Neuroscience

Motor Systems

Cerebellum “little brain”

Significance
 
• Reception of information from the periphery concerning the position
of the body parts (proprioceptive information + muscle tone +
vestibular information…+ position of the head in space).
• Coordination… + prediction of movement…  skilled manipulation
of muscles…(feed-forward mechanisms…calculates trajectories…An error-
correcting movements/ “predictive control” of movements).
• Comparison + integration of information with plans for movement
received from the cortex… smooth coordinated motor actions…
• Plays a significant role in cognition (language and helps to coordinate &
predict mental concepts)…

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Cerebellum… • 10 % total volume of the brain.
• > ½ of total number of neurons in CNS.
• 40x more axons projecting to it than
exit from it.
• Its activity occurs subconsciously.

 Somatosensory
 Visual
 Auditory
 Vestibular
 Proprioceptive
 Neocortex/corticospinal tracts

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Cerebellum…

Integrator

Coordinator +
Predictor

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Functional anatomy
 
• Located in the posterior cranial fossa.
• Separated from the cerebrum by tentorium cerebelli.
• Overlies the 4th ventricle.
• Connected to the brainstem by three cerebellar peduncles.
• superior, middle, and inferior cerebellar peduncles
• All tracts to and from the cerebellum travel through the cerebellar
peduncles.

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Cerebellum:
Surfaces:
o Superior surface
o Inferior surface
o Anterior surface
Hemisphere:
o Right hemisphere
o Left hemisphere
• Midline structure: vermis
Folia ….. parallel, longitudinal folds/convolutions
o Organized into groups of folia
o 10 lobules
o 3 lobes

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Lobes
Lobes: 3

• Anterior (I-V)
• Posterior (VI-IX)
• Flocculonodular (X)

Two deep transverse fissures:

o Primary fissure: ant. & post. lobe.
o Posterolateral fissure: flocculonodular
lobe.

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Anatomical lobes of the cerebellum
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Cerebellar peduncles
• Connects cerebellum to the
dorsal aspect of the brain stem.
1. Superior cerebellar peduncle
/SCP  midbrain (Brachium
conjunctivum)

2. Middle cerebellar peduncle

/MCP  pons (Brachium
pontis)

3. Inferior cerebellar peduncle

ICP  medulla (Restiform
body)

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Afferent tracts to and efferent tracts from the cerebellum

Afferents
SCP Anterior spinocerebellar tract
Acoustic and optic information
MCP Pontocerebellar tract
ICP Vestibulocerebellar tract
Olivocerebellar tract
Posterior spinocerebellar tract
Efferents
Dentatorubrothalamic tract
Dentatothalamic tract
------
Cerebellovestibular tract
Cerebelloolivary tract

 Most cerebellar output fibers arise from the deep nuclei & project
through the superior cerebellar peduncle.
 Exception: Purkinje cells in the flocculonodular lobe
(directly project to vestibular nuclei in the brain stem).

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Cerebellum

Motor planning for

extremities/LCST

Proximal limb + trunk coordination

Distal limb coordination ACST + Reticulospinal +
LCST+ Rubrospinal tract Vestibulospinal + Tectospinal tracts

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 Medial  Lateral
• Fastigial
• Globose
• Emboliform
• Dentate

• Vestibular & visual inputs

• Projects to the vestibular nuclei in
the brain stem.
• Participates in balance + vestibular
reflexes + eye movements.
Functional components of the cerebellum
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Cerebellum
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Physiological anatomy of the cerebellum

1. Vestibulocerebellum/archicerebellum/flocculonodular lobe
• Oldest part of the cerebellum.
• Consists of the flocculonodular lobe.
• Connected to the vestibular apparatus through the vestibular nuclei in
the brain stem (semicircular canals & otolith organs).
• Receives mossy fiber visual input (from pretectal nuclei + 1° & 2°
visual cortex).
• Concerned with maintenance of equilibrium & controlling eye and
head movement.
NB
• Vestibulocerebellum  spinocerebellum cerebrocerebellum

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 Primary fissure

Spinocerebellum
Anterior lobe
Regulation of muscle tone,
coordination of skilled voluntary
movement

Posterior lobe
Planning & initiation of
voluntary activity

Flocculonodular
Cerebrocerebellum
Lobe (FN lobe)
Maintenance of balance, control
of eye movements Vestibulocerebellum

Folia
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2. Spinocerebellum/paleocerebellum
• Consists of the vermis + intermediate/medial paravermal parts of the
cerebellar hemispheres.
• Receives signals from somatosensory + proprioceptors (touch,
pressure & limb position)…spinal cord.
• Vermis receives visual, auditory, and vestibular input as well as
somatic sensory input from the head & proximal parts of the body.
• Concerned with the control of posture + muscle tone + gross
movement.
• Provides some immediate feedback based on sensory input from the
muscles…
• Projects through the ICP to fastigial & interposed nuclei.
• Is somatotopically organized
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3. Cerebrocerebellum/neocerebellum/pontocerebellum
 
• Consists of the lateral parts of the cerebellar hemispheres.
• Connected to the cerebral cortex/thru dentate nucleus (projects to
motor, premotor & pre-frontal cortices).
• Concerned with the planning & control of skilled voluntary
movements initiated by the cerebral cortex/contralateral cerebral cortex.
• Working memory…
NB: All these divisions are interconnected by interneurons.
• The structure of the cerebellum
o Outer layer  gray matter  cerebellar cortex
o Inner layer  white matter  cerebellar white matter

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Cerebellum…

Functional organization of the spinocerebellum, cerebrocerebellum & vestibulocerebellum.

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Cytoarchitecture of the cerebellar cortex
Three layers:
1. Molecular layer…. lies deep to the pia mater
• Outermost layer + processing layer.
• Interconnecting fibers/parallel fibers.

• Contains cell bodies & dendrites of stellate + basket cells

(+++ extensive dendrites of Purkinje cells).
• Inhibitory inter-neurons, dispersed among the excitatory axons of
granule cells and the dendrites of inhibitory Purkinje cells
• Contains the axons of the granule cells/parallel fibers.

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 A

E A
Cytoarchitecture and types of neurons
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 E A
A
Cerebellar cytoarchitecture
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2. Purkinje cell layer

• Middle layer + output layer.

• Giant Purkinje cells + processes of stellate & basket cells.
• Have large cell bodies and fanlike dendrites, extend upward into the
molecular layer…
(climbing fibers + excitatory & inhibitory interneurons).
• Purkinje cell axons conduct the entire output of the cerebellar
cortex (to the deep nuclei in the underlying white matter/or to the
vestibular nuclei in the brain stem).
• This output is entirely inhibitory and mediated by the
neurotransmitter γ-aminobutyric acid (GABA).

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Cytoarchitecture… Cerebellum

A
A
E
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3. Granule cell layer
• Deepest layer + input layer
• Granule cells (1010 cells!)
• Golgi cells (+++ + chandelier cells)
• Cells of Lugaro (5-HT), form inhibitory synapse with Golgi +
basket + stellate cells
• Unipolar brush cells, glutamatergic/glycine in FN lobe.
• Mossy fibers terminate in this layer
• Contains synaptic complexes cerebellar glomeruli.
• Space where the bulbous terminal of a mossy fiber makes
synaptic contact with golgi and granule cells.

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 Cerebellar glomerulus
• Mossy fibers
• Granule cell
• Golgi cell

Cerebellar layers and cerebellar glomerulus

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Cells of the cerebellar cortex
• 5 types of cells:
o Basket (-)
o Golgi (-)
o Granule cells (+)
o Purkinje (-)
o Stellate (-)
1. Basket cells(-)
• Inhibitory, GABAergic neurons, located in the molecular layer.
• Fiber branching is  to the Purkinje cell dendritic tree.
• Inputs: mossy + climbing fibers

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Cerebellum…

Cellular anatomy
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Cerebellar cortex and its component cell layers
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2. Golgi cell (-)
• Located in the granule cell layer.
• Processes radiate into all other layers.
• Inhibitory interneurons/GABAergic
• Synapse in the cerebellar glomerulus (+ parallel + mossy + climbing
fibers).
3. Granule cell (+)
• Very abundant & located in the granular layer/glutamatergic
interneurons
• Inputs: mossy fibers within the cerebellar glomerulus.
• Form parallel fibers (axons bifurcate at molecular layer… T-
shaped configuration).
• Course through the bushy dendrites of Purkinje cells
• Forms glutamate releasing excitatory synapses
• Stellate cells, basket cells, and golgi cells
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Cerebellar cortex and its component cell layers
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4. Purkinje cell (-) ….. are multipolar neurons
• Largest cell in the cerebellar cortex.
• Cell bodies are located in the Purkinje cell layer.
• Excitatory inputs: climbing fibers + parallel fibers of granule cells.
(200,000 to 106 granule cells)
• Inhibitory inputs: basket cells + stellate cells.
• Purkinje axon is the efferent pathway of the cerebellum.
• Projects via the deep cerebellar nuclei…. dentate nucleus
• Inhibitory/GABAergic synapses.
• Axons of some purkinje cells go right past the deep cerebellar nuclei
and synapse in the vestibular nuclei.
• Extracerebellar projection…. flocculonodular lobe/FNL

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5. Stellate cells

• Inhibitory interneurons/molecular layer.

• Inhibitory synapses with the dendrites of the Purkinje cells (taurine)
• Excitatory synapses with parallel fibers of the granule cells.

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Cerebellar neurons

- -
Afferent neurons

+ -

-
Efferent neuron

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Fibers of the cerebellar cortex

Three fiber types:

• Climbing fibers (+)
• Mossy fibers (+)
• Parallel fibers (+)
• Modulatory fibers…

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Wiring of cerebellar cortex

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1. Climbing fibers
• Originate from the inferior olivary nuclear complex.
• Sensory information from periphery & cerebral cortex.
• 1Climbing fiber: 1Purkinje cell (excitatory).
• Event detection, motor learning,.
• Glutamate/aspartate

NB
• Each neuron within the inferior olivary nuclear complex gives rise to
about 10 climbing fibers.
• Each climbing fiber contacts 1 to 10 Purkinje neurons, but each Purkinje
neuron receives synaptic input from only a single climbing fiber.

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2. Mossy fibers
• Cell bodies in spinal cord + brain stem.
• Afferents from the spinal cord + posterior column nuclei + trigeminal
system + pontine nuclei + vestibular nuclei…cerebral cortex.
• Form excitatory synapses on the dendrites of granule cells in the
granular layer.
• Encode magnitude & duration of peripheral stimuli or centrally
generated behaviors….
• Cerebellar glomerulus
o Golgi cell + granule cell dendrites + bulbous axon terminal of mossy
fibers.
o First processing station for cerebellar afferents.
o The synapse with granule cells is under inhibitory control of Golgi cell
axons.
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Cerebellar glomerulus

 Each mossy fiber forms 50 glomerulus/folium.

 Each granule cell receives input from 3-5 mossy fibers.
 A granule cell 3-5 short dendrites and each forms a glomerulus.
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3. Parallel fibers
• Synapse with the Purkinje cell dendrites (+++basket + stellate cells)
• Parallel fibers run perpendicular to the plane of the Purkinje cell
dendritic tree.

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4. Modulatory fibers

• Neural projections to all cortical layers/molecular layer

• Locus ceruleus  NE
• Raphe magnus  5-HT
• Brain stem  ACh
• Hypothalamus  Histamine

NB
• Two afferent fiber systems encode information differently.
• Parallel pathways compare excitatory and inhibitory signals.

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How does Purkinje neuron firing affect movement?

• 50APs/sec in the absence of sensory input

(Rapid, precise, tight temporal control)
• Purkinje neurons are inhibitory, thus when they slow or stop firing their targets are excited.

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Synaptic organization of cerebellar microcircuit
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Wiring of cerebellar cortex

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 Canonical circuit
• Climbing fibers
• Mossy fibers Excitatory

• Basket cells
• Golgi cells Inhibitory/GABA
• Purkinje cells
• Stellate cells

• Granule cells: excitatory/Glutamate

- • Deep nuclear cells: excitatory
Sculpting inhibition
• Allows cerebellum to smooth & coordinate
movement.

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Wiring of cerebellar cortex

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Cerebellar deep nuclei

• 4 paired deep cerebellar nuclei.

• Relay + processing stations for information coming from the
cerebellar cortex to targets outside the cerebellum.
• Nuclei (LM):
o Dentate….. largest deep nucleus
o Emboliform
Interposed n
o Globose
o Fastigial

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Organization of deep cerebellar nuclei Dentate nuclei
• Project contralaterally through the
SCP to contralateral thalamus & to
motor cortex.
• Influence planning & initiation of
voluntary movement.
Emboliform & Globose nuclei
• Project mainly to the contralateral
red nuclei & a small group is
projected to the motor cortex.
• Red Nuclei  Rubrospinal Tract
control of proximal limb muscles.
Fastigial nuclei
• Project to the vestibular nuclei
& to the pontine & medullary RF.
• Vestibulospinal & Reticulospinal
tracts (+ controls all musculature
involved in stance and gait).

Interposed nucleus = [Emboliform n. + globose n.]

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Deep cerebellar nuclei… outputs

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 Cerebellar cortex: Fractured somatotopy

Somatotopic representation/homunculus Functional areas of cerebellar cortex

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• Anterior lobe: extremities
• Posterior lobe: extremities + head
• Midline: trunk

Somatosensory projection areas in the cerebellar cortex

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Principal afferent inputs to cerebellum

Cerebral cortex

Pons Cerebellum

Vestibular Inferior Spinal

inputs Olive Cord

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 Corticobulbar
projections

Mossy fibers

Climbing fibers

Afferent inputs
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Afferent inputs from brainstem & cerebral cortex

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Tracts
• Cerebellum receives + interprets proprioceptive information.
• Coordinates balance/+ vestibular nuclei.
• Connections with the forebrain  coordination of limb movement + fine
movement + eye-hand coordination.
I. Afferents
• Vestibular system/balance.
• Spinal cord/proprioceptive information. Mossy fibers
• Pontine nuclei about cortical information.

• Inferior olivary nuclear complex Climbing fibers.

(Mainly contralateral/+bilateral projections…)

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Cerebellar Afferents

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Cerebellar Afferents

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Principal afferent tracts to the cerebellum
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Spinocerebellar tracts
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 II. Efferents
• Information leaving the cerebellum is relayed through the
deep cerebellar nuclei.
• Project to the cerebral cortex via the red nucleus and the
thalamus, vestibular nuclei and the olivary nuclear complex.
• All information leaving the cerebellum travels through the
superior and inferior cerebellar peduncles

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Principal efferent outputs of cerebellum

Cerebellar cortex

Deep nuclei

Vestibular Inferior Red

Thalamus
Nuclei Olive Nucleus

Cerebral
Cortex

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 • Direct  vestibular nuclei
• Indirect  CDN…brainstem, thalamus…
cerebral cortex…

Efferent outputs
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Purkinje cell projections from the cerebellar cortex to the deep cerebellar nuclei & the vestibular nuclei
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Cerebellar efferents

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Cerebellar efferents

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Principal efferent tracts from the cerebellum
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Efferent outputs of cerebellum

Outputs from fastigial nucleus

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Efferent outputs of cerebellum

Outputs from Purkinje cells thru interposed nucleus

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Efferent outputs of cerebellum

Outputs to cerebral cortex thru dentate nucleus

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Functional relationships
• Cerebellum  feedback + feed-forward mechanisms.
• Inputs from the inferior olivary nuclear complex /IONC go to all areas
of the cerebellum  modulate the output from the Purkinje cells…
• Inputs to & outputs from the cerebellum are segregated, and each
functional loop is independent of the others.
Feedback mechanism
• Plan for movement is compared with sensory + proprioceptive
signals resulting from movement…
• Requires an accurate comparison between the actual movement +
planned movement.
• Correct errors that have already occurred + cannot predict & prevent
errors.
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Feed-forward mechanism

• Allows quick reactions because errors are predicted and prevented

before they can happen…it involves anticipation of errors from previous
experience.
• Relies on learning from previous experience/associative learning.
• Calculates likely outcomes + predicts the sensory consequences 
correcting the ongoing movement before an error can occur.
•  not only for our movements but also for movements of objects and
people around us.
• Cerebellar coordination, timing & learning.

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Vestibulocerebellar connections…
• Stability & balance/state of body equilibrium.
• Components:
o Vestibular nuclei
o FNL
o Inferior parts of the paravermal area +
o Fastigial nucleus.
• Afferent inputs:
o Vestibular apparatus
o Vestibular nucleus
o Primary visual cortex
o Superior colliculus…. +

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• Efferent outputs:
o Vestibular nuclei.
• Function:
o Influences vestibulospinal tracts
that control postural muscles
(to maintain equilibrium).
o Influences vestibulo-ocular reflex…

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Spinocerebellar connections/
paleocerebellum
• Proprioception…
Components:
o Anterior lobe (vermis +
paravermis)
o Vermis
o Superior paravermal area
Afferent inputs:
o Ia + II, Ib, Clarke nucleus
o Acc. cuneate nucleus…
posterior spinocerebellar and
cuneocerebellar tracts through
the ICP to the spinocerebellum.
NB
• Spinal border cells + ‘efference copy’
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Function:
• Influences lateral muscles
(paramedian zone)
• Influences axial muscles (vermis).

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Spinocerebellum/Vermis
• Influences corticobulbar tracts that
control facial/tongue muscles
• Influences vestibulospinal, reticulospinal
+ ventral corticospinal tracts that control
axial muscles.

Efferents

• Via fastigial nucleus to:

o Vestibular nuclei
o Reticular formation
o Motor cortex

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Spinocerebellum/Vermis/Anterior lobe
• Influences rubrospinal, LCST that
control limb muscles/movement.

Efferents
• Via interposed/emboliform + globose
nuclei to:
o Red nucleus…thalamus/VL
o Motor cortex/LCST…
NB.
 Cerebellum controls the ipsilateral side of the
body thru the projection to the contralateral
motor cortex (coordination of limbs is
lateralized).

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Cerebrocerebellar connections/
neocerebellum
• Comprises the lateral aspects of
the posterior lobes.
• Input: pontine nuclei (contralateral
side, MCP, contralateral olivary
nuclear complex).

• These reciprocal connections with the

cerebral cortex put the cerebellum in a
position to coordinate & streamline motor
output from the cortex.

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Reciprocal connections:
• Thru the dentate nucleus  red nucleus 
dentatorubrothalamic tract  VL  motor
+ sensory area of cortex.
• Influences corticospinal & corticobulbar
tracts to regulate complex movements.
Functions
• Necessary for eye-hand coordination
(visual input + calculates the trajectory of
movement needed to reach or manipulate a
target, feedback & feed-forward mech.).
a. Sensory consequence: neocerebellum
predicts sensory consequence of a
movement thru comparison with past
experience.

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b. Voluntary movement
• Planning + automatization of voluntary
movements (fine-tuning of motor patterns).
c. Coordination of motor activity &
cognition
o Modulates but does not generate language
and cognition…
o Interface between cognition & motor
output…
o Language… linguistic coordination…
o Fluidity of language…
o Automatization of syntax & grammar…
Prediction of sentence structure &
flow…
Cerebrocerebellar circuitry
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Cerebellum and nonmotor function…
 The cerebellum is able to subserve cognitive & emotional functions because it is
anatomically interconnected with the associative and paralimbic cortices.
 The convergence of inputs from multiple regions of the cerebral cortex to adjacent
areas within the cerebellum facilitates cerebellar regulation of supramodal
functions.
 The cerebellar contribution to cognitive & emotional processes is one of modulation
rather than generation.
 The cerebellum performs computations for cognitive functions similar to those for
the sensorimotor system—but the nature of the information being modulated is
different.
 The disruption of the cerebellar influences on higher functions can lead to dysmetria
of thought or impairment of mental agility.
 The cerebellum may play an important role in the development of cognitive &
emotional function.
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Summary

Feedback pathways: between the cerebrum, cerebellum, spinal cord and the brainstem nuclei
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Summary

Cerebral and cerebellar control of voluntary movements

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Clinical correlates
 
The cerebellar syndrome
 
Lesions involve large errors in movements…

 Same side of the lesion (3As)

• Asthenia
Potentiating motor signal is lost…
• Atonia (hypotonia, a diminished resistance to passive limb displacements).
o Loss of facilitatory effect of dentate and interpositus nuclei
o Pendular reflexes
NB
o Astasia-abasia, an inability to stand or walk.
o Astasia is loss of the ability to maintain a steady limb or body posture across
multiple joints.
o Abasia is loss of the ability to maintain upright stance against gravity.

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• Ataxia: the abnormal execution of multi-jointed voluntary movements,
characterized by lack of coordination

 Dysmetria (Failure of the timing and damping functions)

Hypermetria
Hypometria
 Kinetic tremor (intention tremor)
 Adiadokokinesia/dysdiadochokinesia…
 Decomposition of movement…
 Dysarthria…
 Rebounding…
 Nystagmus…
 Staggering gait…

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