Cerebellum 

Francico Olucha Bordonau

Universitat de València-Estudi General

Cerebellum ................................................................................................................................................... 1
Development ............................................................................................................................................ 1
Macroscopic structure .............................................................................................................................. 2
Cytoarchitectonic structure ....................................................................................................................... 3
Cerebelar circuits ...................................................................................................................................... 3
Systems and projections of the cerebellum .............................................................................................. 4
Vestibulo-cerebellum ........................................................................................................................... 4
Spino-cerebellum.................................................................................................................................. 5
Ponto cerebellum .................................................................................................................................. 6

The cerebellum is a projection of the dorsal rhomboencephalon that has a highly repetitive organization.
Although any sensory impulse can reach the cerebellum, it is not involved in conscious sensory
perception, but perception and other sensory information is used in automatic coordination of somatic
motor function, regulation of muscle tone and maintaining balance.

Development 
The cerebellum originates from the anterior edges of the rhombic lips. The cells of the anterior rhombic
lip proliferate and migrate dorsally being placed on the outside while their axonic processes remain in the
deep layer. Some cells migrate less and are grouped into the deepest part intermingled between the axonic
bundles making up the three pairs of deep nuclei. The lower segment of the rhombic lips migrate
ventrally to lie below the Tegmentum forming the nuclei of the pons which relay information from the
neocortex to the cerebellum. Another group of cells migrate more caudally to originate the bulbar
(medullary) inferior olivary complexes. The olivary nuclei consist of various bulbar neuronal groups that

Figure 1.- Development of the Cerebellum from a dorsal view (A) of the rhombic lips and from its
anterior part, a group of cells migrate ventrally to originate thenuclei of the pons and inferior
olivary nuclei, (B and C). On the other hand, another group of neurons migrate dorsally to originate
the cerebellar cortex and the deep cerebellar nuclei.

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are located ventrally and lateral with respect to the pyramidal tract. The fibers of the hipoglossum nucleus
appear between the pyramids and the olivary complex.

Macroscopic structure  
The cerebellum is attached to the brainstem by three pairs of cerebellar bundles, the superior, middle and
inferior cerebellar peduncles. The superior cerebellar peduncle consists mainly of efferent fibers from the
cerebellum, however, the middle and inferior peduncles contain mainly afferent fibers in the cerebellum.
The cerebellum is a single mass in which it can be differentiate an odd midline band called vermis and
two lateral masses called cerebellar hemispheres. There is a smooth transition between the vermis and
the hemispheres called the paravermis. The cerebellum is crossed by transverse grooves that frame the

Macroscopic structure of the unfolded cerebellum. a) cerebellar vermis, d) lingula e) central lobe, f)
peak, g) slope, h) folium, i) tuber, j) pyramid, k) uvula, l) node, m) alar lobule, n) quadrates anterior
o) quadrates posterior, p) superior lunate, q) posterior lunate, r) aznterior digastric s) posterior
digastric, t) cerebellar tonsil, u) floculus, v) primary horizontal fissure, x) posteroinferior sulcus , y)
anterior lobe, z) posterior lobe, yy) floculo nodular lobe, ab) horizontal groove, ac) posterosuperior
groove, ad) prepiramidal groove
cerebellar laminae. Some grooves are more prominent what allows to distinguish the main cerebellar
lobes. In an unfolded representation two deep grooves, i.e. the primary sulcus and the posterolateral
sulcus, divides the cerebellum into three lobes. The area anterior to the primary sulcus is called the
anterior lobe, the area between the primary sulcus and the posterolateral sulcus is called the posterior lobe
and the area posterior to the posterolateral sulcuus is called the flocculo-nodular lobe. In each lobe, other
minor sulci makes possible to differentiate lesser divisions.
Classical nomenclatures associate the flocculo-nodular lobe to the area managing vestibular information
and it was refered to as archicerebellum. The anterior lobe was named paleocerebellum . Finally, the
posterior lobe was called neocerebellum. Although these divisions are highly questioned as a whole,
several aspects needs to be rescued and sharpen. The archicerebellum corresponds to the area related to
the vestibular system and is presently known as vestibulum-cerebellum. The area mostly related to spinal
afferents corresponds to midline areas of the vermis and paravermis of both paleocerebellum and
neocerebellum. This area is refered to as spinocerebellum. Finally, pontine afferents which relays
neocortex projections are directed mainly to the cerebellar hemispheres of the paleo- and neocerebellum
and are called as cerebro-cerebellum or ponto-cerebellum.

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Cytoarchitectonic structure  
The cerebellum consists of two structures, the cerebellar folia and the deep nuclei. The deep nuclei are
three pairs of cellular groups which appears in the middle of fiber tracts. From medial to lateral these
nuclei are the fastigium, the interpossitus and the dentate nuclei. These nuclei are mainly related with the
major divisions of the cerebellum. The fastigium is closer related to the vestibule-cerebellum, the
interpossitus is mostly related to the spinocderebellum and the dentate relates to pontocerebellum.

Transverse section of the rat cerebellum in

which are represented the main layers. a)
molecular layer, b) layer of Purkinje cells,
c) granular layer, d) white matter

Figure 3.- Transverse section of a rat brain at the pons revealing the horitzontal arrangement of the
laminaer. a) vermis, b) paravermis c) cerebellar hemispheres, d) floculus, e) mid cerebellar peduncle,
f) fibrous layer, g) granular layer, h) layer of Purkinje cells,i) molecular layer, j) superior cerebellar
peduncle. k) motor nucleus of the facial, l) locus coeruleus, m), spinal trigeminal nucleus, n) spinal
trigeminal tract, or) dorsal tegmental nucleus, p) statoacoustic nerve VIII, q) cochlear nuclei.

The folia cerebelosa are composed of a superficial cerebellar cortex and a deep medulla. The later
contains the fiber tracts of fibers entering or leaving the cortex. The cerebellar cortex has a more
complicated structure, it is composed of three layers: molecular, the Purkinje cell layer and granular cell
layer.
• The molecular layer is composed of the dendritic trees, bundles of fibers and dispersed cells. In
the deepest part of the molecular layer and in the cells, just over the limit with the Purkinje cell
layer it can be found the basket cells.
• The Purkinje cell layer forms a monolayer of cells. The dendritic trees of these cells is very
dense in arborization and successive branches form a plane that is perpendicular to the axis of
the folia cerebelossa. The axon leaves the neuronal body from its deepest part. Basket cells are
interposed between the arborizations of the Purkinje cells and in turn send axons which form
nests terminals surrounding the neuronal bodies of Purklinje cells.
• In the granular cell layer there are two types of neurons, granuklar cells and Golgi cells. The
granular cells are abundant and small s have only three to six short dendrites that end in a long
part that goes into a glomerulus which is formed surrounding a nerve terminal called the mossy
fiber. The axon leaving the granular cells ascends to the molecular layer where it branches into a
"T" shape giving rise to the parallel fibers that run longitudinally along the folia cerebelosa.
Golgi cells are formed by two tufted dendritic trees and a short axon entering the glomerulus.

Cerebelar circuits 
The cerebellum receives diffuse monoaminergic projections from the locus coeruleus and raphe nuclei,
also receives two types of fibers, climbing fibers and mossy fibers. The efferences from the cerebellar
folia arise fronm the Purkinje cells to the deep nuclei. Finally, these nuclei convey the efferent projections
of the cerebellum over brainstem and thalamic targets.

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 • The climbing fibers originate in the olivary bulbar nuclei. At the level of the deep nuclei they
give rise to a collateral which contact them. The main axonic branch continues until the
cerebellar cortex and brach surrounding the Purkinje cell body and dendritic arborization.

• The mossy fibers have different origins (reticular formation, red nucleus, vestibular nuclei,
nuclei of the pons ...). Mossy fibers branch profusely in the cerebellar white matter and its
terminal branches reach the granular layer where they finish in the core of the glomerulus. Each
glomerulus contains the terminal ends of a mossy fiber and dendrites of granular cells and
terminal axons of Golgi cells. All these structures are surrounded and isolated by surrounding
glial processes. Parallel fibers arise from granular cells and run over the molecular layer
contacting the dendritic trees of the Purkinje and basket cells exciting them (as parallel fibers use
glutamate as transmitter). One parallel fiber may excite a row one or two hundred Purkinje cells
and basket cells interposed between them. As the basket cells are inhibitory, the activation of a
parallel fiber causes the activation of a row of Purkinje cells and inhibition of surrounding rows
parallels to them. Finally, the Purkinje cells project to the deep nuclei. As Purkinje cells are
inhibitory their activation causes inhibition of the deep nuclei.
Thus, it must be concluded that in the cerebellum there is a reverberating circuit with multiple cycles of
activation and inhibition. Those circuits that when performing give rise to a successful motor outcome are
fixed through plastic changes and consolidate along time. This is the way for developing motor skills.

Diagram representing the elements of the

cerebellar plate and the flow of information
between the cellular elements. a) climbing fibers
from the inferior olive nuclei, b) mossy fibers
from other structures as pontine nuclei, reticular
formation, vestibular nuclei, c) granular cells, d)
parallel fibers, e) of Purkinje cells , f) basket
cells, g) Golgi cells, h) deep nuclei.

Systems and projections of the cerebellum 
The classical texts claim for the cerebellum to be composed of three systems related primarily to the
vestibular system, the other on the spinal somatomotor system and finally a third which lies under the
domain of the cerebral cortex. Initial connective data c appear that the three systems were almost
completely segregated, but the application of techniques of electrophysiology and finest track tracing
connections revealed that the three systems are superimposed with each other. However the analysis of
the projections can be done independently.

Vestibulo‐cerebellum 
The flocculo-nodular lobe is the main representative of the vestibulum in the cerebellar cortex and the
nucleus fastigium at the base.
In the coordination of any movement is particularly important information from the body position with
respect to the vertical (the utriculum and saculum afferents) and information from the dynamic

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movements of the head or body (semicircular ducts). This information is melted to others in order to
perform movements.
The vestibular ganglion cells send fibers directly to the components of the vestibule-cerebellum. In
addition, cells of the vestibular nuclei, specially the medial nucleus, relay vestibular information to
different areas of the cerebellum which include the vestibule-cerebellum. These fibers end as mossy
fibers. Both direct and indirect fibers pass through the body of the inferior cerebellar peduncle. The
vestibular system projects to the cerebellum of the same side (right to right and left to left). Also the
vestibule-cerebellum receives climbing fibers from the inferior olivary nuclei.
Mossy fibers do not send collaterals to the deep nuclei as the climbing fibers do. Most Purkinje cells
project to the nucleus fastigium but a few also project directly to the lateral vestibular nucleus. The
nucleuas fastigium projects to the medial and lateral. A major projection of the nucleus fastigium targets
the reticular formation at all levels but especially in the paramedian pontine reticular formation (PPRF).
This projection is bilateral and is in charge for controlling the visual gaze. The cerebellar projection to the
visuomotor nuclei keeps the gaze on a target.

Diagram representing the flow of information of the vestibulum-cerebellum. a) neuron of the

vestibular ganglion, b) medial vestibular nucleus, c) superior vestibular nucleus, d) lateral vestibular
nucleus, e) inferior vestibular nucleus, f) nodulus, g) flocsulus, h) fastigium nucleus, i) nucleus
interpossitus, j) dentate nucleus, k) uncinate tract (Russell), l) cerebello-reticular tracts, m) medial
vestibule-spinal tract, n) lateral vestibule-spinal tract, o) reticular formation

Spino‐cerebellum 
The spinocerebellum overlaps the vestibular information with proprioceptive information arising from
articular capsules, joints, tendons and muscular spindle t6herough the spinal cord. Thus while the
movements are performed, feed-back information from the position and strength of tendons and muscles
is arriving and monitoring to make it more accurate. Althuperior cerebellar peduncle. Then, inside the
cerebellum fibers cross again to arrive to the ipsilateral spinocerebellum.

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The main information arriving to the spinocerebellum arise from the ventral and dorsal spinocerebellar
tracts. These tracts carry proprioceptive information but also somatosensory cutaneous information of the
cutaneous surface of the body so that the spinocerebellum has a representation of the body and the degree
of tension of its components at the same time. The propiocepotive information reaches the nuclei at the
base of the dorsal horn (including the column of
Clarke). Propioceptive information relayed in the
spiunal cord nuclei is being sent to the cerebellum
through two pathways.
• The ventral spinocerebellar tract is composed
of fibers which cross to the contralateral side and
ascend to arrive to the From its origin, the fibers are
going up by dorsolateral funicle without crossing
itself and penetrate the cerebellum by the inferior
cerebellar peduncle. These fibers end as mossy fibers.
The core extends back only to the bottom of C8,
therefore, this treatment carries fibers of the trunk and
lower limb. This treatment adds fiber dprovinents
accessory cuneiform kernel.
• The dorsal spinocerebellar fibers ascend
directly to the ipsilateral spinocerebellum.
Over the spinocerebellar cortex two somatotopic
representations of the body can be found. In the
anterior representation the feet and distal parts of the
body appear in the anteriormost part corresponding
with the língula and the head over the folium. On the
second representation the head is represted over the
tuber and the feet in the nodule.
There is also a representation of afferent fibers of the
inferior olivary nuclei through climbing fibers.

Schema representing the information flow of
the spinocerebellum. a) proprioceptors of
muscles and joints, b) the direct dorsal
spino-cerebellar tract, c) gracilis and
cuneïforme bundles, d) the crossed ventral
spinocerebellar tract, e) cuneïforme nucleus,
f) juxtarestiformis tract, g) nuclus
interpositus, h) nucleuys dentatus, i) nucleus
fastigium, j) superior cerebellar peduncle, k)
decussatio of the superior cerebellar
peduncle, l) ventral lateral nnucleus of the
thalamus, m) red nucleus, n) reticular
formation, o) vestibular nuclei, p) vermis
and paravermis
Purkinje cells from the vermis project mainly to the
fastigium nucleus and those from the paravermis
project to the interpositus. Most of fastigial fibers are
integrated into the uncinatus tract which projects to
the contralateral reticular formation additional fibers
project to the vestibular nuclei. The nucleus
inteerpositus send fibers which course through the
superior cerebellar peduncle and at the level of the
midbrain crosse to the contralateral side to reach the
red nucleus. This nucleus in tur send alfibers which
again crosse to the ipsilateral side to form the
rubrospinal tract which courses to the spinal cord. A
part of the interpositus fibers also run sostrally to get
the thalamus.

Ponto cerebellum 
The ponto cerebellum is the main area of the posterior cerebellar hemispheres. Its main afferents comes
from the pointine nuclei. Pontine nuclei receive its main afferents from the four neocortical lobes (frontal,
temporal parietal and occipital). The most massive projections come from the precentral and postcentral
giri (areas 1,2,3,4,5 and 6 which correspond to the sensorimotor system). The coticopontine fibers have a

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topographic distribution along all the descending tract from the cortex. The frontopontine tract run in the
anterior arm of the internal capsule. In turn, the parieto-pontine, occipito-pontine, temporopontine and a
few frontopontine are consecutively arranged along the cerebral peduncles until arriving the pontine
nuclei. Pontine nuclei cross to the contralateral side to get the contralateral middle cerebellar peduncle to
project the deep dentate nucleus and cerebellar cortex.
Purkinje cells project to the dentate nucleus. In turn, this deep nucleus send their axons though the
superior cerebellar peduncle and project consecutively to the reticular formation, the red nucleus and
more importantly to the ventrolateral thalamic nucleus. This nucleus send projections back to the cortex.

Diagram representing the flow of information in the cerebro-cerebellum. a)

motor cortex, b) pontine nuclei, c) cerebellar hemispheres, d) dentate nucleus, e)
nucleus inteerpossitus, f) nucleus fastigium, g) red nucleus, h) ventral lateral
nucleus of the thalamus