If someone's motor cortex is destroyed (by a stroke, for
example), he or she loses the ability to make precise movements,
especially of the hands and fingers. Learning of new movements
is not strongly affected by damage to the cerebral cortex.
The memory of motor sequences learned previously is also largely
spared, though these movements will be executed more clumsily.
These observations show that it is the cerebellum
rather than the cortex that plays an important role in learning
and remembering of movements, also known as procedural
memory.
Area 4 of the precentral gyrus
is not the only area in the cortex that contributes to the pyramidal system. But is the one where movements can be successfully triggered by lower-intensity electrical stimuli. In other words, electrical stimuli that are insufficient to produce movements when applied to other areas of the cortex are sufficient to do so when applied to Area 4.
THE MOTOR CORTEX
So many different structures in the brain are involved in motor functions
that some people even say that practically the entire brain contributes
to body movements. Though the motor cortex is usually associated
with Areas
4 and 6, the control of voluntary movements actually involves
almost all areas of the neocortex.
The primary motor cortex is
the anatomical region composed of Area 4 of the precentral
gyrus. Its location was confirmed in the mid-20th century in
brain operations performed by neurosurgeons such as Dr. Wilder
Penfield, in Montreal. While performing operations to alleviate
patients' epileptic symptoms, Penfield stimulated various areas
of the cortex to identify vital ones that should not be removed.
In this process, he discovered that stimulations applied to
the precentral gyrus triggered highly localized muscle contractions
on the contralateral
side of the body and that there was a somatotopic representation
of the corresponding parts of the body in Area 4 in the primary
motor cortex (see box below) .
Penfield also showed that cortical Area 6, just rostral to Area
4, has two other somatotopic representations that induce complex
movements when stimulated. The first is in the lateral portion
of Area 6 and is called the premotor area (PMA).
It helps to guide body movements by integrating sensory information,
and it controls the muscles that are closest to the body's main
axis.
The second somatotropic representation is in the supplementary motor area
(SMA), in the medial part of Area 6. The SMA is involved
in planning complex movements and in co-ordinating movements involving both hands.
Dr. Penfield's experiments
in stimulating the cortex enabled him to develop a complete
map of the motor cortex, known as the motor homunculus (there
are also other kinds, such as the sensory homunculus).
The most striking aspect of this map is that the areas
assigned to various body parts on the cortex are proportional
not to their size, but rather to the complexity of the
movements that they can perform. Hence, the areas for
the hand and face are especially large compared with
those for the rest of the body. This is no surprise,
because the speed and dexterity of human hand and mouth
movements are precisely what give us two of our most
distinctly human faculties: the ability to use tools
and the ability to speak.
The functions of the
basal ganglia are complex and still largely unknown. People
who have Parkinson's disease, characterized by trembling
and by difficulty in initiating movements, show a deficiency
of dopamine in
their basal ganglia. Because these structures play an important
role in determining various aspects of movement, their
malfunctioning results in the motor problems associated
with Parkinson's disease.
Some abnormalities also are found in the basal ganglia of
people who have Huntington's Disease or Tourette Syndrome.
These patients experience involuntary movements that cause
all sorts of grimaces, tics, and spasms.
THE BASAL GANGLIA
The term "basal
ganglia" refers to a group of several structures in
the brain: the caudate nucleus, the putamen, the globus
pallidus, and the subthalamic nucleus. The substantia nigra,
a midbrain structure that has many interconnections with
the basal ganglia, is not actually part of this grouping
but is often associated with it.
The basal ganglia are involved in a complex
loop that connects them to various areas of the cortex.
The information from the frontal, prefrontal, and parietal
areas of the cortex passes through the basal ganglia, then
returns to the supplementary motor area via the thalamus.
The basal ganglia are thus thought to facilitate movement
by channelling information from various regions of the cortex
to the SMA. The basal ganglia may also act as a filter, blocking
the execution of movements that are unsuited to the situation.
Not all of the circuits involving the basal ganglia are motor circuits, however.
Many are instead involved in memorizing and in cognitive and emotional processing.
A great deal about the basal ganglia remains unknown. They seem to play a far
larger role than just their contribution to motor control.
The cerebellum
also acts as a learning
and memorizing machine,
thanks to its modifiable neural connections that continuously
compare everything they are programmed to do with the results
that they are actually achieving. When this comparison does
not allow the expected result to be achieved satisfactorily,
the cerebellum's activity modifies the sequence of movements
in a compensatory manner to make them more effective. This procedural
memory thus develops automatically with practice,
without the help of any conscious control.
The cerebellum also
appears to play a major role in learning how to co-ordinate
the various segments of the body. The movement of each
segment of your body affects the next, because of its mass.
The cerebellum therefore apparently learns how to calibrate
its commands to the muscles in terms of strength and duration
in order to correct in advance for the effects of these
interactions along the path of motion.
THE
CEREBELLUM
The cerebellum appears to play several roles. It stores
learned sequences of movements, it participates in fine
tuning and co-ordination of movements produced elsewhere
in the brain, and it integrates all of these things to
produce movements so fluid and harmonious that we are
not even aware of them.
To do all this, the cerebellum maintains
close communications with the cortex. The motor, somatosensory,
and posterior parietal areas of the cortex project massive
numbers of axons to the nuclei of the pons, located in the
brainstem. The neurons of the pons then project their axons
into the cerebellum. This corticopontocerebellar tract forms
an extremely dense nerve bundle containing about 20 million
axons, just about 20 times more than the pyramidal bundle!
The
two hemispheres of the cerebellum then sends signals
back to the motor cortex via interconnections in the
ventrolateral nucleus (VLc) of the thalamus. The cerebellar
hemispheres thus influence the muscles of the arms and
legs via the cortex and the lateral
motor system.
The two hemispheres of the cerebellum are not divided neatly
in two like the two hemispheres of the cerebrum. The medial
portion constitutes what is known as the cerebellar vermis.
This vermis does not display any lateralization. It projects
axons to the brainstem which, via the ventromedial
system, help to maintain posture.
The brain mechanisms
that go into planning and executing a movement are far
more complex than the motor cortex's simply issuing a command
and the motor neurons' executing it.
For example, suppose that you go to pick up a glass of water
that you think is cool and refreshing, but is actually boiling
hot. As soon as you touch the glass, you pull your hand back
immediately, by reflex,
without thinking about it.
But suppose that next, your child tries to grab this glass,
which you already know is hot. In this case, because your
child's safety is so important to you, you can consciously
overcome the reflex to pull your hand away. Instead, using
your voluntary motor control, you grab the glass yourself
and put it where your child can't reach it.
Lastly, if someone tells you that the glass is made of fine
crystal and not ordinary glass, you will probably handle
it more carefully. In other words, your brain will take this
information into account and adapt your method of grasping
the glass accordingly.
All of these facts demonstrate that the execution of a movement
is not simply a matter of the brain's sending a "Go!"
command to some motor neurons in the spinal cord, but rather
the result of a highly elaborate construct. Moreover. the
remarkable adaptability of motor activity demonstrates the
involvement of powerful regulatory
and feedback mechanisms.
THE ACTIVATION SEQUENCE FOR
THE MOTOR AREAS
The information
processing that the brain must perform to initiate a voluntary
movement can be divided into three steps. The first step
is to select an appropriate response to
the current situation, out of a repertoire of possible
responses. This response, which corresponds to a particular
behavioural objective, is determined in a global, symbolic
fashion.
The second step is to plan the movement in physical terms. This
step consists in defining the characteristics of the selected response as the
sequence of muscle contractions required to carry it out.
The third step is to actually execute the movement. It is in
this step that the motor neurons are activated that trigger the observable mechanics
of the movement.
Consequently,
the control messages issued by the motor cortex are themselves
triggered by messages from other
cortical areas. The motor cortex also communicates
closely with subcortical structures such as the basal
ganglia and the cerebellum,
through the thalamus, which acts as a relay.
In light of what we now know about the sequence in which
the motor areas of the cortex are activated, we can deconstruct
the classic sequence "Ready? Set. Go!" in terms
of localized activity in the brain.
In the "Ready?" phase, the parietal and frontal
lobes become active first, with a contribution from the
subcortical structures involved in vigilance and attentiveness.
The "Set" command then activates the supplementary
and premotor cortical areas, where the strategies for
movement are developed and maintained until the "Go!" signal
is given. The "Go!" signal may come from an
outside source, as it does in an actual race, or it may
come from inside the person doing the running, who decides
for himself or herself that all the conditions are present
to start running. The "Go!" command then applies
information from subcortical structures such as the basal
ganglia that will influence Area 6, and then eventually
the primary cortex, which will cause the action to be
carried out.