PHYSIOLOGY OF MOTOR CONTROL IN

ACTION SYSTEM

PRESENTED BY: LAXMI BHATTARAI

MPT II
 Content

• Background • Cerebellum and its anatomy and its

• Introduction to motor cortex parts

• Part of the motor cortex
• Details about the corticospinal tract
• Supplementary Motor and Premotor
Areas
• Higher-Level Association Areas
• Basal ganglia and its anatomy and its
parts
• What are the functional differences
between the basal ganglia and the
cerebellum?
• Mesencephalon and Brainstem
• References

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 Objective
• To know the details about the action system and its parts.
• To know about the motor cortex and its role in motor control.
• To know what is the role of the cerebellum in motor control.
• To understand the interconnected pathways of the motor cortex, cerebellum, basal ganglia,
and brain stem, as well as their respective roles in motor control.

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 Background
• The action system includes areas of the nervous system such as the motor cortex, brainstem,
cerebellum, and basal ganglia, which perform processing essential to the coordination of
movement.

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 Motor Cortex
• Primary Motor Cortex and Corticospinal Tract
• The motor cortex is situated in the frontal lobe and consists of a number of different processing
areas, including the primary motor cortex (MI) and two premotor cortical areas, including the
supplementary motor area (SMA), (occasionally called MII), and the premotor cortex.
• These areas interact with sensory processing areas in the parietal lobe and basal ganglia and
cerebellar areas to identify where we want to move, plan the movement, and execute our actions.
• All three of these areas have their own somatotopic maps of the body, so if different regions are
stimulated, different muscles and body parts move. The primary motor cortex (Brodmann’s area
4) contains a very complex map of the body.
• The motor map, or motor homunculus (shown in Fig. 3.12B), is similar to the sensory map in how
it distorts the body's representations. In both cases, the areas that require the most detailed control
(the mouth, throat, and hand), allowing finely graded movements, are most highly represented.

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• Inputs to the motor areas come from the basal ganglia, the cerebellum, and sensory areas,
including the periphery (via the thalamus), SI, and sensory association areas in the parietal
lobe. Interestingly, MI neurons receive sensory inputs from their muscles and the skin above
the muscles.
• Outputs from the primary motor cortex contribute to the corticospinal tract (also called the
pyramidal tract) and often make excitatory monosynaptic connections onto alpha motor
neurons, in addition to polysynaptic connections to gamma motor neurons, which control
muscle spindle length. In addition to their monosynaptic connections, corticospinal neurons
make many polysynaptic connections through interneurons within the spinal cord.

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 Corticospinal Tract
• The corticospinal tract, shown in Figure 3.13, includes neurons from primary motor cortex
(about 50%), and premotor areas including supplementary motor cortex, dorsal and ventral
premotor cortex, and even somatosensory cortex.
• The fibers descend ipsilaterally from the cortex through the internal capsule, the midbrain,
and the medulla. In the medulla, the fibers concentrate to form “pyramids,” near the junction
of the medulla and the spinal cord, most (90%) cross to form the lateral corticospinal tract,
controlling precise movements of the distal muscles of the limbs.
• The remaining 10% continue uncrossed to form the anterior (or ventral) corticospinal tract,
controlling less precise movements of the proximal muscles of the limbs and trunk.
• The majority of the anterior corticospinal neurons cross just before they terminate in the
ventral horn of the spinal cord. Most axons enter the ventral horn and terminate in the
intermediate and ventral areas on interneurons and motor neurons.

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 Supplementary Motor and Premotor Areas
• Each of these areas send projections to primary motor cortex and also to the spinal cord. there
are direct monosynaptic connections from premotor neurons to motor nuclei of the hand and
proximal limb muscles, suggesting that these neurons can control movements separately from
the primary cortex. In addition, these areas receive largely distinct inputs from the thalamus and
other cortical areas. This suggests that they may have very different functions.
• Each of these areas controls different aspects of motor planning and motor learning. Movements
that are initiated internally are controlled primarily by the SMA. This area also contributes to
activating the motor programs involved in learned sequences. The learning of sequences
themselves also involves the presupplementary motor area. The presupplementary motor area is
the rostral extension of the SMA. However, when sequences become overlearned with extensive
training, the control of the movement sequence can be transferred to the primary motor cortex.

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• Movements that are activated by external stimuli (e.g., a visual cue such as a traffic light
changing from red to green) are controlled primarily by the lateral premotor area (dorsal
and ventral premotor cortex). These areas control how stimuli are to be used to direct the
action, specifically associating a given sensory event with a movement to be made. This is
defined as associative learning.
• The study state that premotor and supplementary motor areas differ in their activity
depending on how the movement is initiated and guided , premotor neuron premotor
neurons were more active when a sequential task was visually guided, while
supplementary motor area neurons were more active when the sequence was remembered
and self-determined.
• It is hypothesized that the functional specialization of the SMA and PM based on different
phylogenetic origins, with the SMA being specialized for controlling internally referenced
motor output and the PM area specialized for control of externally referenced motor act.
• Studies also indicate that premotor lesions cause impairment of retrieval of movements in
accordance with visual cues, while SMA lesions disrupt retrieval of self-initiated
movements (Passingham, 1985; Passingham et al., 1989).

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• The supplementary motor area receives inputs from the putamen of the basal ganglia
complex, while the premotor area receives inputs from the cerebellum. In Parkinson’s disease
there is massive depletion of dopamine in the putamen, and patients with Parkinson’s disease
have difficulty with self-initiating movements such as walking. Thus, Parkinson’s disease
may cause impaired input to the supplementary cortex, which results in bradykinesia or
slowness in initiating movement (Marsden, 1989).
• Research suggests that two separate pathways from the parietal cortex to the premotor areas
control reaching and grasping. The reaching pathway originates in the parieto-occipital area
and terminates in the dorsal premotor area, with some neurons synapsing in other areas en-
route. This pathway uses visual information about object location in three-dimensional space
to control the direction of reaching movements. The grasping pathway originates in the dorsal
extrastriate area of the occipital cortex and terminates in the ventral premotor area, with
relays to other areas. This pathway uses visual information about object characteristics
(shape, size, etc.) to control hand shaping for grasping (Krakauer & Ghez, 2000).

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 Higher-Level Association Areas
• Association Areas of the Frontal Regions
• The association areas of the frontal regions (areas rostral to Brodmann’s area 6) are important
for motor planning and other cognitive behaviours.
• The prefrontal cortex may be divided into the principal sulcus and the prefrontal convexities.
Experiments have indicated that the neurons of the principal sulcus are involved in the
strategic planning of higher motor functions. In contrast, lesions in the prefrontal convexity
cause problems in performing any kind of delayed response task..
• Lesions in other prefrontal regions cause patients to have difficulty changing strategies
when asked to. Even when they are shown their errors, they fail to correct them

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 Cerebellum
• The cerebellum is considered one of three important brain areas contributing to the
coordination of movement, in addition to the motor cortex and basal ganglia.
• The cerebellum’s input and output connections are vital to its role as error detector, It receives
information from other modules of the brain related to the programming and execution of
movements (corticopontine areas). This information is often referred to as “efference copy” or
“corollary discharge” when it comes from the primary motor cortex, since it is hypothesized
to be a direct copy of the motor cortex output to the spinal cord.
• The cerebellum also receives sensory feedback information (reafference) from the receptors
about the movements as they are being made (spinal/trigeminal somatosensory inputs, visual,
auditory, and vestibular inputs).
• After processing this information, outputs (Fig. 3.15B) from the cerebellum go to the motor
cortex and other systems within the brainstem to modulate their motor output. In addition to
its role in motor control processes, research has also suggested that the cerebellum may have
important nonmotor functions, including cognition.

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 Anatomy of Cerebellum
• The cerebellum consists of an outer layer of gray matter (the cortex), internal white matter
(input and output fibers), and three pairs of deep nuclei: the fastigial nucleus, the interposed
nucleus, and the dentate nucleus. All the inputs to the cerebellum go first to one of these three
deep cerebellar nuclei and then to the cortex. All the outputs of the cerebellum go back to the
deep nuclei, before going on to the cerebral cortex or the brainstem.
• The cerebellum can be divided into three phylogenetic zones (see Fig. 3.15). The oldest zone
corresponds to the flocculonodular lobe and is functionally related to the vestibular system.
The phylogenetically more recent areas to develop are (a) the vermis and intermediate part
of the hemispheres and (b) the lateral hemispheres, respectively. These three parts of the
cerebellum have distinct functions and distinct input-output connections.

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 Flocculonodular Lobe/
vestibulocerebellum
• Receives inputs from both the
visual system and the vestibular
nuclei.
• It functions in the control of the
axial muscles, which are used in
equilibrium control.
• If a patient experiences
dysfunction in this system, one
observes an ataxic gait, wide-based
stance and nystagmus.
Vermis and Intermediate
Hemispheres/Spinocerebellum
• Receive proprioceptive and cutaneous inputs from the
spinal cord (via the spinocerebellar tracts) and visual,
vestibular, and auditory information.
• four spinocerebellar tracts that relay information from
the spinal cord to the cerebellum. Two tracts relay
information from the arms and the neck, and two relay
information from the trunk and legs. Inputs are also
from the spino-olivo-cerebellar tract, through the
inferior olivary nucleus (climbing fibers). These inputs
are important in learning
• The outputs pathways go to the (a) brainstem reticular
formation, (b) vestibular nuclei, (c) thalamus and motor
cortex, and (d) red nucleus in the midbrain.
• function in the control of the actual execution of
movement: they correct for deviations from an intended
movement by comparing feedback from the spinal cord
with the intended motor command. They also modulate
muscle tone.
• the spinocerebellum is involved in feedforward
mechanisms to regulate movements.
Lateral Hemispheres/Cerebrocerebellum
• Receives inputs from the pontine nuclei in the
brainstem, which relay information from wide
areas of the cerebral cortex (sensory, motor,
premotor, and posterior parietal). Its outputs are
to the thalamus and the motor, premotor, and
prefrontal cortex.
• It is involved in planning or preparing movement
and evaluating sensory information for action as
part of the motor learning process.
• in movement execution and fine-tuning of
ongoing movement via feedback information. It
appears that the lateral hemispheres of the
cerebellum participate in programming the motor
cortex for the execution of movement.
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Role of Cerebellar Involvement in Nonmotor Tasks
• Research has suggested that the lateral cerebellum may have important nonmotor functions,
including cognition. The cerebellum's unique cellular circuitry has been shown to be perfect for the
long-term modification of motor responses, including simple types of learning, such as adaptation.
• Cerebellar learning also appears to occur in the vestibular-ocular reflex (VOR) circuitry, which
includes cerebellar pathways. The VOR keeps the eyes fixed on an object when the head turns. In
experiments in which humans wore prismatic lenses that reversed the image on the eye, an
adaptation of the gain of the VOR occurred over time, with the size of the reflex progressively
reducing and then reversing in direction. This modification of the reflex did not occur in patients
with cerebellar lesions. The cerebellum may also contribute to associative learning, and specifically,
classical conditioning, as lesions to the cerebellum constrain the ability of animals to acquire and
retain the eye-blink reflex.

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• Studies have shown that the right lateral cerebellum becomes active when subjects read verbs
aloud but not when they read nouns, implying that something about the cognitive processing
of verb generation requires the cerebellum, whereas the same processing of other words does
not. Correlated with this, certain patients with cerebellar deficits also showed difficulty in
these verb-generation tasks and in learning and performing a variety of tasks involving
complex nonmotor (cognitive) cortical processing.
• Research on learning problems in patients with cerebellar lesions has shown that while they
had normal scores on the Wechsler Memory Scale, they had problems with some learned
responses. In particular, problems were found in recalling habits, defined as automatic
responses learned through repetition. This is opposite to the learning problems seen in
patients with severe amnesia (resulting from hippocampal and/or midline diencephalic
damage), who do not learn tasks that rely on conscious recall of previous experience but show
normal improvement on a variety of skill learning tasks that involve repetition (Squire, 1986;
Fiez et al., 1992).

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 Basal Ganglia
• The basal ganglia complex consists of a set of nuclei at the base of the cerebral cortex,
including the putamen, caudate nucleus, globus pallidus, subthalamic nucleus, and
substantia nigra. Basal literally means “at the base,” or in other words, “just below the
cortex.”
• As with patients with cerebellar lesions, patients with damage to the basal ganglia are not
paralyzed, but have problems with the coordination of movement.

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 Anatomy of Basal Ganglia
• The globus pallidus has two segments, internal and external (GPi and GPe), and is situated next to
the putamen, while the substantia nigra is situated a little more caudally, in the midbrain, as shown
in the top half of Figure 3.16. The internal segment of the globus pallidus (GPi) and the substantia
nigra (SN) are the major output areas of the basal ganglia. Their outputs terminate in the prefrontal,
supplementary, and premotor cortex areas, by way of the thalamus. The fi nal nucleus, the
subthalamic nucleus (STN), is situated just below the thalamus.
• The connections within the basal ganglia are complex. The striatum receives direct excitatory
cortical inputs and projects to the output nuclei of the basal ganglia (the GPi) and the substantia
nigra pars reticulata (SNr) through two major projection systems, “direct” and “indirect” pathways
(Nambu et al., 2002). The direct pathway begins with projections from the cortex to the striatum,
which project monosynaptically to the GPi/SNr.
• The Gpi projects to the thalamus, which then projects back to the cortex. The connections from the
cortex to the striatum are excitatory, while the connections from the striatum to the GPi and from
the GPi to the thalamus are inhibitory. The connections from the thalamus back to the cortex are
excitatory.

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• The cortex excites the striatum, which then inhibits the GPi through the direct pathway. The
GPi is normally tonically active and inhibitory to the thalamus. When the GPi is inhibited,
this reduces the thalamus’s tonic inhibitory influence (called “disinhibition”), increasing
excitatory drive to the cortex and reinforcing the desired movement.
• The indirect pathway begins with projections from the cortex to the striatum, from the
striatum to the external segment of the Gpe, which projects to the STN, and then to the Gpi.
The Gpi projects to the thalamus, which projects back to the cortex. The projections from the
striatum to the Gpe, and from GPe to STN are inhibitory, while the projections from the STN
to the Gpi are excitatory.
• Input from the cortex excites the striatum, which then inhibits Gpe. Since Gpe is inhibitory to
the STN, the STN becomes more active and excites the Gpi. Increased activation of the Gpi
inhibits the thalamus, and as a result the thalamus does not excite the cortex. In this way,
activation of the indirect pathway by the striatum causes a relative inhibition of movement.

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• The basal ganglia really consist of four different functional circuits that also include the
thalamus and the cortex. These comprise the skeletomotor circuit (including the premotor
cortex, supplementary motor cortex and primary motor cortex), the oculomotor circuit
(including the frontal and supplementary eye fi elds of the cortex), the prefrontal circuits, and
the limbic circuit. The existence of these different functional circuits explains the variety of
movement disorders involving the dysfunction of basal ganglia (DeLong, 2000).

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Role of the Basal Ganglia
• The skeletomotor circuit contributes to both the preparation for and execution of movement.
• It has also been hypothesized that the circuitry of the basal ganglia may play a role in
selectively activating some movements as it suppresses others (Alexander & Crutcher, 1990).
Nambu and associates (2002) also propose that the direct, indirect, and hyperdirect pathways
within the basal ganglia play an important role in selectively activating some movements
while inhibiting others. They suggest that when a voluntary movement is about to be initiated
by cortical mechanisms, a corollary signal is transmitted simultaneously from the motor
cortex to the GPi through the cortico-STN-pallidal hyperdirect pathway. Activation of the GPi
neurons results in the inhibition of large areas of the thalamus and cortex that are related to
both the selected motor program and other competing programs.
• Another corollary signal is sent via the direct pathway to the GPi to inhibit a specific
population of pallidal neurons in the center area, which disinhibit their targets and release
only the selected motor program. Finally, the third corollary signal possibly through the
indirect pathway reaches the GPi to activate neurons therein and causes inhibition of
competing motor programs. Through this sequential information processing, only the selected
motor program is initiated, executed, and terminated at the appropriate time, whereas other,
competing, programs are inhibited.
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• The oculomotor circuit is involved in the control of saccadic eye movements. The prefrontal
circuit and the limbic circuits are involved in nonmotor functions. The prefrontal circuits
contribute to executive functions, including organizing behaviors using verbal skills in
problem solving and mediating socially appropriate responses. Lesions in this area contribute
to obsessive–compulsive disorder. The limbic circuit is involved in the control of motivated
behavior (involving circuits for reinforcing stimuli for behaviors) and procedural learning.
• Most disorders of the basal ganglia involve problems with action rather than perception. They
may involve either hyperactivity/impulsivity (e.g., Huntington’s disease or obsessive–
compulsive disorder) or reduced activity and flat affect (e.g., Parkinson’s disease, depression)
(DeLong, 2000).

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What are the functional differences between the basal
ganglia and the cerebellum?
• Research suggests that the basal ganglia may be particularly concerned with internally
generated movements, while the cerebellum is involved in visually triggered and guided
movements. For example, experiments have shown that in the internal globus pallidus, cells
that project to the supplementary motor area are activated during internally generated
movements.
• This is consistent with clinical data demonstrating that patients with Parkinson’s disease have
a great deal of difficulty with internally generated movements (Georgiou et al., 1993; Morris
et al., 1996). It is interesting to note that patients with Parkinson’s disease with frozen gait
syndrome (difficulty initiating or maintaining gait) are able to use visual cues to improve their
walking abilities. The above research suggests that this may be due to the use of alternative
pathways from the cerebellum to trigger and guide the movements.

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 Mesencephalon and Brainstem
• The nuclei and pathways from the mesencephalon and brainstem to the spinal cord mediate
many aspects of motor control as part of descending pathways from the cerebral cortex,
cerebellum, and basal ganglia.
• This includes the generation of locomotor rhythms, the regulation of postural tone, the
integration of sensory information for posture and balance, as well as contributions to
anticipatory postural control accompanying voluntary movements.
• Stimulation of the mesencephalic locomotor region (and also the subthalamic locomotor
region initiates locomotion and adjusts stepping movements). Signals from this system are
relayed to the spinal cord central pattern generators for locomotion via the medial reticular
formation and reticulospinal pathways (including the pontomedullary locomotor strip).
These pathways and brainstem centers are shown in Figure 3.17A.

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• The brainstem has important centers for controlling the facilitation and inhibition of muscle
tone important for the control of posture. These muscle tone facilitatory and inhibitory
systems within the brainstem are shown in Figure 3.17, A and B.
• when the brainstem reticular formation is inactivated by pharmacologic means, anticipatory
postural adjustments that would normally be activated to stabilise a voluntary movement
initiated through activation of the motor cortex are no longer activated. This indicates the
importance of brainstem nuclei in anticipatory postural control.
• Basal ganglia– cortical–spinal pathways are important to the control of voluntary movements,
while basal ganglia–brainstem–spinal cord pathways contribute to automatic control of
movements such as locomotion and postural tone mainly via pathways originating in the
substantia nigra.

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• Tracts for motor control that originate in the brainstem consist of the medial pathways
(controlling posture and balance), including the vestibulospinal tract (discussed in the
vestibular section of this chapter), the reticulospinal tract (discussed above), the tectospinal
path (from the superior colliculus, and mediating head and eye movement), and the lateral
pathways, controlling goal-directed movements, and including the rubrospinal pathway from
the red nucleus. Figure 3.19 summarizes the major descending spinal tracts from both the
cortex and the brainstem.

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 References
• Shumway-Cook A, Woollacott MH. Motor control: translating research into clinical practice.
Lippincott Williams & Wilkins; 2007.
• Lazaro RT, Reina-Guerra SG, Quiben M, editors. Umphred's Neurological Rehabilitation:
Umphred's Neurological Rehabilitation-E-Book. Elsevier Health Sciences; 2019 Dec 5.

• O'Sullivan SB, Schmitz TJ, Fulk G. Physical rehabilitation. FA Davis; 2019 Jan 25.

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Thank you