Overview: Functions of the Cerebellum

Figure 5.1
Cerebellum

The cerebellum (little brain) is a structure that is located at the back of the brain,
underlying the occipital and temporal lobes of the cerebral cortex (Figure 5.1). Although
the cerebellum accounts for approximately 10% of the brains volume, it contains over
50% of the total number of neurons in the brain. Historically, the cerebellum has been
considered a motor structure, because cerebellar damage leads to impairments in
motor control and posture and because the majority of the cerebellums outputs are to
parts of the motor system. Motor commands are not initiated in the cerebellum; rather,
the cerebellum modifies the motor commands of the descending pathways to make
movements more adaptive and accurate. The cerebellum is involved in the following
functions:
Maintenance of balance and posture. The cerebellum is important for making
postural adjustments in order to maintain balance. Through its input from vestibular
receptors and proprioceptors, it modulates commands to motor neurons to compensate
for shifts in body position or changes in load upon muscles. Patients with cerebellar
damage suffer from balance disorders, and they often develop stereotyped postural
strategies to compensate for this problem (e.g., a wide-based stance).
Coordination of voluntary movements. Most movements are composed of a number
of different muscle groups acting together in a temporally coordinated fashion. One
major function of the cerebellum is to coordinate the timing and force of these different
muscle groups to produce fluid limb or body movements.
Motor learning. The cerebellum is important for motor learning. The cerebellum plays
a major role in adapting and fine-tuning motor programs to make accurate movements
through a trial-and-error process (e.g., learning to hit a baseball).

Cognitive functions. Although the cerebellum is most understood in terms of its

contributions to motor control, it is also involved in certain cognitive functions, such as
language. Thus, like the basal ganglia, the cerebellum is historically considered as part
of the motor system, but its functions extend beyond motor control in ways that are not
yet well understood.

5.2 Cerebellar Gross Anatomy

The cerebellum consists of two major parts (Figure 5.2A). The cerebellar deep nuclei (or
cerebellar nuclei) are the sole output structures of the cerebellum. These nuclei are
encased by a highly convoluted sheet of tissue called the cerebellar cortex, which
contains almost all of the neurons in the cerebellum. A cross-section through the
cerebellum reveals the intricate pattern of folds and fissures that characterize the
cerebellar cortex (Figure 5.3). Like the cerebral cortex, cerebellar gyri are reproducible
across individuals and have been identified and named. We will only be concerned with
some of the larger divisions of the cerebellar cortex.

Figure 5.2
(A) Cerebellar
deep nuclei and
cerebellar cortex
in an idealized
brain section. (B)
External
morphology of the
cerebellum.

Figure 5.3
Midsagittal crosssection of
cerebellum
showing the three
primary lobes of
the cerebellum.

Divisions of the cerebellum. Two major fissures running mediolaterally divide the
cerebellar cortex into three primary subdivisions (Figure 5.2B and Figure 5.3).
The posterolateral fissure separates the flocculonodular lobe from the corpus
cerebelli, and the primary fissure separates the corpus cerebelli into a posterior
lobe and an anterior lobe (Figure 5.4). The cerebellum is also divided sagittally into
three zones that run from medial to lateral (Fig. 5.4). The vermis (from the Latin word
for worm) is located along the midsagittal plane of the cerebellum. Directly lateral to
the vermis is the intermediate zone. Finally, the lateral hemispheres are located
lateral to the intermediate zone (there are no clear morphological borders between the
intermediate zone and the lateral hemisphere that are visible from a gross specimen).

Figure 5.4
Divisions of
cerebellum.
Click PLAY to
see
schematic
unfolding
of
cerebellum.

Cerebellar nuclei. All outputs from the cerebellum originate from the cerebellar
deep nuclei. Thus, a lesion to the cerebellar nuclei has the same effect as a complete
lesion of the entire cerebellum. It is important to know the inputs, outputs, and
anatomical relationships between the different cerebellar nuclei and the subdivisions of
the cerebellum (Figure 5.5).

Figure 5.5
Input and
output
pathways of
the
cerebellum.
Click on the
names of
each
cerebellum
functional
subdivision
(cerebrocere
bellum,
spinocerebel
lum, and

vestibulocer
ebellum) to
view each
pathway in
isolation.
The
cerebellar
deep nuclei
are the sole
outputs of
the
cerebellum.

1. The fastigial nucleus is the most medially located of the cerebellar nuclei. It
receives input from the vermis and from cerebellar afferents that carry
vestibular, proximal somatosensory, auditory, and visual information. It projects
to the vestibular nuclei and the reticular formation.
2. The interposed nuclei comprise the emboliform nucleus and the globose
nucleus. They are situated lateral to the fastigial nucleus. They receive input
from the intermediate zone and from cerebellar afferents that carry spinal,
proximal somatosensory, auditory, and visual information. They project to the
contralateral red nucleus (the origin of the rubrospinal tract).
3. The dentate nucleus is the largest of the cerebellar nuclei, located lateral to the
interposed nuclei. It receives input from the lateral hemisphere and from
cerebellar afferents that carry information from the cerebral cortex (via the
pontine nuclei). It projects to the contralateralred nucleus and
the ventrolateral (VL) thalamic nucleus.
4. The vestibular nuclei are located outside the cerebellum, in the medulla.
Hence, they are not strictly cerebellar nuclei, but they are considered to be
functionally equivalent to the cerebellar nuclei because their connectivity
patterns are identical to the cerebellar nuclei. The vestibular nuclei receive input
from the flocculonodular lobe and from the vestibular labyrinth. They project to
various motor nuclei and originate the vestibulospinal tracts.
In addition to these inputs, all cerebellar nuclei and all regions of cerebellum get special
inputs from the inferior olive of the medulla (discussed below).
It is convenient to remember that the anatomical locations of the cerebellar nuclei
correspond to the cerebellar cortex regions from which they receive input. Thus, the
medially located fastigial nucleus receives input from the medially located vermis; the
slightly lateral interposed nuclei receive input from the slightly lateral intermediate
zone; and the most lateral dentate nucleus receives input from the lateral hemispheres.
Cerebellar peduncles. Three fiber bundles carry the input and output of the
cerebellum.

1. The inferior cerebellar peduncle (also called the restiform body) primarily
contains afferent fibers from the medulla, as well as efferents to the vestibular
nuclei.
2. The middle cerebellar peduncle (also called the brachium pontis) primarily
contains afferents from the pontine nuclei.
3. The superior cerebellar peduncle (also called the brachium conjunctivum)
primarily contains efferent fibers from the cerebellar nuclei, as well as some
afferents from the spinocerebellar tract.
Thus, the inputs to the cerebellum are conveyed primarily through the inferior and
middle cerebellar peduncles, whereas the outputs are conveyed primarily through the
superior cerebellar peduncle. The inputs arise from the ipsilateral side of the body, and
the outputs also go to the ipsilateral side of the body. Note that this is true even for the
outputs to the contralateral red nucleus. Recall from the chapter on descending motor
pathways that the rubrospinal tract immediately crosses the midline after the fibers
leave the red nucleus. Thus, cerebellar output to the red nucleus affects the ipsilateral
side of the body by a double-crossed pathway. Unlike the cerebral cortex, the
cerebellum receives input from, and controls output to, the ipsilateral side of the body,
and damage to the cerebellum therefore results in deficits to the ipsilateral side of the
body.

5.3 Functional Subdivisions of the Cerebellum

The anatomical subdivisions described above correspond to three major functional
subdivisions of the cerebellum.
Vestibulocerebellum. The vestibulocerebellum comprises the flocculonodular
lobe and its connections with the lateral vestibular nuclei. Phylogenetically, the
vestibulocerebellum is the oldest part of the cerebellum. As its name implies, it is
involved in vestibular reflexes (such as the vestibuloocular reflex; see below) and in
postural maintenance.
Spinocerebellum. The spinocerebellum comprises the vermis and the intermediate
zones of the cerebellar cortex, as well as thefastigial and interposed nuclei. As its
name implies, it receives major inputs from the spinocerebellar tract. Its output
projects to rubrospinal, vestibulospinal, and reticulospinal tracts. It is involved in the
integration of sensory input with motor commands to produce adaptive motor
coordination.
Cerebrocerebellum. The cerebrocerebellum is the largest functional subdivision of the
human cerebellum, comprising the lateralhemispheres and the dentate nuclei. Its
name derives from its extensive connections with the cerebral cortex, via the pontine
nuclei (afferents) and the VL thalamus (efferents). It is involved in the planning and
timing of movements. In addition, the cerebrocerebellum is involved in the cognitive
functions of the cerebellum.

5.4 Histology and Connectivity of Cerebellar Cortex

The cerebellar cortex is divided into three layers (Figure 5.6). The innermost layer, the
granule cell layer, is made of 5 x 1010 small, tightly packed granule cells. The middle
layer, the Purkinje cell layer, is only 1-cell thick. The outer layer, the molecular layer,
is made of the axons of granule cells and the dendrites of Purkinje cells, as well as a
few other cell types. The Purkinje cell layer forms the border between the granule and
molecular layers.

Figure 5.6
Cerebellar
circuitry.
This basic
pattern is
repeated
throughout
all regions
of the
cerebellum.

Granule cells. Granule cells are very small, densely packed neurons that account for
the huge majority of neurons in the cerebellum. Indeed, cerebellar granule cells
account for more than half of the neurons in the entire brain. These cells receive input
from mossy fibers and project to the Purkinje cells.

Figure 5.7
Front view of
Purkinje cell.
Click PLAY to see
side view of the
Purkinje cell.
This view shows
that the cell is
virtually flat in this
dimension. Note
the parallel fibers
of the granule cells
that run
perpendicularly to
the Purkinje cell.

Purkinje cells. The Purkinje cell is one of the most striking cell types in the
mammalian brain. Its apical dendrites form a large fan of finely branched processes
(Figure 5.7). Remarkably, this dendritic tree is almost two-dimensional; looked at from
the side, the dendritic tree is flat (click PLAY on Figure 5.7). Moreover, all Purkinje cells
are oriented in parallel. This arrangement has important functional considerations, as
we shall see below.
Other cell types. In addition to the major cell types (granule cells and Purkinje cells),
the cerebellar cortex also contains various interneuron types, including the Golgi cell,
the basket cell, and the stellate cell.
Connectivity. The cerebellar cortex has a relatively simple, stereotyped connectivity
pattern that is identical throughout the whole structure. Figure 6 illustrates a simplified
diagram of the connectivity of the cerebellum. Cerebellar input can be divided into two
distinct classes.
1. Mossy fibers originate in the pontine nuclei, the spinal cord, the brainstem
reticular formation, and the vestibular nuclei, and they make excitatory
projections onto the cerebellar nuclei and onto granule cells in the cerebellar
cortex. They are called mossy fibers because of the tufted appearance of their
synaptic contacts with granule cells. There is a large degree of divergence in the
mossy fiber-granule cell connection, as each mossy fiber innervates hundreds of
granule cells. The granule cells send axons up toward the cortical surface. Each
axon bifurcates in the molecular layer, sending a collateral in opposite directions.
These fibers, called parallel fibers, run parallel to the folds of the cerebellar
cortex, where they make excitatory synapses with Purkinje cells along the way
(Figure 5.7, rotated view after PLAY). The two-dimensional arbors of the Purkinje
cell dendrites are oriented perpendicular to the parallel fibers. Thus, the
arrangement of Purkinje cells and parallel fibers resembles telephone lines
running between telephone poles. Each parallel fiber makes contact with
hundreds of Purkinje cells; because of the high degree of divergence of the
mossy fiber-granule cell synapses, the firing of each Purkinje cell can be
influenced (disynaptically) by thousands of mossy fibers.

2. Climbing fibers originate exclusively in the inferior olive and make excitatory
projections onto the cerebellar nuclei and onto the Purkinje cells of the
cerebellar cortex. They are called climbing fibers because their axons climb and
wrap around the dendrites of the Purkinje cell like a climbing vine. Each Purkinje
cell receives a single, extremely powerful input from a single climbing fiber. In
contrast to mossy fibers and parallel fibers, each climbing fiber contacts only 10
Purkinje cells on average, making ~300 synapses with each Purkinje cell. Thus,
the climbing fiber is a restricted, but extremely powerful, excitatory input onto
Purkinje cells.
The Purkinje cell is the sole source of output from the cerebellar cortex. It is
important to note that Purkinje cells make inhibitory connections onto the cerebellar
nuclei. (Note the distinction between the Purkinje cells, which constitute the sole output
of the cerebellar cortex, and the cerebellar nuclei, which constitute the sole output of
the entire cerebellum.) Almost all of the spikes generated by the Purkinje cell are
caused by its parallel-fiber inputs. These inputs cause the Purkinje cell to fire at a high
resting rate (~70 spikes/sec), tonically inhibiting its cerebellar nucleus targets. The
powerful inputs from climbing fibers occur less frequently (~1 spike/sec); thus, they
have a minor influence on the overall firing rate of the Purkinje cell. The Purkinje cell
spikes that are generated by climbing fibers are calcium-spikes, however, which allow
the climbing fibers to initiate a number of calcium-dependent changes in the Purkinje
cell. As described below, one important change appears to be a long-lasting change in
the strength of the parallel-fiber inputs to the Purkinje cell.

5.5 Damage to Cerebellum Produces Movement Disorders

Much of what is known about cerebellar function comes from studies of patients with
cerebellar damage. In general, such patients display uncoordinated voluntary
movements and problems maintaining balance and posture. The following are some
symptoms of cerebellar damage (we will discuss more symptoms in the next chapter):
1. Decomposition of movement. Most of our movements involve the coordinated
activity of many muscle groups and different joints to produce a smooth
trajectory of the body part through space. Patients with cerebellar dysfunction
are unable to produce these coordinated, smooth movements. Instead, they
often break the movements down into their component parts in order to execute
the desired trajectory. For example, touching ones finger to ones nose requires
the coordinated activity of shoulder, elbow, and wrist joints. Cerebellar patients
must first perform the shoulder movement, then the elbow movement, and
finally the wrist movement in sequence, rather than as one, uniform motion.
2. Intention tremor. When making a movement to a target, cerebellar patients
often produce an involuntary tremor that increases as they approach closer to
the target. For example, if reaching for a cup, the hand starts out in a direct line
toward the cup; as it gets closer, however, the hand begins to move back and
forth as it attempts to make contact with the cup.

3. Dysdiadochokinesia. Patients have difficulty performing rapidly alternating

movements, such as hitting a surface rapidly and repeatedly with the palm and
back of the hand.
4. Deficits in motor learning. Experimental studies have demonstrated that
cerebellar damage causes deficits in motor learning in both human patients and
experimental animals. One prominent experimental model is
the vestibuloocular reflex (VOR). This reflex allows us to maintain gaze on an
object when the head is rotated (Figure 5.8). Vestibular signals detect the head
movement, and send signals through the cerebellum to the eye muscles to
precisely counter the head rotation and maintain a stable center of gaze. The
motor commands to the eyes must be calibrated precisely with experience, and
this calibration appears to be the job of the cerebellum. Experiments have been
performed in which subjects wore prisms that magnified the visual image. When
the subjects heads were moved, the VOR caused the visual image to shift on the
retina rather than remaining stable. Over days, however, the VOR slowly
adjusted, such that the proper compensatory eye movements were made to
keep the retinal image stable when the head was rotated. In experimental
animals, lesions to the cerebellum prevent this adjustment of the VOR.

Figure 5.8
Vestibuloocular reflex (VOR) and cerebellar
learning. Click PLAY to begin demonstration. Under
normal conditions, when a human or animal subject
rotates the head back and forth, the eyes rotate in
an equal and opposite direction in order to keep the
image stable on the retina. The vestibular system
provides the input regarding the head movement,
and the motor system has to learn the precise
output commands in order to keep the image
stable. When magnifying glasses are placed on the
animal, the eyes do not move fast enough to
compensate for the increased speed of movement
of the magnified image, and thus the image moves
along the retina (termed retinal slip) in the
direction opposite to the movement of the head.
Over time, however, the motor system learns to
move the eyes faster (e.g., the gain of the eye
movement command is increased), and the image
becomes stable again. When the goggles are
removed, the eyes now move too quickly, causing
retinal slip in the same direction as head
movement. With time, the system will learn to
calibrate the VOR again. Patients and experimental
animals with damage to the vestibulocerebellum are
not able to adapt their VOR to the addition and
removal of the goggles, demonstrating the role of
the cerebellum in this form of motor learning.

A second example of cerebellum-dependent motor learning involves the execution of

accurate, coordinated movements. Subjects wore prism goggles that shifted the visual
image to the right, and they were asked to then throw balls at a target on the wall.
Because of the prisms, the accuracy of the subjects was initially quite low, as the balls
consistently hit to the left of the target. With repeated practice, however, the subjects
became more and more accurate at hitting the target. When the goggles were
removed, the subject now began to throw the balls to the right of the target, because
their motor programs had been recalibrated to use the shifted visual input. Over time,
once again, they gradually increased their accuracy. Patients with cerebellar damage
never learned to compensate for the prism, as their balls always landed to the left of
the target when the goggles were worn. When the goggles were removed, they were
immediately accurate at hitting the target, because they never made compensations for
the earlier prism trials.
A third example involves the Pavlovian classical conditioning of the eye blink reflex. In
this task, a neutral stimulus (such as a tone) is paired with a noxious stimulus (such as
a puff of air to the eye) that causes a reflexive eye blink. Over time, experimental
animals will learn to close their eye when the tone occurs, in anticipation of the air puff.
This learned eyelid closure is remarkably well-timed to peak at the expected time of the
puff. Animals with cerebellar damage do not learn to produce the eyelid closure in
response to the tone.

5.6 Cerebellum and Control Systems

What do the various symptoms of cerebellar damage have in common that reveal the
function of the cerebellum? A number of different theories have been proposed. Recall
the discussion in Chapter 1 of the ubiquitous use of sensory information in motor
control. The cerebellum receives extensive sensory input, and it appears to use this
input to guide movements in both a feedback and feedforward control manner.

Feedback control systems

In a feedback controller, a desired output is compared continuously with the actual
output, and adjustments are made during the execution of the movement until the
actual movement matches the desired movement. A common example of a feedback
control system is the thermostat in your home (Figure 5.9).

Figure 5.9
A feedback
control
system,
such as the
thermostat
in your
home, is
sufficient for
slow
movements,
such as
posture. The
myotatic
reflex is an
example of a
feedback
control
system in
the spinal
cord.

The thermostat is set to a desired temperature (e.g., 72), and a thermometer

measures the current temperature in the room. If the thermostat (the comparator)
detects that the room is cooler than the desired temperature, it sends an error signal
that turns on the furnace. If the comparator detects that the room is warmer than the
desired setting, its sends an error signal that turns on the air conditioner.
Feedback control systems can produce very accurate outputs; however, in general they
are slow. In order to change the output, the effector must wait until information is
transmitted from the sensor to the comparator and then to the effector. At this point,
another comparison is made, and the process continues. Consider further the
thermostat example. If the temperature reads 5 cooler than desired, the thermostat

can instruct the furnace to turn on at a moderate heat. It reads the new room
temperature, and, if it is still too cool, it instructs the furnace to deliver more heat, and
so on. Although this will eventually produce an accurate room temperature at the
desired point, it takes a number of cycles to reach that point. One possible solution for
quicker results would be to turn an enormous furnace on full-blast, such that is heats
the room very quickly. This solution, however, can generate another problem. It will
tend to cause the system to oscillate if the feedback pathways are slow. For example,
assume that the furnace can heat the room at the rate of 5 per second, but that it
takes 2 seconds for the thermometer to adjust to the new temperature, and for the new
error signal to turn the furnace off. In those 2 seconds, the furnace has heated the
room up 10, and now it is too warm. So the error signal turns on the air conditioner,
and it cools the room at 5/sec. Of course, it also takes 2 sec to receive the feedback,
and by the time it is told to shut off, it has cooled the room by 10. You can see what
happens: the system will be sent into an endless oscillation of being 5 too hot and 5
too cold. In order for a feedback system to work well, the transmission time of sensory
information through the comparator to the effector must be rapid compared to the time
of the action.
Feedback control systems work well only when the sensory feedback about the actual
output is fast relative to the actual output. If the actual output is faster than the
sensors ability to provide feedback, then the system will tend to oscillate between
overshooting and undershooting the desired output. Thus, a feedback controller is
useful for slow movements, like postural adjustments. The role of the myotatic reflex in
posture maintenance is an example of a feedback controller in the spinal cord, and the
cerebellum plays a role in coordinating these postural adjustments. Feedback control is
not effective for most of the fast movements we make routinely (such as an eye
movement or reaching out for a cup). For these movements, a feedforward controller is
needed.

Feedforward control systems

In a feedforward control system, when a desired output is sent to the controller, the
controller evaluates sensory information about the environment and about the system
itself before the output commands are generated. It uses the sensory information to
program the best set of instructions to accomplish the desired output. However, in a
pure feedforward system, once the commands are sent, there is no way to alter them
(i.e., there is no feedback loop). The advantage of a feedforward system is that it can
produce the precise set of commands for the effector without needing to constantly
check the output and make corrections during the movement itself. The main
disadvantage, however, is that the feedforward controller requires a period of trial-anderror learning before it can function properly. In most biological systems, it is hard
(perhaps impossible) to pre-program all of the possible sensory conditions that the
controller may encounter during the life of the organism. Furthermore, the environment
and conditions under which actions are made are constantly changing, and the
feedforward controller must be able to adapt its output commands to account for these
changes.

Figure 5.10
A
feedforward
control
system is
needed for
fast
movements,
because a
feedback
system is
too slow.

Let us extend the thermostat example to see how a temperature controller operating as
a feedforward system would work to raise the temperature of a room from 70 to 75.
The controller would use diverse sensory information about the environment before
sending its command to the furnace (Figure 5.10). For example, it would read the
current temperature, the current humidity level, the size of the room, the number of
people in the room, and so forth. Based on this information, it would direct the furnace
to turn on for a pre-set period of time, and thats it. There would be no need to
continually compare the current temperature with the desired setting, as the system
has predetermined how long the furnace needs to be working in order to achieve the
desired temperature. How did the controller obtain this information? A feedforward
controller requires a large amount of experience in order to learn the appropriate
actions needed for each set of environmental conditions. If on one trial it turns the
furnace off too soon and the room does not reach the desired temperature, it adjusts its
programming such that the next time it encounters the same environmental conditions,
it turns the furnace on for a longer period of time. Through many such instances of trial
and error learning, the feedforward system creates a look-up table that tells it how
long the furnace needs to be active under the current conditions. The key distinction
between a feedback and feedforward system is that the feedback system uses sensory
information to generate an error signal during the control of a movement, whereas a
feedforward system uses sensory information in advance of a movement. Any error

signal about the final output is used by the feedforward system only to change its
programming of future movements.

The cerebellum may be a feedforward control system

The cerebellar involvement in the VOR may be explained in terms of the learning
requirements of a feedforward controller. When the head moves, a compensatory eye
movement must be made to maintain a stable gaze. The cerebellum receives sensory
input from the vestibular system informing it that the head is moving. It also receives
input from eye muscle proprioceptors and other relevant sources of information about
current conditions in order to make an accurate compensatory eye movement. It
evaluates all of this advance sensory information and calculates the proper eye
movement to exactly counterbalance the head movement. What if the eye movement
does not match the head movement, however, and the visual image moves across the
retina (such as in the experimental condition in which a prism was worn, or in a real-life
situation in which an individual wears new prescription eyeglasses)? The retinal slip
constitutes an error signal to tell the cerebellum that next time these conditions are
met, adjust the eye movement to decrease the retinal slip. This trial and error sequence
will be repeated until the movement is properly calibrated; moreover, these
mechanisms will ensure that the movements stay calibrated.
As another example, the coordination of movements requires that muscle groups be
activated in precise temporal sequence. Not only do the different joints need to be
coordinated temporally, but even antagonist muscles that control the same joint need
precise temporal coordination. For example, an extensor muscle needs to be activated
to start a reaching movement, and the corresponding flexor muscle needs to be
activated at the end of the movement to stop the movement appropriately. The precise
timing of muscle contractions and the force necessary for each contraction varies with
the amount of load placed on a muscle, as well as on the inherent properties of the
muscle itself (e.g., elasticity). These variables are constantly changing throughout life,
as one grows, gains/loses weights, and ages. Moreover, a similar movement will require
different patterns of motor activity depending on the weight being born by the muscle
(for example, if an extended hand is empty or holding a heavy weight). The cerebellum
appears necessary for the proper timing and coordination of muscle groups, very likely
through a trial-and-error learning mechanism discussed previously. Such a role helps
explain the deficits seen in dysdiadochokinesia, in which patients cannot perform rapidly
alternating sequences of movements.
It is believed that the mossy fiber inputs to the cerebellum convey the sensory
information used to evaluate the overall sensory context of the movement. Mossy fibers
are known to respond to sensory stimuli; they are also correlated with different
movements (Figure 5.11). These fibers convey such information as: Where are the
appropriate body parts (proprioceptors), what is the current load on the muscle
(proprioceptors, somatosensory receptors, etc.), what other sensory information can
predict a useful response (e.g., the tone in the eye blink conditioning), what are the
desired movements (motor cortex). The error signal is believed to be conveyed by
the climbing fiber inputs. Climbing fibers are known to be especially active when an
unexpected event occurs, such as when a greater load than expected is placed on a
muscle or when a toe is stubbed. Thus, the large divergence of input from the mossy
fibers to the granule cells to the parallel fibers is believed to create complex

representations of the entire sensory context at present and the desired motor output.
When the desired output is not achieved, the climbing fibers signal this error and
trigger a calcium spike in the Purkinje cell. The influx of calcium changes the connection
strengths between parallel fibers and Purkinje cells, such that the next time the same
behavioral context occurs, the motor output will be modified to more closely
approximate the desired output.

Figure 5.11

Figure 5.12

http://neuroscience.uth.tmc.edu/s3/chapter05.html

The cerebellum may act

as a feedback control
system for slow
movements and a
feedforward controller
for fast movements. In
its function as a
feedforward controller,
the mossy fibers may
provide information
regarding the desired
output from motor
cortex and the advance
sensory information
about the state of the
worlds and the body. The
climbing fibers may
convey information
about movement errors,
which provides a
teaching signal such that
the cerebellum is more
likely to produce the
correct movement the
next time the output is
desired.