Cerebellum

GROSS APPEARANCE OF THE CEREBELLUM

 The cerebellum is situated in the posterior

cranial fossa and is covered superiorly by the
tentorium cerebelli
 It is the largest part of the hindbrain and lies
posterior to the fourth ventricle, the pons, and
the medulla oblongata
 The cerebellum is somewhat ovoid in shape
and constricted in its median part.
 It consists of two cerebellar hemispheres joined
by a narrow median vermis
 The cerebellum is connected to the posterior
aspect of the brainstem by three symmetrical
bundles of nerve fibers called the superior,
middle, and inferior cerebellar peduncles
 The cerebellum is divided into three main lobes:
o Anterior
o Middle
o flocculonodular lobe

 The anterior lobe may be seen on the superior

surface of the cerebellum and is separated from
the middle lobe by a wide V-shaped fissure
called the 
o primary fissure
 The middle lobe (sometimes called the
posterior lobe), which is the largest part of the
cerebellum, is situated between the primary
and uvulonodular fissures
 The flocculonodular lobe is situated posterior
to the uvulonodular fissure
  A deep horizontal fissure that is found along
the margin of the cerebellum separates the
superior from the inferior surfaces
o it is of no morphologic or functional
significance

STRUCTURE OF THE CEREBELLUM Molecular Layer

 The cerebellum is composed of an outer  The molecular layer contains two types of
covering of gray matter called the cortex and neurons: the outer stellate cell and the inner
inner white matter. basket cell
 Embedded in the white matter of each  These neurons are scattered among dendritic
hemisphere are three masses of gray matter arborizations and numerous thin axons that
forming the intracerebellar nuclei run parallel to the long axis of the folia
 Neuroglial cells are found between these
Structure of the Cerebellar Cortex structures
 The cerebellar cortex can be regarded as a large Purkinje Cell Layer
sheet with folds lying in the coronal or
transverse plane.  The Purkinje cells are large Golgi type I
o Each fold or folium contains a core of neurons.
white matter covered superficially by  flask shaped and are arranged in a single layer
gray matter  In a plane transverse to the folium, the
 section made through the cerebellum parallel dendrites of these cells are seen to pass into the
with the median plane divides the folia at right molecular layer, where they undergo profuse
angles, and the cut surface has a branched branching
appearance, called the arbor vitae o The primary and secondary branches
 The gray matter of the cortex throughout its are smooth,and subsequent branches
extent has a uniform structure. are covered by short, thick dendritic
 It may be divided into three layers: spines.
o external layer, the molecular layer  It has been shown that the
 External = Molecular spines form synaptic contacts
o a middle layer, the Purkinje cell layer with the parallel fibers derived
 Middle = purkinje from the granule cell axons
o an internal layer, the granular layer  At the base of the Purkinje cell, the axon arises
 Internal = granular and passes through the granular layer to enter
the white matter
o On entering the white matter, the axon
acquires a myelin sheath, and it
terminates by synapsing with cells of
one of the intracerebellar nuclei
o Collateral branches of the Purkinje axon
make synaptic contacts with the
dendrites of basket and stellate cells of
the granular layer in the same area or in
distant folia.
 A few of the Purkinje cell axons pass directly to
end in the vestibular nuclei of the brainstem.
Granular Layer  The cortex of the vermis influences the 
o movements of the long axis of the
 The granular layer is packed with small cells
body, namely, the:
with densely staining nuclei and scanty
 neck, the shoulders, the thorax,
cytoplasm
the abdomen, and the hips
 Each cell gives rise to four or five dendrites,
which make clawlike endings and have synaptic
contact with mossy fiber input
 The axon of each granule cell passes into the
molecular layer, where it bifurcates at a T
junction, the branches running parallel to the
long axis of the cerebellar folium

 Immediately lateral to the vermis is a so-called

intermediate zone of the cerebellar
hemisphere.
o This area has been shown to control the
muscles of the distal parts of the limbs,
especially the hands and feet
 The lateral zone of each cerebellar hemisphere
appears to be concerned with the:
o planning of sequential movements of
the entire body and is involved with the
conscious assessment of movement
 These fibers, known as parallel fibers, run at errors.
right angles to the dendritic processes of the
Purkinje cells.
o Most of the parallel fibers make
synaptic contacts with the spinous
processes of the dendrites of the
Purkinje cells
o Neuroglial cells are found throughout
this layer
 Scattered throughout the granular layer are
Golgi cells. Intracerebellar Nuclei
o Their dendrites ramify in the molecular
layer, and their axons terminate by  Four masses of gray matter are embedded in
splitting up into branches that synapse the white matter of the cerebellum on each side
with the dendrites of the granular cells of the midline
 From lateral to medial, these nuclei are the
Functional Areas of the Cerebellar Cortex dentate, the emboliform, the globose, and the
fastigial.
 divide up the cerebellar cortex into three
o D-E-G-F
functional areas.
  The dentate nucleus  o They enter the cerebellum mainly
o is the largest of the cerebellar nuclei. It through the inferior and middle
has the shape of a crumpled bag with cerebellar peduncles.
the opening facing medially  The efferent fibers constitute the output of the
o The interior of the bag is filled with cerebellum and commence as the axons of the
white matter made up of efferent Purkinje cells of the cerebellar cortex.
fibers that leave the nucleus through o The great majority of the Purkinje cell
the opening to form a large part of the axons pass to and synapse with the
superior cerebellar peduncle. neurons of the cerebellar nuclei
 The emboliform nucleus (fastigial, globose, emboliform, and
o is ovoid and is situated medial to the dentate)
dentate nucleus, partially covering its o The axons of the neurons then leave
hilus the cerebellum.
 The globose nucleus o A few Purkinje cell axons in the
o consists of one or more rounded cell flocculonodular lobe and in parts of the
groups that lie medial to the vermis bypass the cerebellar nuclei and
emboliform nucleus leave the cerebellum without synapsing
 The fastigial nucleus  Fibers from the dentate, emboliform, and
o lies near the midline in the vermis and globose nuclei leave the cerebellum through
close to the roof of the fourth ventricle; the superior cerebellar peduncle
it is larger than the globose nucleus o DEG  superior cerebellar peduncle
 The intracerebellar nuclei are composed of  Fibers from the fastigial nucleus leave through
large, multipolar neurons with simple the inferior cerebellar peduncle.
branching dendrites. o Fastigial  inferior cerebellar peduncle
 The axons form the cerebellar outflow in the
CEREBELLAR CORTICAL MECHANISMS
superior and inferior cerebellar peduncles.
 The climbing and the mossy fibers constitute
White Matter
the two main lines of input to the cortex and
 There is a small amount of white matter in the are excitatory to the Purkinje cells
vermis  The climbing fibers are the terminal fibers of
 arbor vitae - resembles the trunk and branches the olivocerebellar tracts
of a tree o They pass through the granular layer of
 The white matter is made up of three groups of the cortex and terminate in the
fibers: molecular layer by dividing repeatedly
o Intrinsic o Each climbing fiber wraps around and
o Afferent makes a large number of synaptic
o Efferent contacts with the dendrites of a
 The intrinsic fibers do not leave the cerebellum Purkinje cell.
but connect different regions of the organ  A single Purkinje neuron makes synaptic
o Some interconnect folia of the contact with only one climbing fiber.
cerebellar cortex and vermis on the  However, one climbing fiber makes contact
same side with 1 to 10 Purkinje neurons
o others connect the two cerebellar o A few side branches leave each climbing
hemispheres together. fiber and synapse with the stellate cells
 The afferent fibers form the greater part of the and basket cells
white matter and proceed to the cerebellar  The mossy fibers are the terminal fibers of all
cortex other cerebellar afferent tracts.
o They have multiple branches and exert
a much more diffuse excitatory effect
 o A single mossy fiber may stimulate Cerebellar Cortical Neurotransmitters
thousands of Purkinje cells through the
 excitatory climbing and mossy afferent fibers
granule cells
use glutamate (gammaaminobutyric acid
 stellate, basket, and Golgi cells 
[GABA]) as the excitatory transmitter on the
o they serve as inhibitory interneurons
dendrites of the Purkinje cells.
o they not only limit the area of cortex
 other afferent fibers entering the cortex
excited but influence the degree of
liberate norepinephrine and serotonin at their
Purkinje cell excitation produced by the
endings that possibly modify the action of the
climbing and mossy fiber input
glutamate on the Purkinje cells.
o By this means, fluctuating inhibitory
impulses are transmitted by the
Purkinje cells to the intracerebellar
nuclei, which, in turn, modify muscular
activity through the motor control areas
of the brainstem and cerebral cortex.
o It is thus seen that the Purkinje cells
form the center of a functional unit of
the cerebellar cortex

Cerebellar Peduncles

 The cerebellum is linked to other parts of the

central nervous system by numerous efferent
and afferent fibers that are grouped together
on each side into three large bundles, or
peduncles
o The superior cerebellar peduncles
connect the cerebellum to the midbrain
Intracerebellar Nuclear Mechanisms
o the middle cerebellar peduncles
 The deep cerebellar nuclei receive afferent connect the cerebellum to the pons
nervous information from two sources: o the inferior cerebellar peduncles
o the inhibitory axons from the Purkinje connect the cerebellum to the medulla
cells of the overlying cortex oblongata
o the excitatory axons that are branches
of the afferent climbing and mossy
CEREBELLAR AFFERENT FIBERS
fibers that are passing to the overlying
cortex. Cerebellar Afferent Fibers From the Cerebral Cortex
 In this manner, a given sensory input to the
 The cerebral cortex sends information to the
cerebellum sends excitatory information to the
cerebellum by three pathways:
nuclei, which a short time later receive cortical
o the corticopontocerebellar pathway
processed inhibitory information from the
o the cerebro-olivocerebellar pathway
Purkinje cells.
o The cerebroreticulocerebellar pathway
 Efferent information from the deep cerebellar
nuclei leaves the cerebellum to be distributed
to the remainder of the brain and spinal cord.
 Cerebroreticulocerebellar Pathway

 The corticoreticular fibers arise from nerve cells

from many areas of the cerebral cortex,
particularly the sensorimotor areas
 They descend to terminate in the reticular
formation on the same side and on the
opposite side in the pons and medulla
 The cells in the reticular formation give rise to
the reticulocerebellar fibers that enter the
cerebellar hemisphere on the same side
through the inferior and middle cerebellar
peduncles.
 This connection between the cerebrum and the
cerebellum is important in the control of
voluntary movement.
 Information regarding the initiation of
movement in the cerebral cortex is probably
transmitted to the cerebellum 
o so that the movement can be
monitored and appropriate
adjustments in the muscle activity can
Corticopontocerebellar Pathway be made.

 The corticopontine fibers arise from nerve cells

in the frontal, parietal, temporal, and occipital Cerebellar Afferent Fibers From the Spinal Cord
lobes of the cerebral cortex and descend
 The spinal cord sends information to the
through the corona radiata and internal capsule
cerebellum from somatosensory receptors by
and terminate on the pontine nuclei
three pathways 
 The pontine nuclei give rise to the transverse
o the anterior spinocerebellar tract
fibers of the pons, which cross the midline and
o the posterior spinocerebellar tract
enter the opposite cerebellar hemisphere as
o the cuneocerebellar tract.
the 
o middle cerebellar peduncle Anterior Spinocerebellar Tract
Cerebro-olivocerebellar Pathway  The axons entering the spinal cord from the
posterior root ganglion terminate by synapsing
 The cortico-olivary fibers arise from nerve cells
with the neurons in the nucleus dorsalis
in the frontal, parietal, temporal, and occipital
(Clarke’s column) at the base of the posterior
lobes of the cerebral cortex and descend
gray column.
through the corona radiata and internal capsule
o Most of the axons of these neurons
to terminate bilaterally on the inferior olivary
cross to the opposite side and ascend
nuclei
as the anterior spinocerebellar tract in
 The inferior olivary nuclei give rise to fibers that
the contralateral white column
cross the midline and enter the opposite
o some of the axons ascend as the
cerebellar hemisphere through the inferior
anterior spinocerebellar tract in the
cerebellar peduncle.
lateral white column of the same side
 These fibers terminate as the climbing fibers in
 The fibers enter the cerebellum through the
the cerebellar cortex.
superior cerebellar peduncle and terminate as
mossy fibers in the cerebellar cortex
 Collateral branches that end in the deep Posterior Spinocerebellar Tract
cerebellar nuclei are also given off.
 The axons entering the spinal cord from the
o It is believed that those fibers that cross
posterior root ganglion enter the posterior
over to the opposite side in the spinal
gray column and terminate by synapsing on the
cord cross back within the cerebellum.
neurons at the base of the posterior gray
 The anterior spinocerebellar tract is found at all
column.
segments of the spinal cord, and its fibers
o These neurons are known collectively as
convey muscle joint information from the
the nucleus dorsalis (Clarke’s column).
muscle spindles, tendon organs, and joint
 The axons of these neurons enter the
receptors of the upper and lower limbs.
posterolateral part of the lateral white column
 It is also believed that the cerebellum receives
on the same side and ascend as the posterior
information from the skin and superficial fascia
spinocerebellar tract to the medulla oblongata
by this tract
o Here, the tract enters the cerebellum
through the inferior cerebellar
peduncle and terminates as mossy
fibers in the cerebellar cortex
o Collateral branches that end in the deep
cerebellar nuclei are also given off
 The posterior spinocerebellar tract receives
muscle joint information from the muscle
spindles, tendon organs, and joint receptors of
the trunk and lower limbs.

Cuneocerebellar Tract

 These fibers originate in the nucleus cuneatus

of the medulla oblongata and enter the
cerebellar hemisphere on the same side
through the inferior cerebellar peduncle
 The fibers terminate as mossy fibers in the
cerebellar cortex.
 The cuneocerebellar tract receives muscle joint
information from the muscle spindles, tendon
organs, and joint receptors of the upper limb
and upper part of the thorax
Cerebellar Afferent Fibers From the Vestibular Nerve cerebellar peduncle and again in the rubrospinal
tract close to its origin
 The vestibular nerve receives information from
 By this means, the globose and emboliform
the inner ear concerning motion from the
nuclei influence motor activity on the same side
semicircular canals and position relative to
of the body.
gravity from the utricle and saccule.
 The vestibular nerve sends many afferent fibers
directly to the cerebellum through the inferior
cerebellar peduncle on the same side.
o Vestibular nerve = Afferent directly via
the inferior cerebellar peduncle
 All the afferent fibers from the inner ear
terminate as mossy fibers in the
flocculonodular lobe of the cerebellum.

CEREBELLAR EFFERENT FIBERS

 The entire output of the cerebellar cortex is

through the axons of the Purkinje cells
o Most of the axons of the Purkinje cells
end by synapsing on the neurons of the
deep cerebellar nuclei
o The axons of the neurons that form the Dentothalamic Pathway
cerebellar nuclei constitute the efferent
outflow from the cerebellum.  Axons of neurons in the dentate nucleus travel
 The efferent fibers from the cerebellum through the superior cerebellar peduncle and
connect with the: cross the midline to the opposite side in the
o red nucleus decussation of the superior cerebellar
o thalamus peduncle
o vestibular complex  The fibers end by synapsing with cells in the
o reticular formation contralateral ventrolateral nucleus of the
thalamus.
Globose-Emboliform-Rubral Pathway o The axons of the thalamic neurons
 Axons of neurons in the globose and ascend through the internal capsule and
emboliform nuclei travel through the superior corona radiata and terminate in the
cerebellar peduncle and cross the midline to primary motor area of the cerebral
the opposite side in the  cortex
o decussation of the superior cerebellar  By this pathway, the dentate nucleus can
peduncles influence motor activity by acting on the motor
 The fibers end by synapsing with cells of the neurons of the opposite cerebral cortex
contralateral red nucleus, which give rise to o impulses from the motor cortex are
axons of the rubrospinal tract transmitted to spinal segmental levels
 Thus, it is seen that this pathway crosses twice, through the corticospinal tract.
once in the decussation of the superior  Remember that most of the fibers of the
corticospinal tract cross to the opposite side in
 the decussation of the pyramids or later at the FUNCTIONS OF THE CEREBELLUM
spinal segmental levels.
 The cerebellum receives afferent information
o Thus, the dentate nucleus is able to
concerning voluntary movement from the
coordinate muscle activity on the same
cerebral cortex and from the muscles, tendons,
side of the body
and joints
Fastigial Vestibular Pathway  It also receives information concerning balance
from the vestibular nerve and possibly
 The axons of neurons in the fastigial nucleus
concerning sight through the tectocerebellar
travel through the inferior cerebellar peduncle
tract.
and end by projecting on the neurons of the
 All this information is fed into the cerebellar
lateral vestibular nucleus on both sides
cortical circuitry by the mossy fibers and the
o Remember that some Purkinje cell
climbing fibers and converges on the Purkinje
axons project directly to the lateral
cells
vestibular nucleus
 The axons of the Purkinje cells project with few
 The neurons of the lateral vestibular nucleus
exceptions on the deep cerebellar nuclei.
form the vestibulospinal tract.
 The output of the vermis projects to the
 The fastigial nucleus exerts a facilitatory
fastigial nucleus
influence mainly on the ipsilateral extensor
o intermediate regions of the cortex
muscle tone.
project to the globose and emboliform
Fastigial Reticular Pathway nuclei
o output of the lateral part of the
 The axons of neurons in the fastigial nucleus cerebellar hemisphere projects to the
travel through the inferior cerebellar peduncle
dentate nucleus
and end by synapsing with neurons of the
 A few Purkinje cell axons pass directly out of
reticular formation
the cerebellum and end on the lateral
 Axons of these neurons influence spinal vestibular nucleus in the brainstem
segmental motor activity through the
 It is now generally believed that the Purkinje
reticulospinal tract
axons exert an inhibitory influence on the
neurons of the cerebellar nuclei and the lateral
vestibular nuclei.
 The cerebellar output is conducted to the sites
of origin of the descending pathways that
influence motor activity at the segmental spinal
level
 In this respect, the cerebellum has no direct
neuronal connections with the lower motor
neurons but exerts its influence indirectly
through the cerebral cortex and brainstem.
 Cerebellum functions as a coordinator of
precise movements by continually comparing
the output of the motor area of the cerebral
cortex with the proprioceptive information
received from the site of muscle action
o it is then able to bring about the
necessary adjustments by influencing
the activity of the lower motor neurons
o This is accomplished by controlling the
timing and sequence of firing of the
alpha and gamma motor neurons.
 It is also believed that the cerebellum can send
back information to the motor cerebral cortex
to inhibit the agonist muscles and stimulate
the antagonist muscles, thus limiting the extent
of voluntary movement.