genome architecture: ultra-long-range intrachromosomal contacts (10-100Mb), specific interchromosomal contacts and restructuring of active/inactive
272:(ASD), alterations in the chromatin remodelling of granule cells have been identified. This is due to mutations in genes encoding proteins involved in chromatin remodelling. One of these genes is
292:(which are repressed under physiological developmental conditions). However, these alterations at the chromatin level have no effect on the 3D genome architecture of cerebellar granule cells.
155:
The cell bodies are packed into a thick granular layer at the bottom of the cerebellar cortex. A granule cell emits only four to five dendrites, each of which ends in an enlargement called a
575:
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forming the mature cerebellum (by P20, post-natal day 20 in the mouse). Mutations that abnormally activate Sonic hedgehog signaling predispose to cancer of the cerebellum (
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Tan, L., Shi, J., Moghadami, S., Parasar, B., Wright, C. P., Seo, Y., et al. (2023). «Lifelong restructuring of 3D genome architecture in cerebellar granule cells.
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axons of granule cells rise vertically to the upper (molecular) layer of the cortex, where they split in two, with each branch traveling horizontally to form a
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178:, contacting one of every 3–5 that they pass, making a total of 80–100 synaptic connections with Purkinje cell dendritic spines. Granule cells use
576:"Relevancia de los mecanismos epigenéticos en el neurodesarrollo normal y consecuencias de sus perturbaciones | Revista Médica Clínica Las Condes"
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Goodman, Jared V.; Yamada, Tomoko; Yang, Yue; Kong, Lingchun; Wu, Dennis Y.; Zhao, Guoyan; Gabel, Harrison W.; Bonni, Azad (July 9, 2020).
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Polkinghorn, W (2007). "Medulloblastoma: tumorigenesis, current clinical paradigm, and efforts to improve risk stratification".
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signaling stimulates rapid proliferation of cerebellar granule neuron progenitors (CGNPs) in the external granule layer (EGL).
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cerebellum to make much finer distinctions between input patterns than the mossy fibers alone would permit.
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compartmentalisation (scA/B) throughout life. This genomic dynamic is modulated by cell type-specific
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Hatten, M (1995). "Mechanisms of neural patterning and specification in the developing cerebellum".
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602:"The chromatin remodeling enzyme Chd4 regulates genome architecture in the mouse brain"
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as their neurotransmitter, and therefore exert excitatory effects on their targets.
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Science (American
Association for the Advancement of Science), 381(6662), 1112-1119
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at the global level and could be a cellular strategy to manage space and energy.
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387:. Current Topics in Developmental Biology. Vol. 94. pp. 235–82.
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Granule cells, parallel fibers, and flattened dendritic trees of
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and are among the smallest neurons in the brain. (The term
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https://www.science.org/doi/10.1126/science.adh3253
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316:Llinas RR, Walton KD, Lang EJ (2004). "Ch. 7
27:Thick granular layer of the cerebellar cortex
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268:In neurodevelopmental disorders, including
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163:and inhibitory input from Golgi cells.
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322:The Synaptic Organization of the Brain
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324:. New York: Oxford University Press.
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138:form the thick granular layer of the
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113:Parallel fibers to cerebellar cortex
362:10.1146/annurev.ne.18.030195.002125
393:10.1016/B978-0-12-380916-2.00008-5
190:In normal development, endogenous
25:
383:Roussel, M (2011). "Cerebellum".
279:CHD4 is a protein that modulates
119:Anatomical terms of neuroanatomy
840:Oligodendrocyte progenitor cell
502:"A theory of cerebellar cortex"
198:development occurs during late
518:10.1113/jphysiol.1969.sp008820
211:and in genetically engineered
1:
1923:Ventral spinocerebellar tract
73:small cell with few dendrites
1876:Dorsal spinocerebellar tract
950:Postganglionic nerve fibers
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1901:Trigeminocerebellar fibers
945:Preganglionic nerve fibers
618:10.1038/s41467-020-17065-z
559:doi:10.1126/science.adh325
1895:Vestibulocerebellar tract
1455:Olfactory receptor neuron
1119:Neurofibril/neurofilament
320:". In Shepherd GM (ed.).
270:autism spectrum disorders
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136:Cerebellar granule cells
110:Postsynaptic connections
1639:Vallecula of cerebellum
574:Legüe, Marcela (2022).
96:Presynaptic connections
35:Cerebellar granule cell
1912:Pontocerebellar fibers
1697:Deep cerebellar nuclei
1402:Neuromuscular junction
1265:III or Aδ or fast pain
237:3D genome architecture
1928:Dentatothalamic tract
1886:Cuneocerebellar tract
1881:Olivocerebellar tract
606:Nature Communications
1558:Flocculonodular lobe
1420:Meissner's corpuscle
1385:Postsynaptic density
1282:Efferent nerve fiber
1270:IV or C or slow pain
1212:Afferent nerve fiber
1038:Satellite glial cell
456:Nat Clin Pract Oncol
1891:Juxtarestiform body
1805:Unipolar brush cell
1766:Purkinje cell layer
1655:Alar central lobule
1425:Merkel nerve ending
1790:Granule cell layer
1551:Horizontal fissure
1460:Photoreceptor cell
1430:Pacinian corpuscle
1361:Electrical synapse
1315:Lower motor neuron
1310:Upper motor neuron
1031:Internodal segment
971:Connective tissues
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350:Annu Rev Neurosci
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207:) in humans with
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1107:
1105:
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1097:
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1079:
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1022:
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985:
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869:
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851:
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528:
523:
519:
515:
512:(2): 437–70.
511:
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481:
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473:
469:
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457:
450:
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402:9780123809162
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249:, but not by
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200:embryogenesis
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38:
33:
30:
19:
1921:(midbrain):
1839:White matter
1815:Mossy fibers
1800:Granule cell
1789:
1781:Fañanas cell
1765:
1746:
1709:
1661:
1649:
1616:
1594:
1365:Gap junction
1287:Motor neuron
1081:Axon hillock
1057:nerve fibers
1011:Schwann cell
921:
904:
882:
800:Medium spiny
713:White matter
701:Tissue Types
609:
605:
595:
583:. Retrieved
579:
569:
554:
509:
505:
492:
459:
455:
449:
384:
378:
353:
349:
321:
317:
278:
267:
258:
240:
230:combinations
229:
222:
213:mouse models
189:
171:
168:unmyelinated
165:
161:mossy fibers
156:
154:
144:granule cell
135:
134:
100:Mossy fibers
29:
1958:Human cells
1874:(medulla):
1854:Arbor vitae
1757:Basket cell
1688:Grey matter
1380:Active zone
1345:Termination
1195:Interneuron
1099:Telodendron
1007:Myelination
989:Endoneurium
984:Perineurium
805:Interneuron
795:Von Economo
743:Decussation
738:Nerve tract
708:Grey matter
612:(1): 3419.
585:January 16,
356:: 385–408.
254:methylation
186:Development
104:Golgi cells
1953:Cerebellum
1947:Categories
1795:Golgi cell
1717:Emboliform
1711:interposed
1646:Hemisphere
1520:cerebellum
1450:Nociceptor
1190:Multipolar
1139:Nissl body
1016:Neurilemma
979:Epineurium
764:Cell Types
506:J. Physiol
318:Cerebellum
296:References
226:David Marr
196:Cerebellum
166:The thin,
81:excitatory
64:Cerebellum
1864:Peduncles
1729:Fastigial
1662:posterior
1617:posterior
1563:Flocculus
1465:Hair cell
999:Neuroglia
961:Funiculus
850:Microglia
823:Astrocyte
780:Pyramidal
733:Lemniscus
626:2041-1723
439:ignored (
429:cite book
290:enhancers
286:promoters
243:chromatin
180:glutamate
151:Structure
90:glutamate
1919:Superior
1910:(pons):
1872:Inferior
1847:Internal
1813:Fibers:
1650:anterior
1595:anterior
1250:Ia or Aα
1180:Unipolar
1129:Dendrite
1114:Axolemma
1109:Axoplasm
893:Ganglion
833:Tanycyte
785:Purkinje
772:Neuronal
755:Meninges
750:Neuropil
644:32647123
500:(1969).
484:24461280
476:17464337
421:21295689
219:Function
78:Function
60:Location
1963:Neurons
1722:Globose
1705:Dentate
1610:Lingula
1527:Surface
1518:of the
1516:Anatomy
1392:Autapse
1353:Synapse
1200:Renshaw
1175:Bipolar
1052:Neurons
905:Ventral
876:General
790:Granule
635:7347877
536:5784296
527:1351491
412:3213765
370:7605067
55:Details
1908:Middle
1622:Folium
1605:Culmen
1591:Vermis
1568:Nodule
1245:fibers
883:Dorsal
642:
632:
624:
534:
524:
498:Marr D
482:
474:
419:
409:
399:
368:
328:
1632:Uvula
1627:Tuber
1534:Lobes
1168:Types
1065:Parts
934:White
915:Ramus
898:Ramus
815:Glial
480:S2CID
247:genes
123:[
70:Shape
1091:Axon
1073:Soma
929:Gray
910:Root
888:Root
640:PMID
622:ISSN
587:2024
532:PMID
472:PMID
441:help
417:PMID
397:ISBN
366:PMID
326:ISBN
288:and
274:CHD4
102:and
1305:SVE
1300:GVE
1295:GSE
1240:SVA
1235:SSA
1230:GVA
1225:GSA
868:PNS
693:CNS
630:PMC
614:doi
561:».
522:PMC
514:doi
510:202
464:doi
407:PMC
389:doi
358:doi
251:CpG
1949::
1648::
1593::
1009::
638:.
628:.
620:.
610:11
608:.
604:.
578:.
557:.
544:^
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508:.
504:.
478:.
470:.
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433::
431:}}
427:{{
415:.
405:.
395:.
364:.
354:18
352:.
340:^
304:^
276:.
215:.
1897:)
1893:(
1508:e
1501:t
1494:v
1363:/
1284:/
1214:/
1151:/
1054:/
952:)
943:(
677:e
670:t
663:v
646:.
616::
589:.
538:.
516::
486:.
466::
460:4
443:)
423:.
391::
372:.
360::
334:.
127:]
20:)
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