Cerebellum: Key Cerebellar Functions

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Cerebellum

Cerebellar
Signs

Key Cerebellar Functions


• Comparison of intent and action (ie.,
errors) and generates corrective signals
• Motor learning and adaptation
• Motor cognition and general cognition
(new evidence; controversial)

• Plays a role in automating and


optimizing behavior

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Lecture Plan:
• Structural and functional overview
• Principal pathways into and out of the
cerebellum
• Experimental approaches to reveal:
– Anticipatory control
– Motor learning
– Mental processes underlying movement
control

Motor Hierarchy

1° Som sensory

Intent
Actual

Exerts influence
at all levels

Cerebellar Functional Anatomy

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Folia

Cerebellar Cortex

Deep Cerebellar Nuclei:


Dentate

Interposed
Fastigial

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Input-output Organization
Deep Cerebellar
Nuclei:
Cerebellar Fastigial
cortex Interposed

+
Dendate

Cortex

+ - Nuclei

+ Output

Vestibular
nuclei

Extrinsic inputs:
mossy fiber
climbing fiber

Cerebellar
Divisions Spinocerebellum:
Vermis
Spinocerebellum Intermediate hem.
(Vermis + Intermed.
Intermed. Hem) Cerebrocerebellum:
Control of limbs Lateral hem.
and trunk
Cerebrocerebellum
(Lateral hemisphere)
Planning of movement+
IVth vent Vermis
Vestibulo-cerebellum Intermediate hem.
(Floculo-nodular lobe) Lateral hem.
Control of eye &
head movements
Balance

Floculo-nodular lobe

Molecular layer

Purkinje layer
Granular layer

White matter

Nissl-stained section through cerebellar cortex

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Cerebellar Cortex
Inputs
Climbing fibers
Mossy fibers
Output
Purkinje neurons
Interneurons
Granule neurons
Stellate neurons
Basket neurons
Golgi neurons

Apical
dendrites

Purkinje Cell
body
neuron

Cerebellar Cortex
Inputs
Climbing fibers
Mossy fibers
Output
Purkinje neurons
Interneurons
Granule neurons +
Stellate neurons -
Basket neurons -
Golgi neurons -

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Cerebellar Cortex: activation by
mossy fibers
Inputs
Climbing fibers
Mossy fibers
Output
Purkinje neurons
Interneurons
Granule neurons
Stellate neurons
Basket neurons
Golgi neurons

+
-
to Deep Cerebellar
Nuclei

Cerebellar Cortex: activation by


climbing fibers
Inputs
Climbing fibers
Mossy fibers
Output
Purkinje neurons
Interneurons
Granule neurons
Stellate neurons
Basket neurons
Golgi neurons

+
-
to Deep Cerebellar Nuclei

Inferior olivary nucleus: source of all


climbing fibers
to cerebellum,
via inferior
peduncle

Mossy fibers from all other sources

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Cerebellar Cortex: Inhibitory
interneurons
Inputs
Climbing fibers
Mossy fibers
Output
Purkinje neurons
Interneurons
Granule neurons
Stellate neurons
Basket neurons
Golgi neurons

Functional
divisions

Spinocerebellum

Vermis
Intermediate
hemisphere Spinocerebellum
Cerebrocerebellum Vermis
Lateral hemisphere Intermediate hem
Cerebrocerebellum
Lateral hemisphere
Inter
Dentate posed

Fastigial
Vestibulo-
cerebellum Vestibulo-
cerebellum
Floculo-nodular
lobe
To frontal To To To
motor areas lateral medial vestibular
sysetms sysetms nuclei

Motor Motor Eye mvt &


planning execution balance

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Medial & lateral systems

Interposed
nuclei

Fastigial
Vestibulocerebellum
via vestibular nuclei

Intermediate and Lateral


Hemispheres
Planning and
limb control Thalamus and
Cortical motor areas

Brain stem nuclei

Cerebellar cortex
& Deep nuclei

Ipsilateral

Intermediate and Lateral Vermis & Vestibulocerebellum


Hemispheres

Thalamus and
Cortical motor areas

Cerebellar cortex
& Deep nuclei
Brain stem nuclei
Axial control

Ipsilateral Bilateral

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Functions of the Cerebellum

• Feed- forward or predictive motor control


(nuts & bolts of skillful movements)
• Motor learning/adaptation
• Non- motor functions:
– Active tactile exploration
– Higher brain functions (cerebellar cognitive-
affective syndrome)

Anticipatory control
• Anticipating the motor consequences of an event
– See stop light and brake
– Predict baseball location during batting
– Anticipating duck location in an video game
• Cerebellum’s role:
– Fairly low level routines
– Correlations & associations
• Implemented via lateral and medial pathways

Feed-Forward or
Predictive Control
Normal:
• Perturbation extends arm,
stretching biceps
• Muscle action flexes arm and
tends to restore arm position
• Anticipatory contraction of
triceps (extensor) prevents
flexion overshoot
– Occurs during triceps
shortening; not stretch reflex
– BRAKE
• Result: arm position
stabilized at start position

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Feed-Forward or
Predictive Control
Block cerebellar function (cool):
• Perturbation extends arm,
stretching biceps
• Biceps contraction prolonged
• Muscle action flexes arm but
well beyond initial arm position--
OVERSHOOT
• Delayed triceps contraction
– Reverts to simple stretch reflex
– LOSS of brake (antic. control)
• Result: cycle of flexion-
extension (similar to
cerebellar tremor)

Feed-Forward or
Predictive Control

Delayed triceps (antagonist)


produces oscillations

Motor
Learning

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Non-motor Function
Passive stimulation Discriminate roughness

Manipulate only Manipulate + discriminate

Cerebellar Motor Functions

• Implemented via lateral and medial pathways,


especially the corticospinal tract
• Incorporated into motor programs via frontal motor
areas (SMA, premotor cortex…)
• Becomes part of motor strategy via prefrontal cortex

1° motor Executing
cortex movements
Premotor Planning
cortex movements
Dorsolateral Movement
prefrontal strategies
cortex Working memory

MD

All work together


during a motor task: VA/VL Lateral
-anticipatory control during
cerebellum
motor learning
-using sensory information
to guide movement Intermediate
cerebellum

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Cerebellar Cognitive Affective
Disorder
• Lesions of the posterior lobe and vermis
• Impairment of executive functions
– Planning, verbal fluency, abstract reasoning
• Difficulties with spatial cognition
– Visuo-spatial organization, visual memory
• Personality changes
– Blunting of affect, inappropriate behaviors
• Language disorders
– Agrammatism

Anterior
cingulate cortex Emotions/affect

Dorsolateral
prefrontal Cognition
cortex

MD

Lateral
cerebellum

Conclusions

• Cerebellar lesions produce


– Incoordination & errors not weakness
– Lose ability to anticipate errors
– Lose ability to correct
• Motor learning
• Not just motor

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