Motor system

• Posture and movement depend on a

combination of involuntary reflexes
coordinated by the spinal cord and voluntary
actions controlled by higher brain centers.
 Organization of Motor Function
by the Spinal Cord
• Posture and movement ultimately depend on
contraction of some skeletal muscles while,
simultaneously, other muscles remain relaxed.
• Recall that activation and contraction of
skeletal muscles is under the control of the
motoneurons that innervate them.
• The motor system is designed to execute this
coordinated response largely through reflexes
integrated in the spinal cord.
 Motor Units
• A motor unit is defined as a single
motoneuron and the muscle fibers that it
innervates.
• The number of muscle fibers innervated can
vary from a few fibers to thousands of fibers,
depending on the nature of the motor activity.
• Thus, for eye movements requiring fine
control, motoneurons innervate only a few
muscle fibers.
• For postural muscles involved in large
movements, motoneurons innervate
thousands of muscle fibers.
• A motoneuron pool is the set of motoneurons
innervating fibers within the same muscle.
• The force of contraction of a muscle is graded
by recruitment of motor units (size principle).
• For example, small motoneurons innervate a
few muscle fibers, and, because they have the
lowest thresholds, they fire first.
• Small motoneurons also generate the smallest
amounts of force.
• On the other hand, large motoneurons
innervate many muscle fibers.
• They have the highest thresholds to fire action
potentials; thus, they fire last.
• Because large motoneurons innervate many
muscle fibers, they also generate the greatest
amounts of force.
• The size principle states that as more motor
units are recruited, progressively larger
motoneurons are involved and greater tension
will be generated.
 Types of Motoneurons
• There are two types of motoneurons:
1. α motoneurons and
2. γ motoneurons.
• α Motoneurons innervate extrafusal skeletal
muscle fibers.
• Action potentials in α motoneurons lead to
action potentials in the extrafusal muscle fibers
they innervate, which results in contraction.
• γ Motoneurons innervate specialized
intrafusal muscle fibers, a component of the
muscle spindles.
• The overall function of the muscle spindle is to
sense muscle length; the function of the γ
motoneurons innervating them is to adjust the
sensitivity of the muscle spindles (so that they
respond appropriately as the extrafusal fibers
contract and shorten).
• α Motoneurons and γ motoneurons are
coactivated (activated simultaneously) so that
muscle spindles remain sensitive to changes in
muscle length even as the muscle contracts
and shortens.
 Types of Muscle Fibers
• There are two types of muscle fibers:
1) Extrafusal fibers and
2) Intrafusal fibers.
• Extrafusal fibers constitute the majority of skeletal
muscle, are innervated by α motoneurons, and are
used to generate force.
• Intrafusal fibers are specialized fibers that are
innervated by γ motoneurons and are too small to
generate significant force.
• Intrafusal fibers are encapsulated in sheaths, forming
muscle spindles that run parallel to the extrafusal
fibers.
 Muscle Spindles
• Are distributed among the extrafusal muscle
fibers, and they are especially abundant in
muscles utilized for fine movements (e.g.,
muscles of the eye).
• Are spindle-shaped organs composed of
intrafusal muscle fibers and innervated by
sensory and motor nerve fibers.
• Are attached to connective tissue and
arranged in parallel with the extrafusal muscle
fibers.
Intrafusal Muscle Fibers of Muscle Spindles

• There are two types of intrafusal fibers

present in muscle spindles:
I. Nuclear bag fibers and
II. Nuclear chain fibers
• Generally, both types of fibers are present in
every muscle spindle, but nuclear chain fibers
are more plentiful than nuclear bag fibers.
• There are five or six nuclear chain fibers per
muscle spindle, compared with two nuclear
bag fibers.
• Nuclear bag fibers are larger, and their nuclei
are accumulated in a central (“bag”) region.
• Nuclear chain fibers are smaller, and their
nuclei are arranged in rows (“chains”).
 Innervation of Muscle Spindles
• Muscle spindles are innervated by both
sensory (afferent) and motor (efferent)
nerves.
• Sensory innervation of the muscle spindle
consists of a single group Ia afferent nerve,
which innervates the central region of both
the nuclear bag fibers and the nuclear chain
fibers, and group II afferent nerves, which
primarily innervate the nuclear chain fibers.
• Recall that group Ia fibers are among the
largest nerves in the body; thus, they have
among the fastest conduction velocities.
• These fibers form primary endings in a spiral-
shaped terminal around the central region of
the nuclear bag and nuclear chain fibers.
• Group II fibers have intermediate diameters
and intermediate conduction velocities.
• Group II fibers form secondary endings
primarily on the nuclear chain fibers.
• Motor innervation of the muscle spindle
consists of two types of γ motoneurons:
A. dynamic and
B. static.
• Dynamic γ motoneurons synapse on nuclear
bag fibers in “plate endings.”
• Static γ motoneurons synapse on nuclear
chain fibers in “trail endings,” which spread
out over longer distances.
• γ Motoneurons are smaller and slower than
the α motoneurons that innervate the
extrafusal fibers.
• The function of the γ motoneurons (either
static or dynamic) is to regulate the sensitivity
of the intrafusal muscle fibers they innervate.
 Function of Muscle Spindles
• Muscle spindles are stretch receptors whose
function is to correct for changes in muscle
length when extrafusal muscle fibers are
either shortened (by contraction) or
lengthened (by stretch).
• Thus, muscle spindle reflexes operate to
return muscle to its resting length after it has
been shortened or lengthened.
• To illustrate the function of the muscle spindle
reflex, consider the events that occur when a
muscle is stretched.
a) When a muscle is stretched, the extrafusal
muscle fibers are lengthened.
• Because of their parallel arrangement in the
muscle, the intrafusal muscle fibers also are
lengthened.
b) The increase in length of the intrafusal fibers
is detected by the sensory afferent fibers
innervating them.
• The group Ia afferent fibers (innervating the
central region of nuclear bag and nuclear
chain fibers) detect the velocity of length
change, and
• The group II afferent fibers (innervating the
nuclear chain fibers) detect the length of the
muscle fiber.
• Thus, when the muscle is stretched, the
increase in the length of the intrafusal fibers
activates both group Ia and group II sensory
afferent fibers.
c) Activation of the group Ia afferent fibers
stimulates α motoneurons in the spinal cord.
• These α motoneurons innervate extrafusal
fibers in the homonymous (same) muscle and,
when activated, cause the muscle to contract
(i.e., to shorten).
• Thus, the original stretch (lengthening) is
opposed when the reflex causes the muscle to
contract and shorten.
• γ Motoneurons are coactivated with the α
motoneurons, ensuring that the muscle
spindle will remain sensitive to changes in
muscle length even during the contraction.
 Spinal Cord Reflexes
• Spinal cord reflexes are stereotypical motor responses
to specific kinds of stimuli, such as stretch of the muscle.
• The neuronal circuit that directs this motor response is
called the reflex arc.
• The reflex arc includes
o the sensory receptors;
o the sensory afferent nerves, which carry information to
the spinal cord;
o the interneurons in the spinal cord; and
o the motoneurons, which direct the muscle to contract or
relax.
• The stretch reflex is the simplest of all spinal
cord reflexes, having only one synapse
between sensory afferent nerves and motor
efferent nerves.
• The Golgi tendon reflex is of intermediate
complexity and has two synapses.
• The most complex of the spinal cord reflexes is
the flexor-withdrawal reflex, which has
multiple synapses.
 Stretch Reflex
• The stretch (myotatic) reflex is exemplified by
the knee-jerk reflex.
• The following steps occur in the stretch reflex,
which has only one synapse between the
sensory afferent nerves (group Ia afferents)
and the motor efferent nerves (α
motoneurons):
a. When the muscle is stretched, group Ia
afferent fibers in the muscle spindle are
activated and their firing rate increases.
• These group Ia afferents enter the spinal cord
and synapse directly on and activate α
motoneurons.
• This pool of α motoneurons innervates the
homonymous muscle.
b. When these α motoneurons are activated, they
cause contraction of the muscle that was
originally stretched (the homonymous muscle).
• When the muscle contracts, it shortens, thereby
decreasing stretch on the muscle spindle.
• The muscle spindle returns to its original length,
and the firing rate of the group Ia afferents
returns to baseline.
c. Simultaneously, information is sent from the
spinal cord to cause contraction of synergistic
muscles and relaxation of antagonistic
muscles.
• The stretch reflex is illustrated by the knee-
jerk reflex, which is initiated by tapping the
patellar tendon, causing the quadriceps
muscle to stretch.
• When the quadriceps and its muscle spindles
are stretched, group Ia afferent fibers are
stimulated.
• These group Ia afferent fibers synapse on and
activate α motoneurons in the spinal cord.
• These α motoneurons innervate and cause
contraction of the quadriceps (the muscle that
originally was stretched).
• As the quadriceps muscle contracts and
shortens, it forces the lower leg to extend in
the characteristic knee-jerk reflex.
 Golgi Tendon Reflex
• The Golgi tendon reflex is a disynaptic spinal
cord reflex, which is also called the inverse
myotatic reflex (inverse or opposite of the
stretch reflex).
• The Golgi tendon organ is a stretch receptor
found in tendons, which senses contraction
(shortening) of muscle and activates group Ib
afferent nerves.
• Golgi tendon organs are arranged in series
with the extrafusal muscle fibers (contrasting
the parallel arrangement of muscle spindles in
the stretch reflex).
• The steps in the Golgi tendon reflex are
described as follows:
i. When the muscle contracts, the extrafusal
muscle fibers shorten, activating the Golgi
tendon organs attached to them.
• In turn, the group Ib afferent fibers that
synapse on inhibitory interneurons in the
spinal cord are activated.
• These inhibitory interneurons synapse on the
α motoneurons.
ii. When the inhibitory interneurons are
activated (i.e., activated to inhibit), they inhibit
firing of the α motoneurons, producing
relaxation of the homonymous muscle (the
muscle that originally was contracted).
iii. As the homonymous muscle relaxes, the
reflex also causes synergistic muscles to relax
and antagonistic muscles to contract.
 Flexor-Withdrawal Reflex
• The flexor-withdrawal reflex is a polysynaptic
reflex that occurs in response to tactile,
painful, or noxious stimulus.
• Somatosensory and pain afferent fibers
initiate a flexion reflex that causes withdrawal
of the affected part of the body from the
painful or noxious stimulus (e.g., touching a
hand to a hot stove and then rapidly
withdrawing the hand).
• The reflex produces flexion on the ipsilateral
side (i.e., side of the stimulus) and extension
on the contralateral side.
• The steps involved in the flexor-withdrawal
reflex are explained as follows:
1. When a limb touches a painful stimulus (e.g.,
hand touches a hot stove), flexor reflex
afferent fibers (groups II, III, and IV) are
activated.
• These afferent fibers synapse on multiple
interneurons in the spinal cord (i.e.,
polysynaptic reflex).
2. On the ipsilateral side of the pain stimulus,
reflexes are activated that cause flexor
muscles to contract and extensor muscles to
relax.
• This portion of the reflex produces flexion on
the ipsilateral side (e.g., withdrawal of the
hand from the hot stove).
3. On the contralateral side of the pain stimulus,
reflexes are activated that cause extensor
muscles to contract and flexor muscles to
relax.
• This portion of the reflex produces extension
on the contralateral side and is called the
crossed-extension reflex.
• Thus, if the painful stimulus occurs on the left
side, the left arm and leg will flex or withdraw
and the right arm and leg will extend to
maintain balance.
4. A persistent neural discharge, called an
afterdischarge, occurs in the polysynaptic
reflex circuits.
• As a result of the afterdischarge, the
contracted muscles remain contracted for a
period of time after the reflex is activated.
 HIGHEST LEVEL OF MOTOR CONTROL
• CEREBRAL CORTEX:
• The highest level of motor control is exerted
through motor cortex and two major
descending pathways emerging from the
motor areas
• Voluntary movements are directed by the
motor cortex, via descending pathways.
• The motivation and ideas necessary to
produce voluntary motor activity are first
organized in multiple associative areas of the
cerebral cortex and then transmitted to the
supplementary motor and premotor cortices
for the development of a motor plan
• The motor plan will identify the specific
muscles that need to contract, how much they
need to contract, and in what sequence.
• The plan then is transmitted to upper
motoneurons in the primary motor cortex,
which send it through descending pathways to
lower motoneurons in the spinal cord.
• The planning and execution stages of the plan
are also influenced by motor control systems
in the cerebellum and basal ganglia.
• The motor cortex consists of three areas:
primary motor cortex, supplementary motor
cortex, and premotor cortex
 Motor cortex
• Areas of motor cortex include:
Primary motor cortex (Brodmann’s area 4):
• It is organized in terms of movements rather
than the individual muscles, e.g. stimulation
reveals discrete isolated movements on
opposite half of the body.
Premotor cortex:
• It is located immediately anterior to the lateral
portion of the primary cortex.
• It includes Brodmann’s area 6, 8, 44 and 45.
 Supplementary motor cortex: is located in
the medial surface of frontal lobe rostral to
the primary motor area
 Functional role of motor cortex in
control of voluntary movements
• Supplementary motor cortex is responsible for
generating the idea for a movement.
• There it plans the movements.
• Lateral cerebellum and basal ganglia are also
involved in the planning and programming of
movements.
• Primary motor cortex is responsible for the
execution of movement.
• Programmed patterns of motor neurons are
activated in the motor cortex.
• Premotor cortex co-ordinates the voluntary
activity:
• Area 6: co-ordinates the proximal and axial
muscles during a motor task.
• It ensures that the skilled movement are
accurate and smooth.
• Area 8: also called frontal eye field, is involved
in the co-ordination of eye movements.
• Area 44 and 45: also called Broca’s motor
speech areas ( are engaged in co-ordination of
the activity of musculature involved in speech.
• E.g activation of vocal cords simultaneously
with the movement of mouth and tongue
during speech.
 Descending motor pathways from motor
cortex
• The motor tract originating from the cerebral
cortex traditionally has been called the
pyramidal tract because it traverses the
medullary pyramids on its way to the spinal
cord.
• This path is the corticospinal tract (CST).
• All other descending motor tracts emanating
from the brainstem were generally grouped
together as the extrapyramidal system.
 Pyramidal tracts
• The CST has approximately 1 million nerve fibres
with an average conduction velocity of
approximately 60m/s using glutamate as their
transmitter substance.
• Corticospinal tracts include 30% fibers arising from
the primary cortex (area 4), 30% from premotor
area (area 8) and supplementry cortex and 40%
arising from the somatic sensory cortex (Brodmann
areas 1, 2, and 3).
• All the above fibres form the fibres of upper motor
neurons of the motor pathway.
• After originating from the cerebral cortex, the
corticospinal tract fibers descend as a part of
corona radiata and then pass through the
posterior limb of the internal capsule and then
downwards through the brain stem forming
pyramids in the medulla (hence the name
pyramidal tracts).
• In the lower part of medulla about 80% fibres
of each pyramid decussate in the mid line to
reach opposite side.
• The fibers of corticospinal tracts are divided into
two separate tracts:
1) Lateral corticospinal tract is constituted by 80%
of fibers which have crossed to opposite side.
• The lateral corticospinal tract fibers descend the
full length of spinal cord through the posterior
part of lateral white funiculus.
• Most of these fibers terminate in the
internuncial neurons of the spinal grey matter.
• The internuncial neurons carry the impulses to
the motor neurons situated in the ventral grey
horn.
• Some fibers of the tract terminate directly on
the ventral horn cells.
• The axons of the ventral motor neurons supply
the skeletal muscles directly by passing
through the ventral nerve root.
• The neurons giving origin to the fibers of
pyramidal tract along with their axons
constitute the upper motor neurons.
• The ventral motor neurons in the spinal cord
along with their axons constitute the lower
motor neurons.
2) Anterior corticospinal tract is formed by 20%
uncrossed pyramidal fibers.
• These fibers descend down through the
anterior white funiculus of the same side.
• The anterior corticospinal tract fibers do not
reach further than the mid-thoracic region.
• On reaching the appropriate level of the spinal
cord, the fibers of this tract cross the midline
(through the anterior white commissure) to
reach grey matter on the opposite side of the
cord and terminate in a manner similar to that
of the fibers of the lateral corticospinal tract.
• Thus, the corticospinal fibers of both the
lateral as well as the anterior tracts ultimately
connect the cerebral cortex of one side with
ventral horn cells in opposite half of the spinal
cord.
 Functions

• The cerebral cortex controls voluntary fine

skilled movements of the body through the
corticospinal tracts.
• The pyramidal tract fibres also send collaterals
to other areas of the motor control systems
thus communicating motor command to the
basal ganglia, cerebellum and the brain stem.
• In their course through the brain stem, some
of the fibers (corticonuclear fibers) terminate
directly on the motor nuclei of cranial neurons
controlling facial muscles.
• Since these fibers perform the same function
as pyramidal tracts, they are also considered
part of the pyramidal system.
 Extrapyramidal tracts
• The descending tracts of spinal cord other
than the pyramidal tracts are collectively
called extrapyramidal tracts. These include:
 Rubrospinal tract,
Vestibulospinal tract,
 Reticulospinal tract,
Tectospinal tract and
Olivospinal tract
Rubrospinal tract:
• Origin. This tract arises from the large cells
(nucleus magnocellularis) or red nucleus in the
mid brain.
• Course. After arising from the red nucleus, the
fibers of this tract cross to opposite side in the
lower part of the segmental of mid brain
(ventral segmental decussation).
• Then, the tract descends through the pons
and medulla and follows a course similar to
that of lateral corticospinal tract in the lateral
funiculus of the spinal cord.
• Termination. The fibers terminate mainly on
interneurons along with the corticospinal
fibers.
• Functions. This tract exhibits facilitatory
influence on the flexor muscles and inhibitory
influence on the extensor muscles of the body.
• The red nucleus also receives the corticorubral
fibers from the ipsilateral motor cortex.
• The corticorubro-spinal tract thus formed may
act as an alternate route of pyramidal system
to exert influence on the lower motor
neurons.
Vestibulospinal tracts:
• There are two vestibulospinal tracts:
I. lateral and
II. medial.
I. Lateral vestibulospinal tract:
• Origin. Fibers of this tract arise from the lateral
vestibular (Deiters’) nucleus.
• These fibers are somatotopically arranged.
• Fibers to cervical segments arise from the
cranioventral part, those to thoracic segments
from the central part and those to lumbosacral
segments from the dorsocaudal part of lateral
vestibular nucleus.
• Location and course. This tract is uncrossed
and lies in the anterior funiculus of the spinal
cord; shifting medially as it descends.
• Termination. The fibers extend up to caudal
segments of the cord and terminate into the
neurons of ventral grey column.
• Through the interneurons these are projected
to the alpha and gamma neurons; some fibers
directly reach the alpha neurons.
• Functions. Vestibular nucleus receives
afferents from the vestibular apparatus mainly
from utricles.
• This pathway is principally concerned with
adjustment of postural muscles to linear
acceleratory displacements of the body.
• Lateral vestibulospinal tract mainly facilitates
activity of extensor muscles and inhibits the
activity of flexor muscles in association with
the maintenance of balance.
II. Medial vestibulospinal tract:
• Origin. The fibers of this tract arise from the
medial vestibular nucleus.
• Location and course. This tract descends
through the anterior funiculus (within the
sulcomarginal fasciculus).
• The fibers are mostly uncrossed but some
fibers are crossed.
• Termination. The fibers end in the anterior
motor neurons directly or through internuncial
neurons of the cervical segments of spinal cord.
• Functions. This part of the vestibular nucleus
receives signals from the vestibular apparatus
mainly from the semicircular canals.
• Functionally, medial vestibulospinal tract is the
donor connection of medial longitudinal
fasciculus.
• This tract provides a reflex pathway for
movements of head, neck and eyes in
response to the visual and auditory stimuli.
Reticulospinal tracts:
• There are two reticulospinal tracts:
A. The medial (pontine) reticulospinal tract and
B. lateral (medullary) reticulospinal tract.
A. Medial (pontine) reticulospinal tract:
• Origin. It arises in the medial pontine reticular
formation.
• Course. The tract descends, mostly uncrossed,
in the anterior funiculus of spinal cord.
• Termination. The fibers terminate in the spinal
grey matter and through internuncial neurons
influence alpha and gamma neurons.
B. Lateral (medullary) reticulospinal tract:
• Origin. The fibers of this tract originate from the
gigantocellular component of medullary
reticular formation.
• Course. These fibers are mostly uncrossed and a
few crossed.
• This tract descends in the lateral funiculus
medial to the lateral corticospinal and
rubrospinal tracts.
• Termination. The fibers terminate in the
internuncial neurons of laminae VII, VIII and IX
of the spinal cord.
• Functions of reticulospinal tracts. The reticular
formation of the brain stem receives input
mostly from the motor cortex through the
corticoreticular fibers which accompany the
corticospinal tracts.
• Thus the corticoreticulospinal tracts form
additional polysynaptic pathways from the
motor cortex to the spinal cord.
• These tracts are concerned with control of
movements and maintenance of muscle tone.
• The reticulospinal tracts, probably, also convey
autonomic information from higher centers to
the intermediate region of spinal grey matter
and regulate respiration, circulation and
sweating.
• The pontine and medullary reticular nuclei
mostly function antagonistic to each other.
Tectospinal tract:
• Origin. Fibers of this tract arise from the
superior colliculi.
• Course. The fibers cross the midline in the
lower part of segmental of the mid brain
forming dorsal segmental decussation.
• Then the tract descends through the pons and
medulla into the anterior white funiculus of the
spinal cord.
• Termination. The fibers terminate in upper
cervical levels by synapsing on the anterior
horn cells through internuncial neurons
located in laminae V and VII of the spinal grey
matter.
• Function. This tract forms the motor limb of
the reflex pathway for turning the head and
moving the arms in response to visual, hearing
or other exteroceptive stimuli.
Olivospinal tract:
• Origin. This tract originates from the inferior
olivary nucleus.
• Course and termination. The tract fibers
descend and terminate ipsilaterally in the
anterior horn cells of the spinal cord.
• Functions. Inferior olivary nucleus receives
afferent fibers from the cerebral cortex,
corpus striatum, red nucleus and spinal cord.
• It influences muscle activity.
• Probably, it is involved in the reflex
movements arising from the proprioceptors.
Medial longitudinal fasciculus:
• Origin. The medial longitudinal fasciculus
(MLF) extends from the mid brain downwards.
• The fibers of this tract take origin from
different area of the brain stem namely:
o Vestibular nuclei,
o Reticular formation,
o Superior colliculus,
o Interstitial nucleus of Cajal and
o Nucleus of posterior commissure.
• Course. The MLF in the brain stem is closely
related to the nuclei of third, fourth, sixth and
twelfth cranial nerves.
• It is also related to the fibers of seventh nerve
(as they wind round the abducent nucleus),
and to some fibers arising from the cochlear
nuclei.
• Below, the MLF becomes continuous with the
anterior intersegmental tract of spinal cord,
which descends through the posterior part of
anterior white funiculus.
• This tract is well defined only in the upper
cervical segments.
• Below this level, the fibers run along with the
fibers of medial vestibulospinal tract.
• Termination. Along with the fibers of the
medial vestibulospinal tract, the fibers of this
tract make connections with ventral horn cells
that innervate the muscles of neck.
• Functions. MLF plays an important role in the
pathway of ocular movements.
• Its function can be summarized as:
 It ensures harmonious movements of the
eyes and neck (head) in response to vestibular
stimulation and auditory stimuli.
 It facilitates simultaneous movements of the
lips and tongue as in speech.
 CEREBELLUM IN THE
CONTROL OF MOVEMENT
• The cerebellum, or “little brain,” lies caudal to
the occipital lobe and is attached to the
posterior aspect of the brainstem through three
paired fiber tracts:
1) inferior,
2) middle, and
3) superior cerebellar peduncles.
• Input to the cerebellum comes from
peripheral sensory receptors, the brainstem,
and the cerebral cortex.
• The inferior, middle, and, to a lesser degree,
superior cerebellar peduncles carry the input.
• The output projections are mainly, if not
totally, to other motor control areas of the
CNS and are mostly carried in the superior
cerebellar peduncle.
• The cerebellum contains three pairs of
intrinsic nuclei:
I. The fastigial,
II. interposed (or interpositus), and
III. dentate.
• In some classification schemes, the interposed
nucleus is further divided into the
 Emboliform and
 Globose nuclei.
 Structural divisions of the cerebellum
correlate with function.
• The cerebellar surface is arranged in multiple,
parallel, longitudinal folds termed folia.
• Several deep fissures divide the cerebellum into
three main morphologic components:
A. anterior,
B. posterior, and
C. flocculonodular lobes, which also correspond
with the functional subdivisions of the
cerebellum.
• The functional divisions are the
vestibulocerebellum, the
spinocerebellum, and the
cerebrocerebellum.
• The lateral cerebellar hemispheres increase in
size along with expansion of the cerebral
cortex.
• The three divisions have similar intrinsic
circuitry; thus, the function of each depends
on the nature of the output nucleus to which
it projects.
• The vestibulocerebellum is composed of the
flocculonodular lobe.
• It receives input from the vestibular system and
visual areas.
• Output goes to the vestibular nuclei in the
brainstem, rather than to nuclei intrinsic to the
cerebellum.
• The vestibulocerebellum functions to control
equilibrium and eye movements.
• The medially placed spinocerebellum consists
of the midline vermis plus the medial portion
of the lateral hemispheres, called the
intermediate zones.
• Spinocerebellar pathways carrying
somatosensory information terminate in the
vermis and intermediate zones in somatotopic
arrangements.
• The auditory, visual, and vestibular systems
and sensorimotor cortex also project to this
portion of the cerebellum.
• Output from the vermis is directed to the
fastigial nuclei, which project through the
inferior cerebellar peduncle to the vestibular
nuclei and reticular formation of the pons and
medulla.
• Output from the intermediate zones goes to
the interposed nuclei and then to the red
nucleus.
• The ultimate target is the motor cortex via the
ventrolateral nucleus of the thalamus.
• It is believed that both the fastigial and
interposed nuclei contain a complete
representation of the muscles.
• The fastigial output system controls antigravity
muscles in posture and locomotion, whereas
the interposed nuclei, perhaps, act on stretch
reflexes and other somatosensory reflexes.
• The cerebrocerebellum occupies the lateral
aspects of the cerebellar hemispheres.
• Input comes exclusively from the cerebral
cortex, relayed through the middle cerebellar
peduncles of the pons.
• The cortical areas that are prominent in motor
control are the sources for most of this input.
• Output is directed to the dentate nuclei and,
ultimately, to the motor and premotor areas
of the cerebral cortex via the ventrolateral
thalamus.
 Intrinsic circuitry of the cerebellum is
regulated by Purkinje cells.
• The cerebellar cortex is composed of five
types of neurons arranged into three layers.
a) The molecular layer is the outermost and
consists mostly of axons and dendrites plus
two types of interneurons:
stellate cells and
basket cells.
b) The Purkinje cell layer contains the Purkinje
cells, whose dendrites reach upward into the
molecular layer in a fanlike array.
• The Purkinje cells are the efferent neurons of
the cerebellar cortex.
• Their action is inhibitory, with GABA being the
neurotransmitter
c) Deep to the Purkinje cells is the granular
layer, containing Golgi cells and small local
circuit neurons, the granule cells.
• The granule cells are numerous; there are
more granule cells in the cerebellum than
neurons in the entire cerebral cortex!
• Afferent axons to the cerebellar cortex are of
two types:
a. Mossy fibers and
b. Climbing fibers.
• Mossy fibers arise from the spinal cord and
brainstem neurons, including those of the
pons that receive input from the cerebral
cortex.
• Mossy fibers make complex multicontact
synapses on granule cells.
• The axons of the granule cells then ascend to
the molecular layer and bifurcate, forming the
parallel fibers.
• These travel perpendicular to and synapse
with the dendrites of Purkinje cells, providing
excitatory input via glutamate.
• Mossy fibers discharge at high tonic rates, 50
to 100 Hz, which increase further during
voluntary movement.
• When mossy fiber input is of sufficient
strength to bring a Purkinje cell to threshold, a
single action potential results.
• Climbing fibers arise from the inferior olive, a
nucleus in the medulla.
• Each climbing fiber synapses directly on the
dendrites of a Purkinje cell and exerts a strong
excitatory influence.
• One action potential in a climbing fiber
produces a burst of action potentials in the
Purkinje cell, called a complex spike.
• Climbing fibers also synapse with basket,
Golgi, and stellate interneurons, which then
make inhibitory contact with adjacent Purkinje
cells.
• This circuitry allows a climbing fiber to
produce excitation in one Purkinje cell and
inhibition in adjacent ones.
• Mossy and climbing fibers also give off
excitatory collateral axons to the deep
cerebellar nuclei before reaching the
cerebellar cortex.
• The cerebellar cortical output (Purkinje cell
efferents) is inhibitory to the cerebellar and
vestibular nuclei, but the final output of the
cerebellum by those nuclei is predominantly
excitatory.
 Cerebellar function
• Current research proposes that the
cerebellum compares the intended movement
with the actual movement and adjusts motor
system output in ongoing movements.
• The cerebellum functions in planning
movements and learning new motor skills and
also has a role in non motor cognitive
processes such as word association.
 BASAL GANGLIA
• The basal ganglia are a group of subcortical
nuclei located primarily in the base of the
forebrain, with some in the diencephalon and
upper brainstem.
• The striatum, globus pallidus (GP),
subthalamic nucleus, and substantia nigra
comprise the basal ganglia.
• Input is derived from the cerebral cortex, and
output is directed to the cortical and
brainstem areas concerned with movement.
• Basal ganglia action influences the entire
motor system and plays a role in the
preparation and execution of coordinated
movements.
• The forebrain (telencephalic) components of
the basal ganglia consist of the striatum,
which is made up of the (caudate nucleus and
the putamen), and the globus pallidus .
• The caudate nucleus and putamen are
histologically identical but are separated
anatomically by fibers of the anterior limb of
the internal capsule.
• The GP has two subdivisions: the
i. external segment (GPe), adjacent to the
medial aspect of the putamen, and the
ii. internal segment (GPi), medial to the GPe.
• The other main nuclei of the basal ganglia are
the subthalamic nucleus in the diencephalon
and the substantia nigra in th mesencephalon.
• The pathways into and out of the basal ganglia
are complex.
• Almost all areas of the cerebral cortex project
topographically onto the striatum including a
critical input from the motor cortex.
• The striatum then communicates with the
thalamus and then back to the cortex via two
different pathways.
• Indirect pathway: In the indirect pathway, the
striatum has inhibitory input to the external
segment of the globus pallidus, which has
inhibitory input to the subthalamic nuclei.
• The subthalamic nuclei project excitatory
input to the internal segment of the globus
pallidus and the pars reticulata of the
substantia nigra, which send inhibitory input
to the thalamus
• The thalamus then sends excitatory input back
to the motor cortex.
• In this pathway, the inhibitory
neurotransmitter is GABA, and the excitatory
neurotransmitter is glutamate.
• The overall output of the indirect pathway is
inhibitory.
• Direct pathway: In the direct pathway, the
striatum sends inhibitory input to the internal
segment of the globus pallidus and the pars
reticulata of the substantia nigra, which send
inhibitory input to the thalamus.
• As in the indirect pathway, the thalamus sends
excitatory input back to the motor cortex.
• Again, the inhibitory neurotransmitter is GABA,
and the excitatory neurotransmitter is
glutamate.
• The overall output of the direct pathway is
excitatory.
• The outputs of the indirect and direct
pathways from the basal ganglia to the motor
cortex are opposite and carefully balanced:
• The indirect path is inhibitory, and the direct
path is excitatory.
• A disturbance in one of the pathways will
upset this balance of motor control, with
either an increase or a decrease in motor
activity.
• Such an imbalance is characteristic of diseases
of the basal ganglia.
• In addition to the basic circuitry of the indirect
and direct pathways, there is an additional
connection, back and forth, between the
striatum and the pars compacta of the substantia
nigra.
• The neurotransmitter for the connection back to
the striatum is dopamine.
• This additional connection between the
substantia nigra and the striatum means that
dopamine will be inhibitory (via D2 receptors) in
the indirect pathway and excitatory (via D1
receptors) in the direct pathway.
• The main function of the basal ganglia is to
influence the motor cortex via pathways
through the thalamus.
• The role of the basal ganglia is to aid in
planning and execution of smooth
movements.
• The basal ganglia also contribute to affective
and cognitive functions.