The Cerebellum

ANAT 321

Descending projections from the motor cortex form the pyramidal

tract. The output from motor cortex is modulated by two
extrapyramidal structures: the cerebellum and the basal ganglia.

Basal
ganglia

Pyramidal tract

Thalamus

Cerebellum

Cerebellum: Compares differences between intention and

action, and makes appropriate adjustments in motor centers
of the cortex.
The cerebellar cortex consists of a large number of virtually
identical functional units, suggesting that it performs similar
functional operations on different inputs.
It receives massive input from sensory systems and from systems
involved in planning and execution of movement.
Its outputs project to premotor and motor systems of cerebral
cortex and brainstem.

Synaptic transmission in functional modules is subject to

modification (long term depression), which is thought to be
important for motor learning.

The cerebellum accounts for around 10% of the total

brain volume, but more than half of the brains neurons.

The surface of the cerebellum is covered with folds called

folia. The folia greatly increase the surface area of the
cerebellar cortex.

The cerebellum is divided into anatomically distinct lobes.

The cerebellum is connected to the brainstem through three large

fiber tracts called cerebellar peduncles.

Superior cerebellar peduncle

(outputs to brainstem and
thalamus)

Inferior cerebellar peduncle

inputs from spinal cord (directly
and through brainstem) and
vestibular nuclei.

Middle cerebellar peduncle

(inputs from cerebral cortex, by
way of pontine nuclei)

The Cerebellar Peduncles

Superior cerebellar
peduncle

Middle cerebellar
peduncle

Inferior cerebellar
peduncle

The cerebellum comprises a cortex and deep nuclei. Inputs to the

cerebellum project to both the cortex and the deep nuclei. Outputs
from the cortex project to the deep nuclei, which, in turn, form the
outputs of the cerebellum.
Cortex

Deep
nuclei

There are three deep cerebellar nuclei, called fastigial, interposed and
dentate.

The cerebellar cortex

The cerebellar cortex consists of five cell types, arranged

in three layers: the molecular layer, the Purkinje cell
layer and the granule cell layer.

The five cell types of neurons in the cerebellar cortex are

arranged in a highly ordered, repeating architecture.

The human cerebellum contains ~ 100 billion granule cells.

Purkinje cells form a monolayer above the granule cells.

Purkinje cell dendrites are two-dimensional

Synaptic organization of the basic cerebellar circuit module

Stellate cell

Parallel fiber

Golgi cell

Basket cell

Granule
cell

Mossy fiber

_ +

Climbing fiber

Purkinje cell

_
+
+

Inferior olive

Deep nuclei

Mossy fibers make excitatory connections with granule cells and with
neurons in the deep cerebellar nuclei. Granule cell axons form parallel
fibers in the molecular layer, which make excitatory synapses on
Purkinje cells.

Parallel fiber

Purkinje cells

Granule
cell

Mossy fiber

Deep nuclei

Parallel fibers are densely packed in the molecular layer. A single parallel
fiber synapses with hundreds of Purkinje cells. A single Purkinje cell
receives excitatory input from hundreds of thousands of parallel fibers.

Parallel fibers

Bundles of parallel fibers, called beams, run transversely and excite the
dendrites of Purkinje cells and basket cells. The basket cells then inhibit the
Purkinje cells flanking the parallel fiber beam.

Mossy fiber activity produces a steady stream of simple spikes in Purkinje

cells, at rates of up to several hundred simple spikes per second. The
frequency of simple spikes is strongly modulated by sensory stimuli and
voluntary movements.

Simple spikes

Each Purkinje cell receives powerful excitatory input from a single climbing
fiber. Each climbing fiber innervates 5 10 Purkinje cells. Climbing fibers
originate in the inferior olive. They send collateral projections to the deep
nuclei

Climbing fiber

Purkinje cell

Inferior olive
From inferior olive

Deep nuclei

Climbing fibers generate complex spikes at fairly regular intervals

(1-3 Hz). The frequency of complex spikes is not strongly
modulated by sensory or motor activity.

Simple spikes

Complex spike

Projection neurons in the inferior olive are electrically coupled,

which may enable arrays of Purkinje cells to fire synchronous
complex spikes.

The grid represents the spatial locations of 29 Purkinje cells from which complex
spikes were recorded while the rat was licking. The cells in red fired synchronously
at one time; those in blue fired synchronously at another time; cells represented
by open circles were not synchronized.

Purkinje cells make inhibitory GABAergic synapses on neurons in the deep

nuclei. Excitatory drive on deep nuclear neurons comes from the mossy
fiber and climbing fiber collaterals. The neurons of the deep nuclei make
excitatory connections with neurons in the brainstem and thalamus.
Parallel fiber

Granule
cell

Mossy fiber

Climbing fiber

Purkinje cell

_
+
+

Deep nuclei

A summary of the cerebellar circuit

Long term depression

Synapses in which simple and complex spikes occur concurrently have
reduced efficacy. This is a form of synaptic plasticity called long-term
depression. It is thought to be important for motor learning.

The cerebellum participates in motor learning, e.g. subjects playing darts

while wearing prism glasses.

The vestibulocerebellum comprises the flocculonodular lobe. It

receives inputs from the semicircular canals and the otolith organs. It
is involved in balance and eye movements.

The spinocerebellum consists of the vermis and the intermediate

hemispheres. It receives massive sensory (mainly proprioceptive) input
in the form of mossy fibers, either directly from the spinal cord or
from the cord by way of the brainstem reticular formation.

The vermis projects to the fastigial nucleus, which in turn projects to

the vestibular nuclei and the reticular formation. These projections
arem mainly involved in posture.

The Intermediate hemispheres project through the interposed nuclei

to primary motor cortex (via the VL and VA thalamic nuclei) and to the
magnocellular region of the red nucleus.

The spinocerebellum uses feedforward mechanisms to

regulate movement. Damage to these feedforward systems
produce intention tremors when patients reach for objects.

Arm position
Control

Inactivated deep nuclei

Biceps
Triceps
0 msec

700 msec

The lateral hemispheres form the cerebrocerebellum. This region is much

larger in humans than in other primates. It is involved in the planning,
programming and timing of complex, precisely coordinated movement
sequences. Lateral cerebellar lesions produce decomposition of
movement.

Activity in the dentate nucleus is greater when the

subject is mentally active during movement.

(A,C) Subjects first passively experienced sandpaper rubbed across the fingers (A) and
then were asked to discriminate the degree of roughness of the sandpaper (B). (B,D)
Subjects were asked to lift and drop a series of objects (C) and then had to identify the
felt object from a similar group near the other hand (D).

Motor cortex-cerebello-rubrocerebellar loops may be

functional units involved in mental rehearsal and motor
learning.
Premotor and
motor cortex
Cerebellar cortex
LV/AV thalamus

Parvocellular
red nucleus

Dentate nucleus

Inferior olive

Damage to the cerebellum causes characteristic disorders

Hypotonia: diminished resistance to passive limb displacement.

Ataxia: Abnormalities in execution of voluntary movements. Lack of
coordination.
Dysdiadochokinesia: Loss of ability to produce a regular rhythm or consistent
force in repetitive movements.
Intention tremors: Tremor during movement. Most noticeable at the end of a
movement.