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UNIT 15. CEREBELLUM

GENERALITIES. CORTICAL SUPPORT NUCLEI. BASAL GANGLIA. CEREBELLUM.

FUNCTION OF THE CEREBELLUM.

- GENERALITIES

So far, we have studied subconscious motor functions embedded in the spinal cord
and brainstem—involuntary or reflex movements; and furthermore, we know how
the motor cortex executes voluntary movements. However, the motor control
function of the cortex requires the unavoidable collaboration of other structures
located in different regions of the brain, namely: the cortical nuclei of the brainstem,
the basal ganglia and the cerebellum.

Figure 15-1. Lateral and dorsal view of the cortical nuclei of the brainstem.

- CORTICAL SUPPORT NUCLEI.

At the level of the brainstem, a series of nerve nuclei are located that
collaborate in motor activity and act as an emergency mechanism in
nervous system of domestic animals Topic 1 5 Cerebellum

case of cortical injury. In order of importance we highlight: the red nucleus, the
substantia nigra, the bridge nuclei and the olivary nucleus (Fig.15-1).

1) The red core, located in the midbrain, is highly developed in

domestic mammals and has giant pyramidal cells (cells of Betz), similar in
morphology and distribution, to those of the motor cortex. These neurons give rise to
rubrospinal tract (extrapyramidal pathway), which crosses to the opposite side of
the brainstem and continues parallel to the corticospinal tract in search of the
interneurons and motor neurons of the spinal cord. The rubrospinal pathway is even
faster than the corticospinal pathway and serves as an accessory route for
transmission of cortical signals to the spinal cord. Therefore, if we were to destroy the
corticospinal tract, coarse and coarse movements could still be developed, since to
carry out fine movements the cortex must intervene directly.

The red nucleus, like the motor cortex, has somatotopic representation of all
the muscles of the body. Therefore, experimental electrical stimulation of a single
point in the red nucleus will cause the contraction of a muscle or small group of
muscles.

On the other hand, in the red nucleus there is a greater number of dynamic
pyramidal neurons than static pyramidal ones, contrary to what occurs in the motor
cortex; therefore, the red nucleus is more related to the initiation of movements
while the motor cortex is related to the maintenance of movements. In addition, this
nucleus is functionally linked to the cerebellum, helping to regulate faster muscle
movements.

2) The black substance It is a nuclear formation located in the mesencephalon,

on the border between the cerebral peduncles and the mesencephalic roof, and
whose function is to help the basal ganglia regulate muscle tone. The experimental
destruction of the neurons of the substantia nigra causes a muscular tremor, similar
to that produced in the human species in theParkinson's disease.

3) The reticular bridging nuclei are small neuronal groups

Located at the level of the bridge of the brainstem and in addition to controlling the
antigravity muscles, they function as exchange centers for the pyramidal motor pathways
and with the cerebellum through the reticulocerebellar pathway.

4) The olive core (bulbar olive), located in the medulla oblongata, functions
as a relay station between brain and mesencephalic centers and the cerebellum, via
the olivocerebellar pathway.

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- BASAL GANGLIA
The basal ganglia are nuclear, bilateral and subcortical formations, present in
all vertebrates and located in close relationship with the cerebral cortex, cortical
nuclei of the brainstem, cerebellum and thalamus.
In mammals, the basal ganglia consist primarily of the caudate nucleus,
putamen and the pale globe. In birds, because they lack a motor cortex, the basal
ganglia take on a unique development, taking over much of the functions that the
motor cortex performs in mammals.

Severe lesions in the motor cortex are known to cause loss of movement
(paralysis), mild lesions in the motor cortex cause loss of fine and precise movements,
however, lesions in the basal ganglia or cerebellum tend to cause abnormalities. of
movement and not to eliminate it. Even a decorticated dog or cat, in addition to its
great muscular hypertonus, is still capable of surviving by coordinating certain vital
activities, such as eating, fighting, defending itself, etc.; but if we destroyed its basal
ganglia, its hypertonus would be exaggerated and the animal would almost only
conserve the motor activity typical of a mesencephalic animal, with hardly any chance
of survival.

The basal ganglia (Fig.15-2-B-) act in close relationship with the cerebral cortex
to control not only motor functions but also sensitive ones; To do this, the basal
ganglia receive input from a large area of the motor cortex (-Love-), of the cerebral
cortex (-Ac-) and the brainstem (-At-), and send signals that are exchanged in the
nuclei of the thalamus (-C-), reach the motor cortex (-D-) from which motor signals are
directed to the brainstem (-AND-); at this level, they cross to the opposite side and
reach the motor neurons of the spinal cord and from here to the muscles (-F-). It is
interesting to note that the basal ganglia are activated with maximum frequency not
by signals from the motor cortex but by signals that arrive directly from the cerebral
cortex or the brainstem (afferent pathways) and, furthermore, this activation takes
place before muscle contraction to occur.

Figure 15-2. Sagittal view of the basal ganglia in

monkey. See explanation in the text.

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There is a firm belief that the basal ganglia have an essential role in the
initiation of movements, with the help of the red nucleus, and in the control of slow
movements (since the cerebellum is in charge of the fast ones). In addition, the basal
ganglia are primarily responsible for the inhibition of motor tone, with the help of the
substantia nigra. These inhibitory signals are sent directly to the motor cortex and
brainstem.

- CEREBELLUM

The cerebellum It is the region of the brain where the correlation between
anatomy and its function has been determined with greater accuracy, and even the
diagram of its intricate nervous circuits has been known with total accuracy. In our
domestic animals, the cerebellum is anatomically located above the brainstem and is
divided into three lobes:rostral (neocerebellum), flow (paleocerebellum) and
flocculonodular(archicerebellum). The latter is the most primitive of the entire
cerebellum and works in collaboration with the vestibular apparatus of the inner ear,
in the control of balance (Fig.15-3).

Representation topographical
of the cerebellum. In the same way
that the sensory cortex, the motor
cortex and the red nuclei have
topographical or somatotopic
representation of the different parts of
the body, so it is with the cerebellum.
However, the somatotopy is not
organized in the lobes but along the
longitudinal axis, showing three
defined longitudinal zones (Fig.15-3); a
narrow central areavermis, in the
midline; in this area it Figure 15-3. Schematic representation of
cerebellum.

they localize most of the cerebellar functions of control for the muscular movements
of the axis of the body (neck, back and hip). Toward each side of the vermis is a large
cerebellar hemisphere divided into an intermediate zone orparavermal zoneand one
side area. The paravermal zone of each hemisphere is related to the control of
muscular contractions of the extremities (hands and legs) on the opposite side of the
body. However, the lateral zones are related to the preparation of the movements
that the animal is immediately ready to carry out.cerebellar cortex.

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It wasCajalwho identified the neurons of the cerebellar cortex and their

connections. Thus, in the cerebellar cortex five types of neurons can be found,
distributed in three well-differentiated layers (Fig.15-4): to) In the granular layer,
more internal, are located the granule cells, small and abundant and the cells of
Golgi. The axons of the granules ascend to the outermost (molecular) layer and there
they bifurcate and give rise to the so-called parallel fibers. The dendrites of the cells
Golgi they also extend to the molecular layer. b) The middle layer contains large
neurons cells of Purkinje, with huge dendritic arborizations to the molecular layer
and axons that are the only fibers in the cortex that go to the deep cerebellar nuclei.
c) Lastly, the molecular Layer, outermost, it is traversed by the parallel fibers of the
granule cells and includes dendritic arborizations of the granule cells.Golgi and of
Purkinjeand contains two other classes of neurons,starry Y in basket.

Figure 15-4. Neurons of the cerebellar cortex in the dog.

The cerebellum, in addition to the cerebellar cortex, has an internal region occupied
by numerous input and output fibers and also has threedeep cerebellar nucleiand
bilateral:fastigial, interposedY jagged. The fastigial nucleus is related to the vermis and
the flocculonodular lobe, the dentate nucleus is related to the lateral zone, and the
interposed nucleus is related to the paravermal zone.

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pathways to the cerebellum. (Fig.15-5) The most important pathway iscortico-

pontocerebellar pathway(-Love-) that brings information mostly from the motor cortex
and some from the rest of the cortex
sensory, crosses the bridge of the
brainstem and reaches the cerebellum.
The pathways that provide information
from the brainstem are:via

reticulocerebellar(-At-) that starts from the

reticular formation and theolivocerebellar
pathway(-At-), which runs from the olivary
nucleus to the cerebellum.

From outside the brain, the routes

of greatest contribution of information
are:vestibulocerebellar pathway(-Ap-),
which provides information directly
from the vestibular apparatus to the
flocculonodular lobe of the cerebellum;
and thespinocerebellar pathway(-A.M-
) that provides information from the
Figure 15-5. Sagittal view of the monkey cerebellum.
proprioceptors, through the spinal
See explanation in the text.
cord, to know the state of contraction of
the animal's muscle groups.

The deep nuclei of the cerebellum receive signals from the input pathways;
These go in two directions: a) directly to one of the deep nuclei, and b) to the
corresponding area of the cerebellar cortex, above the deep nucleus.

Outflow pathways from the cerebellum. These originate in the deep nuclei of
the cerebellum (fastigial, interposed, and dentate) in addition to the vestibular nuclei
of the medulla oblongata and pons, which also function as "deep nuclei" given their
direct connection to the flocculonodular lobule of the cerebellum.

- FUNCTION OF THE CEREBELLUM

The cerebellum has long been called the "silent area" of the brain, since its
electrical stimulation does not cause sensation or movement. However, its removal
causes almost total incoordination even though it does not produce paralysis of any
muscle. The cerebellum is especially useful for the control of the most rapid and
skillful muscular activities of our animals.

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domestic; like the basal ganglia they are for the control of slow movements.

Unlike the basal ganglia, the cerebellum receives great info via the
spinocerebellar pathwayFig.15-5) of the proprioceptors that report the length and
tension of the muscles and the state of the joints at any time, compares them with
the rest of the signals provided by the input pathways and causes an output pathway
that goes to the motor nuclei thalamic (-C-) to subsequently reach the motor cortex (
-D-). When the animal is at motor rest, the outputs of the motor nuclei are minimal;
but when it is in motion, those outputs increase due to the processing of afferent
signals.

But how is it that the cerebellum can be so important if it does not establish
direct control over muscle contraction? The answer lies in the fact that the cerebellum
is responsible forlook out Y set corrective settings of motor activities promoted by
other parts of the brain. To do this, the cerebellum receives input from a large area of
the cerebral cortex (- Master-, -Ac-) and peripheral regions of the body, in order to
determine the instantaneous state of each of its areas: its position, its rate of
movement and the forces acting on it.

The cerebellum compares the current physical state of each body part, as
indicated by sensory information (proprioceptors), with the state that the motor
system is trying to produce. If the two are not favorably related, appropriate
corrective signals are instantly transmitted to the motor system via the fast track of
the pyramidal pathways (-AND-) to motor neurons (-F-), to increase or decrease the
activity of certain specific muscles.

Because the cerebellum has to make important motor adjustments very quickly
in the course of movement, an extensive, very fast-acting input system is needed,
entering the cerebellum both from peripheral parts of the body and from the motor
areas of the brain. . An extensive output system is also needed to work just as quickly
sending signals to the motor system to make the appropriate corrections to the
motor signals.

Cerebellar areas in motor activity.Although it is difficult to give a clear picture

of the motor functions of the cerebellum, we assign to the various regions that
compose it, the following actions (Fig.15-3):

one)vestibular area. Includes the flocculonodular lobe. It is important for

regulation of balance, receiving and sending vestibular information (vestibular nuclei)
necessary for maintaining balance, whether in a static or dynamic situation.

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two) Vermis. Receives proprioceptive information from muscles and joints

that allow the vermis to exercise control over the cortical and brainstem components,
favoring the maintenance of posture, support and support movements, adequate
muscle tone and body balance.

3) paravermal area. Receive somesthetic information and "copies" of the

Execution signals of the motor program, with the possibility of relating them to the
state of the muscle and joints at each moment. It plays an important role in course
rectification of many movements by sending corrections and adjustments via the red
nucleus and rubrospinal tract or via nuclei from the thalamus to the motor cortex and
corticospinal tract.

4) cerebellar hemispheres. The great development of the hemispheres

cerebellar in our primates has been related to the ability to execute rapid, well-
organized and skillful movements. Each hemisphere receives several million afferent
fibers (called mossy fibers) from pontine nuclei that synapse with granule and spleen
cells.Golgi in the granular layer; their outputs go to the nuclei of the thalamus and
from there to the motor cortex.

These hemispheres are thought to play an important role, paralleling that of

the basal ganglia, in transforming a movement plan generated in the association
cortex into an organized motor program in the motor cortex, especially for well-
learned movements. , which include well-organized fast and alternating sequences,
so fast that they hardly allow adjustments or so well learned that they do not require
those adjustments.

It has also been revealed that the cerebellar hemispheres are important in
learning and routinizing new skillful movements, so that they are executed almost
automatically and without having to think about how to do them.

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