Cerebellum

Welke, Ph.D.
 Objectives
*Know the lobular and well as zonular organization of the cerebellum
and how that relates to function
*Know the deep cerebellar nuclei
*Understand the three layered cerebellar architecture
*Be able to basically describe the local circuitry and how it
communicates with the deep cerebellar nuclei.
*Be able to describe the afferent inputs into the cerebellum and
which inputs are mossy or climbing fibers. This includes knowing
the anterior spinocerebellar tract, posterior spinocerebellar tract
and cuneocerebellar tract.
*Understand the big picture to the three circuits: spinocerebellum,
pontocerebellum and vestibulocerebellum. What are they modulating
and what are their major inputs?
*Understand the difference between truncal and appendicular
ataxia, where a lesion would have to occur to produce each and what
test would you use to determine either.
 You should be familiar with these terms:

Posture the position of the body

Station stance, i.e. manner of standing
Gait Manner or style of walking
Akinesia absence (or poor) movement
Dyskinesia fragmentary or incomplete movements
Dystonia disordered tonicity of muscle
Dysmetria improper measuring of distance in
muscular acts
Dysdiadochokinesia impairment of ability to
perform rapid alternating movements
Tremor - rhythmic involuntary movement
Ataxia - lack of finely tuned muscular movements
involved in postural controlinability to coordinate
voluntary movements
 1-LOBES
2-LOCAL CEREBELLAR CORTICAL
CIRCUITRY
3-ASCENDING TRACTS (3) & SENSORY
INPUTS
3-DEEP CEREBELLAR NUCLEI
4-MODULATION OF MOTOR TRACTS &
THEIR MOTOR COROLLARIES
 Arbor Vitae -
pattern of
white matter

Folia
Primary fissure
separates
Anterior &
Posterior Lobes,
which make up
the body of the
cerebellum

Vermis =

Posterolateral
fissure
separates
body from
flocculonodular
lobe
 The 3 lobes are further classified based on their
developmental origins and functions:
______________________
1-Anterior Lobe = Spinocerebellum = Paleocerebellum
-Adjusts ongoing movements & regulates muscle tone
-Main input is the Spinocerebellar Tracts
2-Posterior Lobe = Pontocerebellum = Neocerebellum
-Coordinates planning of movements
-Main input is from sensory & motor cortices
3-Flocculo-nodular Lobe=Vestibulocerebellum=Archicerebellum
-Controls balance and eye movements
-Main inputs are the semicircular canals and the
vestibular nuclei
Deep Cerebellar Nuclei Relate to Function

Fastigial Nucleus
-Flocculo-nodular Lobe - Vestibulocerebellum
Globose & Emboliform Nucleus
-Anterior Lobe - Spinocerebellum
Dentate Nucleus
-Posterior Lobe - Pontocerebellum
Cerebellar Inputs

Cerebellar Outputs
 Cerebellar Cortex Architecture
3 Layers:
1-Molecular Layer (outermost layer)
2-Purkinje Cell Layer
3-Granule Cell Layer (innermost layer)
5 Different Neuronal Types:
1-Stellate Cells
2-Basket Cells
Local circuit
3-Purkinje Cells - output neuron neurons
4-Golgi Cells
5-Granule Cells
Granule Cell Layer - granule & golgi cells
Purkinje Cell Layer - Purkinje Cells
Molecular Layer - Stellate & Basket Cells
-both cell types are inhibitory interneurons
 How the local circuitry works: Cells of the
cerebellar cortex
Granule cell parallel fibers are:
-excitatory on Basket, Stellate & Golgi cells
-excitatory on Purkinje cell dendrites in
molecular layer
Basket cells are:
-inhibitory on Purkinje cell soma in Purkinje layer
Stellate cells are:
-inhibitory on Purkinje cell dendrites in
molecular layer
Golgi cells are:
-inhibitory on mossy fibers in granule cell layer
Purkinje cells are:
-inhibitory on the DCN (use GABA)
-MAJOR OUTPUT NEURON OF THE
CEREBELLAR CORTEX!
 How the local circuitry works: 2 Inputs into
the Cerebellum

Afferents to the cerebellum arise from

numerous sources, however they enter the
cerebellum and terminate in 2 ways:
1-Mossy Fibers
-terminate as excitatory synapses with
dendrites of the granule cells
2-Climbing Fibers
-from inferior olive climb the dendrites
of Purkinje cells & terminate as excitatory
synapses

*Both types of fibers also terminate on

the DCN
 Local Cortical Circuitry

(+)
(+)
Parallel Fiber
(+)

(-)
MOLECULAR
LAYER

Stellate Cell

Basket Cell

PURKINJE
Purkinje Cell

LAYER
(-)

(+)

GRANULE
Granule
LAYER
(+)
Cell

Golgi Cell

(-)
(+)
(-)

Mossy Fibers
(+)
DCN

Climbing Fibers

 Output of the
DCN is the result
DCN and their of the balance of
excitatory excitation from
afferents mossy & climbing
constitute a deep fibers and
excitatory loop inhibition by the
whose output is Purkinje cells
shaped by a
cortical inhibitory
loop that inverts
the sign of the
input signals. The
purkinje neuron
output to the DCN
cell therefore
generates an error
correction signal
that can modify
movements. The
climbing fibers
modify the
efficacy of the
parallel fiber-
Purkinje cell
connection,
producing long-
term changes in
cerebellar output.
(Purves, 2008)
 Principles of Cerebellar Function

1-Functions are IPSILATERAL

-if an afferent input to the cerebellum arises from one
side of the body or world , it terminates in the same
side of the cerebellum
2-Architecture/circuitry of the cerebellar cortex is constant
-different functions of the different lobes are a result
of afferent/efferent input
3-Cerebellar Dysfunction
-Lesions limited to the cerebellum DO NOT produce
paralysisthey produce disequilibrium, ataxia,
dysmetria and changes in muscle tone
 What does it do?

-Evaluates differences between intention and action and adjusts operations
in the motor cortex and brainstem while a movement is in progress and also
during repetitions of the same movement

How does it do it?

1-Receives information about commands & feedback signals associated with
the execution of movement
2-Output from the cerebellum is focused on centers that control motor
neurons (motor cortices & brainstem)
3-Cerebellar circuits can be modifiedimportant for motor adaptation and
learning

 3 Ascending Pathways

1-Anterior Spinocerebellar Pathway
-input from golgi tendon organs & spinal
interneurons, cutaneous receptors & fibers of
descending tracts in LOWER (distal) extremitiesthis
tract is more related to intended movement

2-Dorsal Spinocerebellar Pathway

-conveys signals from muscle spindles & golgi
tendon organs - info about individual muscles in
the LOWER (proximal) extremities

3-Cuneocerebellar Pathway
-conveys signals from muscle spindles & golgi
tendon organs - info about individual muscles in
the UPPER extremities
Anterior Spinocerebellar
-Transmits GTO afferent information
from distal lower limbs to cerebellum
-information about intended movement
from lower extremities

1st Order Neuron = DRG

- Fibers travel via Fasiculus Gracilis
to
2nd Order Neuron = Spinal Border Cells
- Fibers cross in Anterior White
Commissure (AWC), ascend via the
Anterior Spinocerebellar Tract (in the
anterior lateral funiculus), cross again
in the brainstem, and pass through the
Superior Cerebellar Peduncle (SCP) to
the Cerebellum

DRG -> (Fasciculus gracilis) ->

Spinal Border Cells -> (Anterior
Spinocerebellar Tract)-> thru SCP
-> Cerebellum
Dorsal Spinocerebellar
-Transmits muscle spindle and GTO
afferents information from trunk
and proximal lower limb to
cerebellum

1st Order Neuron = DRG

- Fibers travel via Fasiculus Gracilis
to
2nd Order Neuron = Clarke s
Nucleus (Column) (T1-L2)
-Fibers travel rostrally via the Dorsal
Spinocerebellar Tract (in the lateral
funiculus) through the Inferior
Cerebellar Peduncle (ICP) and into the
Cerebellum

DRG -> (Fasciculus gracilis) ->

Clarke s Nucleus -> (Dorsal
Spinocerebellar Tract) -> thru ICP
-> Cerebellum
 Cuneocerebellar
-Transmits muscle spindle and GTO
afferents information from the
upper limb & neck

1st Order Neuron = DRG

- Fibers travel via Fasciculus
Cuneatus to

2nd Order Neuron = Lateral

(external) Cuneate Nucleus
- Fibers travel rostrally via the
Cuneocerebellar Tract and through
the Inferior Cerebellar Peduncle
(ICP) to the Cerebellum

DRG -> (Fasiculus Cuneatus) ->

Lateral (external) Cuneate Nucleus
-> (Cuneocerebellar Tract) -> thru
ICP -> Cerebellum
Spinocerebellar Tracts in Spinal Cord
 General Plan of Cerebellar Pathways
1-Corollary Motor Discharge
-cerebellum receives input from motor
structures
-motor inputs convey the intended results
or goals of the descending motor outputs
2-Sensory Feedback
-cerebellum receives input that conveys
exteroceptive & proprioceptive info about
the results - actual effect of the motor
output on both muscles & the relationship to
the external world
3-Modulatory Output to Motor Centers
-Cerebellum the COMPARES the intended
movement and actual motor response that
results from motor pathways
-Cerebellum then computes adjustments
needed to the motor systems to produce
smooth synergy of motor output we label as
COORDINATION!
 Red Nucleus

Anterior
Lobe of
Central Cerebellum
Tegmental
Tract

Rubrospinal
Tract

Posterior
Spinocerebellar
or
Cuneocerebellar
Tract
 Spinocerebellar Overview
-Afferents from muscle spindles/joint receptors ascend on the
Spinocerebellar tracts and travel through the inferior cerebellar
peduncle (ICP-Dorsal Spinocerebellar or Cuneocerebellar) or
superior cerebellar peduncle (SCP-Anterior Spinocerebellar) to
reach the anterior lobe/intermediate zone.
-Corollary motor fibers also enter the anterior lobe from the
Rubrospinal Tract.
-The cerebellum then compares the intended output of the
Red Nucleus with the state of the muscles and computes the
necessary corrections between the intended Rubrospinal
output and the state of the muscle, tendons & joints
-The anterior lobe projects to the Globose & Emboliform nuclei
and they send efferents out the Superior Cerebellar Peduncle
(SCP) and across the midline to the opposite Red Nucleus to
modulate the output of the rubrospinal tract or other lateral
motor system pathways.

 Pontocerebellar

VL

Posterior
Lobe of
Pontocerebellar

Cerebellum
Fibers

Corticospinal


Tract
 Pontocerebellar Overview
-Afferent sensory information from the cortex (Occipital,
Frontal, Parietal & Temporal Lobes) enters the posterior lobe.
-Corollary motor fibers also enter the posterior lobe from the
Corticospinal Tract. These efferents cross the midline and reach
the cortex of the posterior lobe.
-The cerebellum analyzes the sensory info and what effect that
may have on any voluntary movement along with the intended
Corticospinal output.
-Posterior lobe projects to the dentate nucleus, which then
projects through the superior cerebellar peduncle, crosses the
midline, synapses on VL. VL then projects to motor & premotor
cortices for modulation of the Corticospinal Tract

Flocculo-nodular
Lobe of Cerebellum

Vestibulospinal


Tract
 Vestibulocerebellar Overview
-Afferent sensory input from the vestibular apparatus enters
the cerebellum through the inferior cerebellar peduncle (ICP)
and projects to the Flocculo-nodular Lobe.
-Corollary motor fibers also enter the Flocculo-nodular Lobe
from the Vestibulospinal Tract.
-The cerebellum then compares the vestibular input with the
intended output of the Vestibulospinal Tract and computes
the required corrections.
-Flocculo-nodular lobe projects to the Fastigial Nucleus and
Fastigial Nucleus axons reach the Vestibular Nuclei to regulate
the output of the Vestibulospinal Tract (equilibrium) or other
medial motor system pathways.

 A Note on Pathways

-The pathways discussed are not isolated from one another.
There is considerable overlap, reflecting the action of the
cerebellum in coordinating motor centers.
-There are other connections (tectospinal system & cranial
nerve nuclei III & V) but the specifics are not well understood.
-In any event:
-All the systems are necessary for the cerebellum to
effectively and simultaneously coordinate all descending
motor outputs

 Clinical Correlations

Ataxia = disordered contractions of agonist & antagonist muscles &
lack of coordination between movements at different joints.
Truncal Ataxia
-lesions to vermis / flocculonodular lobe
-loss of equilibrium, wide stance, problems balancing, veering to
one side
-patients have an unsteady drunklike gait
-Tests
-Tandem Gait Test- Walk the line patients fall or
deviate toward the side of the lesion during
walking
-Romberg test is NOT a test of cerebellar function,
but of the proprioceptive system in the dorsal columns a
patient stands with the feet together and the eyes closed.
Swaying before closing the eyes = midline cerebellar lesion
swaying after closing the eyes = damage to afferent systems
(Proprioceptive Systems)
 Appendicular Ataxia
-lesion to cerebellar hemispheres / lateral zone
-affects lateral motor systems - movement of extremities, skilled,
fine voluntary & planned movements
-Errors in force, direction, amplitude and speed
Patients can exhibit:
*Dysdiadochokinesia
-patients with cerebellar dysfunction are unable to pronate and
supinate
*Hpotonia loss of muscle tone
*Dysmetria improper measurement of distance or range for a
movement

A Individual with
appendicular ataxia (B)
has trouble tracing the
B image in A
 *Finger-Nose-Finger (Pass-Pointing) Test


-Used to help detemine
appendicular ataxia
-The patient is asked to
alternately touch their nose
and the examiner s finger as
quickly as possibletests for
irregular wavering
movements caused by limb
weakness & abnormal
movements caused by ataxia
-movement can be initiated,
but execution is poor

*Often times truncal and

appendicular ataxia both exist in
a patient!
 Friedrich s Ataxia
A hereditary disease with degeneration of
specific regions of spinal cord white matter
Spinocerebellar tracts
dorsal columns
corticospinal tracts
 Cognitive Functions

1-Motor Learning
-activity is not limited to modulationcerebellum also has a role in
motor/cerebellar learning for complex motor tasks
2-Classical Conditioning
-A noise, flash of light or some other stimulus preceeds a stimulus that
evokes a viseromotor response. The tone or flash will soon evoke that
visceral response.
-noise -> food - > salivation
-eventually the noise alone will result in salivation
-Cerebellar lesions disrupt this type of simple learning
3-Higher Cognitive Functions
-Evidence suggests that cerebellar lesions produce slight disruptions of
cognition called cognitive discoordination
-Patients have difficulty concentrating, attending to proper cues,
selecting correct words and both learning and performing complex
cognitive tasks

End!