Modulation of movement by the

cerebellum
Subject

Status Completed

Week 9

Property Chapter 19

Organisation of cerebellum

Cerebrocerebellum - the largest subdivision in human.

→ occupies most of the lateral part of cerebellar hemisphere - receive input
(indirect) ~ from areas of the cerebral cortex.

developed in primates & part. prominent in humans.

concerned w. the regulation of high skilled movements ~ planning & execution

of complex spatial + temporal sequences of movement.

Spinocerebellum - occupies the median + paramedian zones of cerebellar

hemispheres → the only part ~ receive input directly from the spinal cord.

more lateral (paramedian) part of spinocerebellum - concerned prim. w.

movements of distal muscles.

vermis - median strip of cerebella hemisphere lies along the midline.

→ concerned prim. w. movements of proximal muscles ~ regulates certain
types of eye movements.

Vestibulocerebellum - 3rd major subdivision → the phylogenetically oldest part of

the cerebellum.

comprise the caudal-inferior lobes of the cerebellum ~ incl. flocculus & nodulus.

Modulation of movement by the cerebellum 1

 → receives input from the vestibular nuclei in brainstem ~ concerned prim. w. the
vestibulo-ocular reflex + regulation of movements → maintain posture & equilibrium.

1. Cerebellar pecundles - the connections btwn the cerebellum & other parts of
nervous sys. are made by 3 pathways.

2. Superior cerebellar pecundle (brachium conjunctivum) - efferent pathway ~

neurons give rise to this pathway located in deep cerebellar nuclei.
→ their axons - project to the motor nuclei of the thalamus ~ relay signals to circuits
of upper neurons in the prim. motor + premotor divisions of cerebral cortex.

efferent axons - project directly to upper neurons in the deep layers of superior
colliculus → control orienting movements of the head and eyes.

3. The middle cerebellar pecundle (brachium pontis) - afferent pathway to

cerebellum → cell bodies that give rise to this pathway in base of contralateral pons
→ form the pontine nuclei.

4. The inferior cerebellar peduncle (resting form body) - smallest ~ but most
complex

contain multiple afferent + efferent pathways.

afferent pathways - axons from vestibular nuclei, spinal cord, several regions of
brainstem tegmentum.

efferent pathways - project to vestibular nuclei + reticular formation.

Projections to the cerebellum

Cortical axons - not project directly into the cerebellum → synapse on neurons in
ipsilateral pontine nuclei.

receive input from a wide variety of sources ~ all areas of the cerebral cortex +
superior colliculus.

Transverse pontine fibers (pontocerebellar fibers) - axons cross the midline →

enter the contralateral cerebellum via the middle cerebellar peduncle.

Modulation of movement by the cerebellum 2

 The size of the cerebral peduncles in ventral portion of human midbrain - prim. due
to the mag. of the projection from the cerebral cortex via the pontine nuclei → enter
cerebellum.

crossed projection of transverse pontine fibers into cerebellum via middle

cerebellum peduncle (mean) → signals originating in one cerebral hemisphere -
sent to neural circuits in opp. cerebellar hemisphere.

Sensory pathways - project to the cerebellum → vestibular axons in 8th cranial

nerve ~ axons from vestibular nuclei in the pons + medulla → project to
vestibulocerebellum.

Somatosensory relay neurons in dorsal nucleus of Clarke in spinal cord + external

cuneate nucleus of caudal medulla → send axons to spinocerebellum.

input - mapped in the spinocerebellum → provide the basis for orderly

representation of body w/in the cerebellum.

→ fine-grain electrophysio. analysis - indicate ea. small area of body ~

represented multiple times by spatially separated clusters of cells.

⇒ ensures the right cerebellum w. left half ~ many areas of the brain concerned w.
contralateral representations

→ cerebellum is concerned w. ipsilateral representations.

process info. related to the same side of body/ space → right cerebellum
is concerned w. right half of body and external space ~ left concerned w. left
half.

→ allows for fine-tuned control & coordination of movements & functions ~

associated w. ea. side of the body.

Proprioceptive signals from face - relayed via the mesencephalic trigeminal

nucleus to spinoacerebellum.

→ provide the cerebellum w. info. from the labyrinth in ear, muscle spindles, other
mechanoreceptors - monitor the position + motion of the body.

Visual + auditory signals - relayed via brainstem nuclei to cerebellum → provide

the cerebellum w. add. sensory signals ~ supplement the prorioceptive info.
regarding body position + motion.

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 Inferior olive

entire cerebellum - receives modulatory inputs from inferior olive → located in

the medulla oblongata.

modulatory inputs - invol. in the learning & memory functions ~ supported by

cerebellar circuitry.

receives input from various brain structures incl. the cerebral cortex,
reticular formation & spinal cord.

Inputs to inferior olive → suggest a role in integrating & processing info. from
diff. parts of nervous systems.

The olivo-cerebellar axons - exit the inferior olive medially ~ cross midline

→ enter the cerebellum on the opp. side thru. inferior cerebellar peduncle.

this pathway ensures info. from inferior olive → reaches the contralateral
(opp. side) cerebellum.

Gap junctions - neurons in inferior olive connected.

imp. for timing & spatial distribution of cerebellar responses to inputs

from inferior olive.

facilitate coordinated signalling - info. processing w/in inferior olive → infl.

cerebellar function.

Projections from the cerebellum

efferent neurons of cerebellar cortex - project to the deep cerebellar nuclei +

vestibular complex.

→ project to upper neurons in brainstem + thalamic nuclei - innervate upper

neurons in the motor cortex.

4 major deep nuclei: dentate nucleus, interposed nuclei, fastigial nucleus.

ea. receives input from diff. region of cerebellar cortex.

The vestibulocerebellum - projects directly to vestibular complex in brainstem → infl.

posture, equilibrium, vestibulo-ocular eye movements.

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 Pathways in dentate nucleus - project prim. to premotor & associational cortices of
the frontal lobe → function in planning & initiating volitional movements.

ea. cerebellar hemisphere - concerned w. the ipsilateral side of the body →

cross the midline ~ motor cortex in ea. hemisphere - govern contralateral
musculature
→ receive info. from the approp. cerebellar hemisphere.

dentate axons - exit the cerebellum via the superior cerebellar peduncle

~ cross the midline at the decussation of the superior cerebellar peduncle in

caudal midbrain.
→ ascend to the contralateral thalamus.

⇒ send axons to eye-movement-related upper neurons in superior colliculus ~ send

collaterals to parvocellular division of red nucleus in midbrain.

(provide a means for cerebellar output to feedback on major source of cerebellar

input.)

Figure 19.6B

Modulation of movement by the cerebellum 5

 Cerebrocerebellum form ‘closed loops’ ~ by sending info. back to non-motor areas
of the cortex.

→ regions of cerebellum - sends projections back to the same cortical areas from its
input signals originated.

provide a mechanism for the cerebellum - modulate its own input.

→ infl. the coordination of non-motor programs.

‘open loops’ - receive input from ultiple cortical areas & funnel output ~ back to
upper neurons in spec. regions of motor & premotor cortices.

Spinocerebellar pathways - direct toward circuits of upper neurons → govern the

execution of movement.

Somatotopic organisation - reflected in the organisation of its efferent projections

→ conform to the mediolateral organisation of motor control in spinal cord.

the fastigial nuclei (underlies vermis near midline)- project via. the inferior
cerebellar peduncle ~ nuclei of the reticular formation & vestibular complex
→ rise to medial tracts governing the axial & proximal limb musculature.

the interposed nuclei (underlies the paramedian subdivision of spinocerebellum)

- send projections via. the superior cerebellar peduncle ~ thalamic circuit

→ project to motor regions in frontal lobe concerned w. volitional movements of

the limbs.

Modulation of movement by the cerebellum 6

 💡 Most of the projection from the cerebellum ~ the eye-movement-related upper
motor neurons in the superior colliculus - arises in the dentate and interposed
nuclei

→ receive their input from the lateral portions of the cerebellar cortex.

Output pathway travels in the superior cerebellar peduncle and crosses the
midline → terminate on upper motor neurons in the deep layers of the
superior colliculus on the contralateral side

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 Thalamic nuclei - receive projections from the cerebrocerebellum +
spinocerebellum → segregated in 2 distinct subdivisions of ventral lateral nuclear
complex.

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 both relays project directly to prim. motor & premotor association cortices.

→ the cerebellum - access to upper neurons - organise the sequence of muscular

contractions underlying complex voluntary movements ~ circuits in frontal
association cortex → exert executive control over planning movements.

These nuclei of vestibular complex - govern the movements of eyes, head, neck → compensate
for linear & rotational accelerations of the head.

Circuits within the cerebellum

Purkinje cell - ultimate destination of afferent pathways ~ cerebeller cortex.

arise in widespread areas of the cerebral cortex → terminates in pontine nuclei

of basal pons → project to contralateral cerebellum.

mossy fibers - axons from pontine nuclei → send collateral branches - synapse
both on neurons in deep cerebellar nuclei & granule cells in layer of cerebellar
cortex.

Modulation of movement by the cerebellum 9

 Cerebellar granule cells - widely held to be most abundant class of neurons in
human brain
→ arise parallel filbers - ascend to the outermost molecular layer of cerebellar
cortex → bifurcate to form T-shapred branched ~ extend to folia → form
excitatory synapses w. the dencritic spines of the underlying Purkinje cells.

Purkinje cells - elaborate dendrites → extend into molecular layer from single
subjacent layer of giant Purkinje cell bodies layer.

ea. P cell - receive input from large number of parallel fibers → ea. fiber can
contact to vast number of P cells.

receive direct innput on their dendritic shafts from climbing fibers ~ arise in
tralateral inferior olive.
→ provide a training signal - modulate synaptic strength of the parallel fiber
connection w. P cells.

Purkinje cells - project to deep cerebellar nuclei ~ comprise only output cells of the
cerebral cortex.

are GABAergic - the output of the cerebellar cortex is inhibitory BUT the deep
cerebellar nuclei → receive excitatory input from mossy and climbing fibers.

Inputs of GABAergic interneurons - modulate the inhibitory act. of P cells.

basket cells (most powerful input) - inhibitory nests of synapse w. the

Purkinje cell bodies.

stellate cells - receive input from parallel fibers → provides inhibitory input
to P cell dendrites.

Golgi cells- the apical dendrites of inhibitory interneurons → have their cell
bodies in granular cell layer.
→receive input from parallel fibers - provide an inhibitory feedback to cells
of origin of parallel fibers.

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 💡 Circuit module of excitatory & inhibitory cells - repeated over every
subdivision of the cerebellum.

ea. subdivision - transformation of signal flow thru. these modules

→ provide the basis for both real-time regulation of movement & longterm
changes in regulation ~ underlies motor learning.

The deep cerebellar nuclei stage

Mossy + climbing fiber inputs - activate neurons in deep cerebellar nuclei →

forming excitatory loop ~ input signals convergence for cerebellar output.

output patterns of deep cerebellar nuclei - not direct replications of input

patterns
→ P cells driven by the same afferent pathways - modify these output
patterns → respond to excitatory inputs w. inhibitory output.

P cells - part of the cortical inhibitory loop w/in the cerebellar cortex incl.
interneurons.
→ control the flow of info. in the cerebellar cortex.

The modulation of cerebellar output ~ by cerebellar cortex → linked to motor

learning. & error correction.

Climbing fibers from inferior olive - convey the message of motor error to P
cells ~ incl. feedback signals from cerebellum via. red nucleus.

→ results in complex spikes in P cells ~ followed by longterm plasticity in

response to parallel fiber inputs.

Longterm depression - decr. in the efficacy of parallel fiber inputs to P cells ~ more
prominent in zebrin II-neg. P cells.

Zebrin II-neg. P cells - exhibit higher basal rates of simple spike activity.

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 Longterm potentiation - more likely in zebrin II-pos. bands ~ rate of simple spike
activity is relatively low.

→ regional diff. in gene expression & physio. properties across cerebellar cortex -
convey varying capabilities in info. processing.

Cerebellar circuitry & the coordination of ongoing movement

The execution of relatively simp. task ~ flexing & extending the wrist back - forth

→ elicits a dynamic pattern of act. in both P cells + deep cerebellar nuclear cells ~
follows the ongoing movement.

recognise the potential errors by comparing patterns of convergent act. ~

concurrently avail. to both cell types.

deep nuclear cells - send corrective signals ~ to upper neurons → maintain/

improve the accuracy of the movement.

The neuronal responses infl. by various aspects of movement incl. relaxation &
contraction of spec. muscles ~ position of joints, direction of next movement.

→ all encoded by changes in the discharge pattern of P cells - modulate the

ongoing output of the deep cerebellar nuclear cells.

Cerebellar injuries + disease - tend to disrupt the modulation + coordination of

ongoing movements.

spec. movement disrupted vary w. the location of the damage.

⇒ difficulty producing smooth, well-coordinated, multi-jointed movement.

Cerebellar ataxia - movement tend to be decomposed into jerky & imprecise
elements.

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 💡 Difficulty in movement - explained as disruption of the cerebellum’s role in
correcting errors in ongoing movements.
⇒ cerebellar error correction - ensures the movements are modified to cope
w. changing circumstances.

Monkey’s study w. cut tendons in one eye’s lateral rectus muscle → contributed to
understanding the cerebellum;s role in motor error reduction.

One eye is weakened - saccades made by that eye initially fall short of visual
targets (hypometric).

→ nervous system corrects these errors - saccade amp. incr. until they become
accurate.

patch cover switched to normal eye ~ saccades by normal eye become large.

Lesions in the vermis of spinocerebellum - impair the ability to recude motor errors
in saccades.

VOR - reflex that maintain stable gaze during head movements.

indv. fitted w. magnifying/ minifying spectacles → alter the size of visual image
on the retina ~ adjust their compensatory eye movements during head
movements.

→ adapt to the distances their eyes move to match the changed visual field.
(response of P cells + deep cerebellar nuclei neurons → support cerebellum -
crucial role in adaptive process)

Damage/ removal of cerebellum - impair VOR’s ability to adapt to new

conditions.

Further consequences of cerebellar lesions

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 These movement errors - always on the same side of the body → damage to to
the cerebellum ~ reflecting the cerebellum’s unusual status

brain structure - sesorimotor info. is represented ipsilaterally.

somatic, visual, other inputs represented topographically w/in the cerebellum

→ allow cerebellar damge to disrupt the coordination of movements - performed
by some muscle group.

→ movement deficits following circumscribed cerebellar damage maybe quite

spec.

Damage spec. affects movement in lower limbs - represented in anterior

spinocerebellum
~ consequences incl. a wide & staggering gait BUT little impairment of arm/ hand
movement.

Cerebellar lesions - lead to lack of coordination of ongoing movements. (BOX 19A)

damage to vestibulocerebellum - impairs the ability to stand upright → cannot

maintain the direction gaze/ fixation (nystagmus).

Disruption to vestibular nuclei - results in a reduction of muscle tone → indv.

w. damage to spinocerebellum ~ diff. controlling walking movement.
→ represents the inappropriate operation of groups of muscles - rely on
sensory feedback to produce smooth, concerted actions.
(dysdiadochokinesia, dysmetria)

Tremor (action/ intention tremors) - accompany over & undershooting of a

movement → due to disruption of the mechanism for detecting & correcting
movement errors.

Lesion of cerebrocerebellum - produce impairments in highly skilled sequences of

learned movements.
→ inability to perform smooth, precisely directed movements.

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