Documente Academic
Documente Profesional
Documente Cultură
Because of its large number of tiny granule cells, the cerebellum contains more neurons than the rest of the
brain put together, but it only takes up 10% of total brain volume.[3] The cerebellum receives nearly
200 million input fibers; in contrast, the optic nerve is composed of a mere one million fibers.
The unusual surface appearance of the cerebellum conceals the fact that the bulk of the structure is made up of
a very tightly folded layer of gray matter, the cerebellar cortex. It has been estimated that if the human
cerebellar cortex could be completely unfolded it would give rise to a layer of neural tissue about 1 meter long
and 10 centimeters wide—a total surface area of 500-1000 square cm, all packed within a volume of 100-150
cubic cm.[4][5] Underneath the gray matter of the cortex lies white matter, made up largely of myelinated nerve
fibers running to and from the cortex. Embedded within the white matter—which is sometimes called the
arbor vitae (Tree of Life) in the cerebellum because of its branched, tree-like appearance—are four deep
cerebellar nuclei.
The cerebellum can be divided according to three different criteria: gross anatomical, phylogenetical, and
functional.
Much of what is understood about the functions of the cerebellum stems from careful documentation of the
effects of focal lesions in human patients who have suffered from injury or disease or through animal lesion
research.
Cellular anatomy
As explained in more detail in the Function section, the cerebellum differs from most other brain areas in that
the flow of neural signals through it is almost entirely unidirectional: there are virtually no backward
connections between its neuronal elements. Thus the most logical way to describe the cellular structure is to
begin with the inputs and follow the sequence of connections through to the outputs.
Deep nuclei
The four deep nuclei of the cerebellum are the dentate, emboliform, globose, and fastigii nuclei and they act as
the main centers of communication, sending and receiving information to and from specific parts of the brain.
In addition, these nuclei receive both inhibitory and excitatory signals from other parts of the brain which in
turn affect the nuclei's outgoing signals.[7](The globose and the emboliform nuclei make up the interposed
nucleus).
Cortical layers
The cytoarchitecture (cellular organization) of the
cerebellum is highly uniform, with connections organized
into a rough, three-dimensional array of perpendicular
circuit elements. This organizational uniformity makes the
nerve circuitry relatively easy to study.
Molecular layer
This outermost layer of the cerebellar cortex contains two
types of inhibitory interneurons: the stellate and basket
cells. It also contains the dendritic arbors of Purkinje
neurons and parallel fiber tracts from the granule cells.
Both stellate and basket cells form GABAergic synapses
Figure 6: Confocal micrograph from mouse
onto Purkinje cell dendrites.
cerebellum expressing green-fluorescent protein in
Purkinje cells
Purkinje layer
The middle layer contains only one type of cell body—that of the large Purkinje cell. Purkinje cells are the
primary integrative neurons of the cerebellar cortex and provide its sole output. Purkinje cell dendrites are
large arbors with hundreds of spiny branches reaching up into the molecular layer (Fig. 6). These dendritic
arbors are flat—nearly all of them lie in planes—with neighboring Purkinje arbors in parallel planes. Each
parallel fiber from the granule cells runs orthogonally through these arbors, like a wire passing through many
layers. Purkinje neurons are GABAergic—meaning they have inhibitory synapses—with the neurons of the
deep cerebellar and vestibular nuclei in the brainstem. Each Purkinje cell receives excitatory input from
100,000 to 200,000 parallel fibers. Parallel fibers are said to be responsible for the simple (all or nothing,
amplitude invariant) spiking of the Purkinje cell.
Purkinje cells also receive input from the inferior olivary nucleus via climbing fibers. A good mnemonic for
this interaction is the phrase "climb the other olive tree", given that climbing fibers originate from the
contralateral inferior olive. In striking contrast to the 100,000-plus inputs from parallel fibers, each Purkinje
cell receives input from exactly one climbing fiber; but this single fiber "climbs" the dendrites of the Purkinje
cell, winding around them and making a large number of synapses as it goes. The net input is so strong that a
single action potential from a climbing fiber is capable of producing a "complex spike" in the Purkinje cell: a
burst of several spikes in a row, with diminishing amplitude,[8] followed by a pause during which simple
spikes are suppressed.
Just underneath the Purkinje layer are the Lugaro cells whose very long dendrites travel along the boundary
between the Purkinje and the granular layers.
Granular layer
The innermost layer contains the cell bodies of three types of cells: the numerous and tiny granule cells, the
slightly larger unipolar brush cells[9] and the much larger Golgi cells. Mossy fibers enter the granular layer
from their main point of origin, the pontine nuclei. These fibers form excitatory synapses with the granule cells
and the cells of the deep cerebellar nuclei. The granule cells send their T-shaped axons—known as parallel
fibers—up into the superficial molecular layer, where they form hundreds of thousands of synapses with
Purkinje cell dendrites. The human cerebellum contains on the order of 60 to 80 billion granule cells, making
this single cell type by far the most numerous neuron in the brain (roughly 70% of all neurons in the brain and
spinal cord, combined). Golgi cells provide inhibitory feedback to granule cells, forming a synapse with them
and projecting an axon into the molecular layer.
The corticopontocerebellar pathway is the largest pathway associated with the cerebellum. Arising in the
cerebral cortex these fibers first terminate ipsilaterally in the pontine nuclei. Then the fibers decussate and
form the middle cerebellar peduncle, terminating in the cerebellar cortex as mossy fibers. This pathway
transmits signals that inform the cerebellum about the movement in progress and the upcoming movement.
This helps the continuous adjustment of motor activity.[12]
The initiation of the movement is relayed to cerebellum via the corticoreticulocerebellar pathway. Those
synapse ipsilaterally in the reticular formation, then via the inferior and middle peduncles into the cerebellar
vermis.[12]
The motor cortex and somatosensory cortex projects onto the ipsilateral inferior and accessory olivary nuclei,
then forming the olivocerebellar tract. Cortico-olivary fibers synapse bilaterally in the inferior olivary nucleus.
The order is preserved in the olivocerebellar tract projections onto the ‘body maps’ in the contralateral
cerebellar cortex. Under resting conditions in animal experiments, groups of olivary neurons discharge
synchronously at 5 to 10 Hz (impulses/s). In the cerebellar cortex, the response of Purkinje cells takes the form
of complex spikes.[13]
The cerebellum send its projections back to the cerebral cortex via the Cerebellothalamic tract.
The cerebellar lateral expansion, or the neocerebellum, may be associated with cognitive functions, and it is
anatomically linked with the lateral prefrontal cortex. It shows greatest activity during speech, with a one-sided
predominance consistent with a possible linkage (via the thalamus) with the motor speech area.[13]
When lesions occur in the association areas linked to the cerebellum by corticopontocerebellar fibres, the
cognitive affective syndrome may occur. This results in cognitive defects in the form of diminished reasoning
power, inattention, grammatical errors in speech, poor spatial sense, and patchy memory loss.[13]
Blood supply
Three arteries supply blood to the cerebellum (Fig. 7): the
superior cerebellar artery (SCA), anterior inferior
cerebellar artery (AICA), and posterior inferior cerebellar
artery (PICA).
In amphibians, lampreys, and hagfish the cerebellum is little developed; in the latter two groups it is barely
distinguishable from the brain-stem. Although the spinocerebellum is present in these groups, the primary
structures are small paired nuclei corresponding to the vestibulocerebellum.[15]
Peduncles
The cerebellum follows the general groups-of-three pattern found in anatomy,[16] with three major input and
output cerebellar peduncles (fiber bundles). These are the superior (brachium conjunctivum), middle
(brachium pontis), and inferior (restiform and juxtarestiform bodies) cerebellar peduncles.
Peduncle Description
While there are some afferent fibers from the anterior spinocerebellar tract that are conveyed to the anterior
cerebellar lobe via this peduncle, most of the fibers are efferents. Thus, the superior cerebellar peduncle is
the major output pathway of the cerebellum. Most of the efferent fibers originate within the dentate nucleus
which in turn project to various midbrain structures including the red nucleus, the ventral lateral/ventral
Superior
anterior nucleus of the thalamus, and the medulla. The dentatorubrothalamocortical (dentate nucleus > red
nucleus > thalamus > premotor cortex) and cerebellothalamocortical (cerebellum > thalamus > premotor
cortex) pathways are two major pathways that pass through this peduncle and are important in motor
planning.
This is composed entirely of afferent fibers originating within the pontine nuclei as part of the massive
corticopontocerebellar tract (cerebral cortex > pons > cerebellum). These fibers descend from the sensory
Middle
and motor areas of the cerebral neocortex and make the middle cerebellar peduncle the largest of the three
cerebellar peduncles.
This carries many types of input and output fibers that are mainly concerned with integrating proprioceptive
sensory input with motor vestibular functions such as balance and posture maintenance. Proprioceptive
information from the body is carried to the cerebellum via the dorsal spinocerebellar tract. This tract passes
through the inferior cerebellar peduncle and synapses within the paleocerebellum. Vestibular information
Inferior
projects onto the archicerebellum.
The climbing fibers of the inferior olive run through the inferior cerebellar peduncle.
This peduncle also carries information directly from the Purkinje cells out to the vestibular nuclei in the
dorsal brainstem located at the junction between the pons and medulla.
There are three sources of input to the cerebellum, in two categories consisting of mossy and climbing fibers,
respectively. Mossy fibers can originate from the pontine nuclei, which are clusters of neurons located in the
pons that carry information from the contralateral cerebral cortex. They may also arise within the
spinocerebellar tract whose origin is located in the ipsilateral spinal cord. Most of the output from the
cerebellum initially synapses onto the deep cerebellar nuclei before exiting via the three peduncles. The most
notable exception is the direct inhibition of the vestibular nuclei by Purkinje cells.
Development
During the early stages of embryonic development, the brain starts to form in three distinct segments: the
prosencephalon, mesencephalon, and rhombencephalon. The rhombencephalon is the most caudal (toward the
tail) segment of the embryonic brain; it is from this segment that the cerebellum develops. Along the
embryonic rhombencephalic segment develop eight swellings, called rhombomeres. The cerebellum arises
from two rhombomeres located in the alar plate of the neural tube, a structure that eventually forms the brain
and spinal cord. The specific rhombomeres from which the cerebellum forms are rhombomere 1 (Rh.1)
caudally (near the tail) and the "isthmus" rostrally (near the front).[17]
Two primary regions are thought to give rise to the neurons that make up the cerebellum. The first region is the
ventricular zone in the roof of the fourth ventricle. This area produces Purkinje cells and deep cerebellar
nuclear neurons. These cells are the primary output neurons of the cerebellar cortex and cerebellum. The
second germinal zone (cellular birthplace) is known as the Rhombic lip, neurons then move by human
embryonic week 27 to the external granular layer. This layer of cells—found on the exterior of the cerebellum
—produces the granule neurons. The granule neurons migrate from this exterior layer to form an inner layer
known as the internal granule layer.[18] The external granular layer ceases to exist in the mature cerebellum,
leaving only granule cells in the internal granule layer. The cerebellar white matter may be a third germinal
zone in the cerebellum; however, its function as a germinal zone is controversial.
Additional images
References
1. Knierim, James. "Chapter 5: Cerebellum" (https://nba.uth.tmc.edu/neuroscience/s3/chapter05.ht
ml). Neuroscience Online: An Electronic Textbook for the Neurosciences.
2. Friede, Reinhard L. (1973-03-01). "Dating the development of human cerebellum". Acta
Neuropathologica. 23 (1): 48–58. doi:10.1007/BF00689004 (https://doi.org/10.1007%2FBF00689
004). ISSN 1432-0533 (https://www.worldcat.org/issn/1432-0533). PMID 4698523 (https://pubme
d.ncbi.nlm.nih.gov/4698523).
3. The Brain From Top To Bottom (http://thebrain.mcgill.ca/flash/d/d_06/d_06_cr/d_06_cr_mou/d_0
6_cr_mou.html)
4. Edwards CR, Newman S, Bismark A, et al. (2008). "Cerebellum volume and eyeblink
conditioning in schizophrenia" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2366060).
Psychiatry Res. 162 (3): 185–194. doi:10.1016/j.pscychresns.2007.06.001 (https://doi.org/10.101
6%2Fj.pscychresns.2007.06.001). PMC 2366060 (https://www.ncbi.nlm.nih.gov/pmc/articles/PM
C2366060). PMID 18222655 (https://pubmed.ncbi.nlm.nih.gov/18222655).
5. Hutchinson S, Lee LH, Gaab N, Schlaug G (2003). "Cerebellar volume of musicians" (http://cerco
r.oxfordjournals.org/cgi/content/full/13/9/943). Cereb. Cortex. 13 (9): 943–9.
doi:10.1093/cercor/13.9.943 (https://doi.org/10.1093%2Fcercor%2F13.9.943). PMID 12902393
(https://pubmed.ncbi.nlm.nih.gov/12902393).
6. Kingsley, RE (2000). Concise Text of Neuroscience (2nd ed.). Lippincott Williams and Wilkins.
ISBN 0-683-30460-7.
7. Harting, J.K. "The Global Cerebellum '97" (http://www.neuroanatomy.wisc.edu/cere/text/P5/intro.
htm). University of Wisconsin Medical School.
8. Häusser, Michael; Clark, Beverley A.; Davie, Jenny T. (2008-07-23). "The origin of the complex
spike in cerebellar Purkinje cells". Journal of Neuroscience. 28 (30): 7599–7609.
doi:10.1523/JNEUROSCI.0559-08.2008 (https://doi.org/10.1523%2FJNEUROSCI.0559-
08.2008). ISSN 0270-6474 (https://www.worldcat.org/issn/0270-6474). PMID 18650337 (https://p
ubmed.ncbi.nlm.nih.gov/18650337).
9. Kinney GA, Overstreet LS, Slater NT (September 1997). "Prolonged physiological entrapment of
glutamate in the synaptic cleft of cerebellar unipolar brush cells" (http://jn.physiology.org/cgi/repri
nt/78/3/1320.pdf) (PDF). J Neurophysiol. 78 (3): 1320–33. PMID 9310423 (https://pubmed.ncbi.nl
m.nih.gov/9310423).
10. Soteropoulos DS, Baker SN (2006). "Cortico-cerebellar coherence during a precision grip task in
the monkey" (http://jn.physiology.org/cgi/reprint/95/2/1194). J Neurophysiol. 95 (2): 1194–206.
doi:10.1152/jn.00935.2005 (https://doi.org/10.1152%2Fjn.00935.2005). PMID 16424458 (https://
pubmed.ncbi.nlm.nih.gov/16424458).
11. Ros H, Sachdev RN, Yu Y, Sestan N, McCormick DA (2009). "Neocortical networks entrain
neuronal circuits in cerebellar cortex" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3137973).
Journal of Neuroscience. 29 (33): 10309–20. doi:10.1523/JNEUROSCI.2327-09.2009 (https://do
i.org/10.1523%2FJNEUROSCI.2327-09.2009). PMC 3137973 (https://www.ncbi.nlm.nih.gov/pm
c/articles/PMC3137973). PMID 19692605 (https://pubmed.ncbi.nlm.nih.gov/19692605).
12. Gartner, Leslie P.; Patestas, Maria A. (2009). Textbook of Neuroanatomy. Wiley-Blackwell.
p. 464. ISBN 9781405103404.
13. Mtui, Estomih; Gruener, Gregory; Dockery, Peter (2016). Fitzgerald's Clinical Neuroanatomy and
Neuroscience (7th ed.). Elsevier. pp. 243–252.
14. Gray, Henry; Lewis, Warren Harmon (1918). Anatomy of the human body (http://archive.org/detai
ls/anatomyofhumanbo1918gray) (20th ed.). Philadelphia: Lea & Febiger.
15. Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA:
Holt-Saunders International. p. 531. ISBN 0-03-910284-X.
16. "List of Three's" (http://www.meddean.luc.edu/lumen/meded/grossanatomy/threes.html).
www.meddean.luc.edu.
17. Muller F, O'Rahilly R (1990). "The human brain at stages 21–23, with particular reference to the
cerebral cortical plate and to the development of the cerebellum". Anat Embryol (Berl). 182 (4):
375–400. doi:10.1007/BF02433497 (https://doi.org/10.1007%2FBF02433497). PMID 2252222 (h
ttps://pubmed.ncbi.nlm.nih.gov/2252222).
18. Smeyne, Richard J.; Goldowitz, Dan (May 1989). "Development and death of external granular
layer cells in the weaver mouse cerebellum: a quantitative study". The Journal of Neuroscience.
9 (5): 1608–20. doi:10.1523/JNEUROSCI.09-05-01608.1989 (https://doi.org/10.1523%2FJNEUR
OSCI.09-05-01608.1989). PMID 2723742 (https://pubmed.ncbi.nlm.nih.gov/2723742).
Text is available under the Creative Commons Attribution-ShareAlike License; additional terms may apply. By using this site,
you agree to the Terms of Use and Privacy Policy. Wikipedia® is a registered trademark of the Wikimedia Foundation, Inc.,
a non-profit organization.