What is the function of the cerebellum in overall motor control?

Cerebellum - influences the rate, range, force, and direction of movements. It also influences
muscle tone and posture, as well as eye movement and balance.

In superior or inferior views, the shape of the cerebellum resembles a butterfly.

The central constricted area is the vermis (= worm), and the lateral “wings” or lobes are the cerebellar hemispheres
(Figure 14.8a, b). Each hemisphere consists of lobes separated by deep and distinct fissures. The anterior lobe and
posterior lobe govern subconscious aspects of skeletal muscle movements. The flocculonodular lobe (flok-ū-lō-NOD-ū-
lar; flocculo- = wool-like tuft) on the inferior surface contributes to equilibrium and balance.

The nervous system uses the cerebellum to coordinate motor control functions at three levels:

1. The Vestibulocerebellum
 This level consists principally of the small flocculonodular cerebellar lobes that lie under the
posterior cerebellum and adjacent portions of the vermis.
 It provides neural circuits for most of the body’s equilibrium movements.

2. The Spinocerebellum
 This level consists of most of the vermis of the posterior and anterior cerebellum plus the adjacent
intermediate zones on both sides of the vermis.
 It provides the circuitry for coordinating mainly movements of the distal portions of the limbs,
especially the hands and fingers.
3. The Cerebrocerebellum
 This level consists of the large lateral zones of the cerebellar hemispheres, lateral to the
intermediate zones.
 It receives virtually all its input from the cerebral motor cortex and adjacent premotor and
somatosensory cortices of the cerebrum.
 It transmits its output information in the upward direction back to the brain, functioning in a
feedback manner with the cerebral cortical sensorimotor system to plan sequential voluntary body
and limb movements.
o These movements are planned as much as tenths of a second in advance of the actual
movements.
o This process is called development of “motor imagery” of movements to be performed.

The Vestibulocerebellum Functions in Association with the Brain Stem and Spinal Cord to Control
Equilibrium and Postural Movements

 The vestibulocerebellum originated phylogenetically at about the same time that the vestibular apparatus in
the inner ear developed.
 loss of the flocculonodular lobes and adjacent portions of the vermis of the cerebellum, which constitute the
vestibulocerebellum, causes extreme disturbance of equilibrium and postural movements
 In people with vestibulocerebellar dysfunction, equilibrium is far more disturbed during performance of
rapid motions than during inactivity, especially when these movements involve changes in direction of
movement and stimulate the semicircular ducts.
o This phenomenon suggests that the vestibulocerebellum is important in controlling balance
between agonist and antagonist muscle contractions of the spine, hips, and shoulders during rapid
changes in body positions as required by the vestibular apparatus.
 One of the major problems in controlling balance is the amount of time required to transmit position signals
and velocity of movement signals from the different parts of the body to the brain.
o Even when the most rapidly conducting sensory pathways are used, up to 120 m/sec in the
spinocerebellar afferent tracts, the delay for transmission from the feet to the brain is still 15 to 20
milliseconds.
  The feet of a person running rapidly can move as much as 10 inches during that time.
o Therefore, it is never possible for return signals from the peripheral parts of the body to reach the
brain at the same time that the movements actually occur.
 How, then, is it possible for the brain to know when to stop a movement and to perform the next sequential
act when the movements are performed rapidly?
o The answer is that the signals from the periphery tell the brain how rapidly and in which directions
the body parts are moving.
o It is then the function of the vestibulocerebellum to calculate in advance from these rates and
directions where the different parts will be during the next few milliseconds.
o The results of these calculations are the key to the brain’s progression to the next sequential
movement.
 Thus, during control of equilibrium, it is presumed that information from both the body periphery and the
vestibular apparatus is used in a typical feedback control circuit to provide anticipatory correction of
postural motor signals necessary for maintaining equilibrium even during extremely rapid motion,
including rapidly changing directions of motion.

Spinocerebellum—Feedback Control of Distal Limb Movements via the Intermediate Cerebellar

Cortex and the Interposed Nucleus

 As shown in Figure 57-8, the intermediate zone of each cerebellar hemisphere receives two types of
information when a movement is performed:
(1) information from the cerebral motor cortex and from the midbrain red nucleus, telling the cerebellum
the intended sequential plan of movement for the next few fractions of a second; and
(2) feedback information from the peripheral parts of the body, especially from the distal proprioceptors of
the limbs, telling the cerebellum what actual movements result.
 After the intermediate zone of the cerebellum has compared the intended movements with the actual
movements, the deep nuclear cells of the interposed nucleus send corrective output signals
(1) back to the cerebral motor cortex through relay nuclei in the thalamus and
(2) to the magnocellular portion (the lower portion) of the red nucleus that gives rise to the rubrospinal
tract.
o The rubrospinal tract in turn joins the corticospinal tract in innervating the lateralmost motor
neurons in the anterior horns of the spinal cord gray matter, the neurons that control the distal
parts of the limbs, particularly the hands and fingers.
o This part of the cerebellar motor control system provides smooth, coordinated movements of the
agonist and antagonist muscles of the distal limbs for performing acute purposeful patterned
movements.
 The cerebellum seems to compare the “intentions” of the higher levels of the motor control system, as
transmitted to the intermediate cerebellar zone through the corticopontocerebellar tract, with the
“performance” by the respective parts of the body, as transmitted back to the cerebellum from the
periphery.
 In fact, the ventral spinocerebellar tract even transmits back to the cerebellum an “efference” copy of the
actual motor control signals that reach the anterior motor neurons, and this information is also integrated
with the signals arriving from the muscle spindles and other proprioceptor sensory organs, transmitted
principally in the dorsal spinocerebellar tract.
 Similar comparator signals also go to the inferior olivary complex; if the signals do not compare favorably,
the olivary–Purkinje cell system, along with possibly other cerebellar learning mechanisms, eventually
corrects the motions until they perform the desired function.

Function of the Cerebellum to Prevent Overshoot and to “Damp” Movements

 Almost all movements of the body are “pendular.”
o For example, when an arm is moved, momentum develops, and the momentum must be overcome
before the movement can be stopped.
o Because of momentum, all pendular movements have a tendency to overshoot.
 o If overshooting occurs in a person whose cerebellum has been destroyed, the conscious centers of
the cerebrum eventually recognize this error and initiate a movement in the reverse direction to
attempt to bring the arm to its intended position.
o However, the arm, by virtue of its momentum, overshoots once more in the opposite direction, and
appropriate corrective signals must again be instituted.
o Thus, the arm oscillates back and forth past its intended point for several cycles before it finally
fixes on its mark.
 This effect is called an action tremor or intention tremor.
 If the cerebellum is intact, appropriate learned, subconscious signals stop the movement precisely at the
intended point, thereby preventing the overshoot and the tremor.
o This activity is the basic characteristic of a damping system.
o All control systems regulating pendular elements that have inertia must have damping circuits
built into the mechanisms.
 For motor control by the nervous system, the cerebellum provides most of this damping function.

Cerebellar Control of Ballistic Movements

 Most rapid movements of the body, such as the movements of the fingers in typing, occur so rapidly that it
is not possible to receive feedback information either from the periphery to the cerebellum or from the
cerebellum back to the motor cortex before the movements are over.
o These movements are called ballistic movements,
 meaning that the entire movement is preplanned and set into motion to go a specific
distance and then to stop.
 Another important example is the saccadic movements of the eyes,
o in which the eyes jump from one position to the next when reading or when looking at successive
points along a road as a person is moving in a car.
 Three major changes occur in these ballistic movements when the cerebellum is removed:
(1) the movements are slow to develop and do not have the extra onset surge that the cerebellum usually
provides;
(2) the force developed is weak; and
(3) the movements are slow to turn off, usually allowing the movement to go well beyond the intended
mark.
o Therefore, in the absence of the cerebellar circuit, the motor cortex has to think extra hard to turn
ballistic movements on and off.
o Thus, the automatism of ballistic movements is lost.
 Considering once again the circuitry of the cerebellum, one sees that it is beautifully organized to perform
this biphasic, first excitatory and then delayed inhibitory function that is required for preplanned rapid
ballistic movements.
 Also, the built-in timing circuits of the cerebellar cortex are fundamental to this particular ability of the
cerebellum.

Cerebrocerebellum—Function of the Large Lateral Zone of the Cerebellar Hemisphere to Plan,

Sequence, and Time Complex Movements

 In humans the lateral zones of the two cerebellar hemispheres are highly developed and greatly enlarged.
o This characteristic goes along with human abilities to plan and perform intricate sequential
patterns of movement, especially with the hands and fingers, and to speak.
o Yet, the large lateral zones of the cerebellar hemispheres have no direct input of information from
the peripheral parts of the body.
o In addition, almost all communication between these lateral cerebellar areas and the cerebral
cortex is not with the primary cerebral motor cortex but instead with the premotor area and
primary and association somatosensory areas.
  Even so, destruction of the lateral zones of the cerebellar hemispheres, along with their deep nuclei, the
dentate nuclei, can lead to extreme incoordination of complex purposeful movements of the hands, fingers,
and feet and of the speech apparatus.
o This condition has been difficult to understand because there is no direct communication between
this part of the cerebellum and the primary motor cortex.
 However, experimental studies suggest that these portions of the cerebellum are concerned with two other
important but indirect aspects of motor control:
(1) planning of sequential movements and
(2) “timing” of the sequential movements.

Planning of Sequential Movements.

 The planning of sequential movements requires that the lateral zones of the hemispheres communicate with
both the premotor and sensory portions of the cerebral cortex, and it requires two-way communication
between these cerebral cortex areas with corresponding areas of the basal ganglia.
 It seems that the “plan” of sequential movements actually begins in the sensory and premotor areas of the
cerebral cortex, and from there the plan is transmitted to the lateral zones of the cerebellar hemispheres.
 Then, amid much two-way traffic between the cerebellum and the cerebral cortex, appropriate motor
signals provide transition from one sequence of movements to the next.
 An interesting observation that supports this view is that many neurons in the cerebellar dentate nuclei
display the activity pattern for the sequential movement that is yet to come while the present movement is
still occurring. Thus, the lateral cerebellar zones appear to be involved not with what movement is
happening at a given moment but with what will be happening during the next sequential movementa
fraction of a second or perhaps even seconds later.
 To summarize, one of the most important features of normal motor function is one’s ability to progress
smoothly from one movement to the next in orderly succession.
 In the absence of the large lateral zones of the cerebellar hemispheres, this capability is seriously disturbed
for rapid movements.

Timing Function for Sequential Movements.

 Another important function of the lateral zones of the cerebellar hemispheres is to provide appropriate
timing for each succeeding movement.
 In the absence of these cerebellar zones, one loses the subconscious ability to predict how far the different
parts of the body will move in a given time.
 Without this timing capability, the person becomes unable to determine when the next sequential
movement needs to begin. As a result, the succeeding movement may begin too early or, more likely, too
late.
 Therefore, lesions in the lateral zones of the cerebellum cause complex movements (e.g., those required for
writing, running, or even talking) to become incoordinate and lacking ability to progress in orderly
sequence from one movement to the next.
o Such cerebellar lesions are said to cause failure of smooth progression of movements.

Extramotor Predictive Functions of the Cerebrocerebellum.

 The cerebrocerebellum (the large lateral lobes) also helps to “time” events other than movements of the
body.
o For example, the rates of progression of both auditory and visual phenomena can be predicted by
the brain, but both of these require cerebellar participation.
o As an example, a person can predict from the changing visual scene how rapidly he or she is
approaching an object.
 A striking experiment that demonstrates the importance of the cerebellum in this ability is the effects of
removing the large lateral portions of the cerebellum in monkeys.
o Such a monkey occasionally charges into the wall of a corridor because it is unable to predict
when it will reach the wall.
 It is quite possible that the cerebellum provides a “time base,” perhaps using time-delay circuits, against
which signals from other parts of the central nervous system can be compared.
 It is often stated that the cerebellum is particularly helpful in interpreting rapidly changing spatiotemporal
relations in sensory information.