Chapter 8: The Sensorimotor System

The sensorimotor system is organized in a hierarchical structure

o Association cortex is the highest level and sends signals down through lower levels of the
sensorimotor system
o Muscles are the lowest level
Sensory input guides motor output
Learning can change the locus of sensorimotor control from conscious to unconscious or automatic
8.1: Three Principles of Sensorimotor Function
8.2: Sensorimotor Association Cortex
Posterior Parietal Cortex: The portion of parietal neocortex posterior to the primary somatosensory
cortex.
o Plays an important role in integrating information about how the body is currently positioned
and where the object is that it will be interacting with.
o Provides spatial information to other areas of the brain
o Directs attention
o Classified as association cortex because it receives information from more than one sensory
system.
o Receives signals from the visual system, the auditory system and the somatosensory system.
o Most of the signals from the posterior parietal cortex go to areas of the motor cortex (Located in
the prefrontal cortex) and to the frontal eye field (small area of the prefrontal cortex that
controls eye movement)
o When the posterior parietal cortex is stimulated at a low level, patients report having an
intention to perform an action. High levels of stimulation are associated with the belief that one
actually performed the action. In neither case was the action performed.
o Damage to the posterior parietal cortex can result in a variety of consequences
Deficits in the perception and memory of spatial relationships
Deficits in accurate reaching and grasping
Deficits in control of eye movement
Deficits in attention
o Composed of several different areas that perform different functions
Dorsolateral Prefrontal Association Cortex: The area of the prefrontal association cortex that
plays a role in the evaluation of external stimuli and the initiation of complex voluntary motor
responses.
o Composed of several different areas that perform different functions
There is some disagreement over how to divide up the posterior parietal cortex and the dorsolateral
prefrontal association cortex.
Apraxia: Disorder of voluntary movement; Typically the problem is only evident when instructed
to perform and action;
o A disorder in which patients have great difficulty performing movements when asked to do so
out of context but can readily perform them spontaneously in natural situations.
Contralateral Neglect: Unable to respond to stimuli contralateral to the side with the lesion; Most
often observed with large lesions on the right side of the brain.
o A disturbance in the patient's ability to respond to visual, auditory or somatosensory stimuli
presented on the side of the body opposite to the site of brain damage, usually on the left side of

the body following damage to the right parietal lobe.

o Patients often behave as though the left side of their world simply doesn't exist
o Patients often don't recognize that they have a problem seeing the world as it actually is.
o Deficits are in response to stimuli presented to the left of their own bodies, referred to as
egocentric left
o Egocentric left is often determined by gravitational coordinates. When a patient tilts his or her
head, the neglected field does not also tilt.
o Patients often ignore the left half of objects, even if the object is within their visual field
o There are two lines of evidence that suggest that patients with contralateral may unconsciously
sense objects to their egocentric left, even though they don't perceive them.
When objects were repeatedly shown in the same location to the egocentric left of the
patients, they tended to look in the same spot, even though they were unaware of the
objects.
Fragmented drawings presented to the right of a patient with contralateral neglect, the
patient can more easily decipher the content of the images if complete versions were
displayed to the left of the patient.
Dorsolateral Prefrontal Association Cortex: Receives projections from the posterior parietal
cortex and sends projections to areas of the secondary motor cortex, primary motor cortex and to
the frontal eye field.
o Plays a role in the evaluation of external stimuli and the initiation of voluntary reactions to them

8.3: Secondary Motor Cortex

Secondary Motor Cortex: Areas of the secondary motor cortex are areas that receive much of their
input from association cortex (ie. Posterior parietal cortex and dorsolateral prefrontal cortex) and send
much of their output to primary motor cortex.
o Over 8 different area of the cortex are known to be secondary motor cortex.
o Supplementary Motor Area: Area of the secondary motor cortex that is within and adjacent to
the longitudinal fissure
3 regions of supplementary motor area
Supplementary Motor Area
Presupplementary Motor Area
Supplementary Eye Field
o Premotor Area: Area of the secondary motor cortex that lies between the supplementary motor
area and the lateral fissure
2 regions of premotor area
Dorsal premotor area
Ventral premotor area
o Cingulate Motor Area: Two small areas of the motor cortex of the cingulae gyrus of each
hemisphere
3 cingulate motor areas
Mirror Neurons: Neurons that fire when an individual performs a particular goal-directed hand
movement or when she or he observes the same goal-directed movement performed by another.
o Discovered in the 1990's
o Mirror neurons provide a potential mechanism for social cognition
o Being able to map other's actions onto your own actions would serve an adaptive advantage in
situations requiring understanding, imitation and cooperation.

o Mirror neurons respond to the understanding of the action, not to any superficial part of the
action itself
o Many ventral premotor neurons fire even when a monkey doesn't perceive the action being
done, but has enough clues to imagine a recreation of the behaviour
o Mirror neurons have been found in the ventral premotor cortex and the inferior portion of the
posterior parietal lobe.
o Some mirror neurons respond to the intention for behaviour rather than the behaviour itself
ie. Some posterior parietal neurons fired when a monkey grasped a piece of food with the
intention of eating it, but didn't fire if the food was being picked up and placed into a bowl
o There is very little direct evidence for the existance of mirror neurons in humans since it is
difficult to record the firing of individual neurons in humans while conducting the required
behavioural tests.
o Despite a lack of direct evidence, indirect evidence suggests that mirror neurons do, in fact,
exist in humans
fMRI have found areas of the human motor cortex that are active when a person performs,
observes or imagines a goal-directed behaviour (Rizzolatti & Fabbri-Destro, 2008;
Rodriguez et. al., 2008)
8.4: Primary Motor Cortex
Primary Motor Cortex: The cortex of the precentral gyrus, which is the major point of departure for
motor signals descending from the cerebral cortex into lower levels of the sensorimotor system.
o Major point of convergence of cortical sensorimotor signsignals
o Major point of departure for signals from cortex
o The primary motor cortex is somatotopic, meaning that it is layed out according to the muscles
of the body, with more area being devoted to regions of the body that require more dexterous
and fine motor control
Since the order of the regions controlled by each segment of primary motor cortex resemble
a person laying down across the brain, this region of the brain is sometmes described as the
'motor humunculus'
Motor Homunculus:
Each site receives sensory feedback from the receptors and joints in the areas of the body
which that site influences
An exception to this rule has been discovered in monkeys - monkeys have at least two
different areas of the primary motor cortex dedicated to the hands, with one receiving
input from the skin rather than the muscles and joints
This adaptation presumably facilitates stereognosis.
Stereognosis: The ability to distinguish objects exclusively by touch and feel.
o Stereognosis depends on a combination of muscular responses and the stimulation
produced by those responses.
o Until recently, each neuron was thought to encode the direction of movement
The evidence for this belief came from the finding that individual neurons fired maximally
when the arm reached in a different direction..
It is now believed that the primary motor cortex is the site of initiation of typical speciestypical movements.
Neurons direct movement towards a target, not necessarily in a single, pre-determined, fixed
direction.
Experiments done with stimulation of the primary motor cortex using longer pulses than

previous studies showed that each area of the motor cortex was capable of initiating
complex strings of actions that resembled recognizable behaviours such as eating.
Implications of findings on movement are:
Signals from every site in the primary motor cortex diverge greatly, so particular site has
the ability to get that body part to a target location regardless of starting position.
The sensorimotor system is inherently plastic, since we do not learn rigidly fixed
directional commands that control our bodily movements.
o Effects of Motor Cortex Lesions
Small lesions to the primary motor cortex often produce only minimal impairment of
movement.
Large lesions may disrupt the patient's ability to isolate the movement of a single body part.
(ie. Move a body part independently of other body movement)
Large lesions may also produce astereotognosia.
Astereotognosia: Deficit in stereognosis. Difficulty identifying objects by feel/touch.
Can reduce the speed, accuracy and strength of patient movement.
Lesions do not eliminate voluntary movement since there are multiple pathways leading in
and out of the primary motor cortex
8.5: Cerebellum and Basal Ganglia
Both the cerebellum and the basal ganglia are important structures in the sensorimotor system, but
neither is integrated directly into the sensorimotor hierarchy
Instead, both systems interact with many aspects of the hierarchy, coordinating their functioning to
produce smooth movement at many levels.
Cerebellum: The metencephalic structure that has been shown to mediate Pavlovian eye-blink
conditioning.
The cerebellum interacts with multiple aspects of the sensorimotor hierarchy
Helps coordinate, control and moderate movement
In the case of blindness or impaired vision, the cerebellum may allow vision-directed movement
despite the patient's inability to consciously see.
The cerebellum accounts for only 10% of the brain's mass, but over 50% of the brain's neurons
are located here.
The cerebellum receives input from four areas
Primary motor cortex
Secondary motor cortex
Brain stem
Motor nuclei
Receives feedback from motor responses via two routes
Somatosensory system (muscles and joints)
Vestibular system (Sensory system that detects changes in the direction and intensity of
head movements and that contribute to the maintenance of balance through its output to the
motor system)
Cerebellum combines input from all of these sources and adjusts movements that deviate from
the path they are meant to follow.
Involved in the timing, fine-tuning and learning of movement.
May also be involved in the timing, fine-tuning and learning of cognitive processes as well.
Diffuse damage to the cerebellum is devastating

Patient loses ability to control accuracy, strength, timing and precision of movements.
Patient loses ability to adapt movements to changing conditions once movement has been
initiated.
Patient has difficulty maintaining constant positions such as holding a glass or standing still.
Effort to do so often results in shaking/tremors.
Severe disturbances in balance, eye movements, speech and walking
Learning new motor sequences is particularly difficult for patients with this type of damage
Cognitive deficits are also associated with cerebellar damage
View is that cerebellum moderates and fine tunes cognitive processes in the same way that it
fine-tunes and moderates movement
Organized systematically in lobes, columns and layars.
Basal Ganglia: A collection of subcortical nuclei (eg. Striatum and globus pallidus) that have
important motor functions.
Contains fewer neurons than the cerebellum, but in many senses is far more complex.
The basal ganglia interacts with multiple aspects of the sensorimotor hierarchy.
Helps coordinate, control and moderate movement
In the case of blindness or impaired vision, the basal ganglia may allow vision-directed
movement despite the patient's inability to consciously see.
The basal ganglia is a heterogenous mass of interconnected nuclei
The Basal Ganglia contribute few fibers to descending motor pathways.
Part of the neural loop that receives input from the cortex and sends output back to the motor
cortex via the thalamus.
Moderates movement and cognitive functions and some of the learning processes associated
with them.
Some neurons from the basal ganglia project to areas of the cortex that are known to be
associated with cognitive functions
In lab experiments, the basal ganglia have been shown to be related to learning behaviours
associated with obtaining reward and avoiding punishment. This type of response is learned
gradually over the course of many trials.
The basal ganglia do not appear to be limited to this type of learning.

8.6: Descending Motor Pathways

There are two descending dorsal motor tracts
o Corticospinal
o Corticorubrosinal
There are two descending ventral motor tracts
o Corticospinal
o Cortico-brainstem-spinal
Both corticospinal pathways (medial and ventral) are direct motor pathways
Dorsolateral Corticospinal Tract and Dorsolateral Corticorubrospinal Tract
o Most neurons of the descending dorsal tracts synapse with neurons in the spinal gray matter
o Dorsolateral Corticospinal Tract: The motor tract that leaves the primary motor cortex,
descends to the medullary pyramids, decusates (Crosses neural pathways so that motor
pathways cross to the opposite side of the body), and then descends in the contralateral
dorsolateral spinal white matter.
1. Departs from the primary motor cortex

2. Descends to the medullary pyramid

3. Decussates
4. Descends down the dorsal white matter of the spinal cord
Controls wrist, hand, fingers and toes.
o Beltz Cells: Large pyramidal neurons of the primary motor cortex that synapse directly on
motor neurons in the lower regions of the spinal cord.
Control leg movement
o Dorsolateral Corticorubrospinal Tract: The descending motor tract that synapses in the red
nucleus of the midbrain, decussates, and descends in the dorsolateral spinal white matter.
Red Nucleus: A motor structure that is located in the tectum of the mesencephalon.
1. Departs the primary motor cortex
2. Tract synapses with neurons in the red nucleus
3. Decussates before the medulla
4. Descends down dorsolateral spinal white matter
This tract controls some facial muscles as well as distal muscles of the arms and legs
o Decussate: To cross, intersect or otherwise form the shape if an X.
Ventromedial Corticospinal Tract and Ventromedial Cortico-brainstem-spinal tract
o Ventromedial Corticospinal Tract: The direct ventromedial motor pathway, which descends
ipsilaterally from the primary motor cortex directly into the ventromedial areas of the spinal
white matter.
Axons branch diffusely and innervate interneuron circuits on both sides of many levels of
the spinal cord.
o Ventromedial Cortico-Brainstem-Spinal Tract: The indirect ventromedial motor pathway,
which descends ipsillaterally from the primary motor cortex directly into the ventromedial areas
of the spinal white matter.
Interacts with various brain stem structures
Descends on both sides of the spinal cord, carrying signals from both hemispheres.
Synapse with interneurons of multiple spinal segments to control limb and trunk movement.
o Tectum: The "roof", or dorsal surface, of the mesencephalon; it includes the superior and
inferior colliculi
o Vestibular Nucleus: The brain stem nucleus that receives information about balance from
receptors in the semicircular canals.
o Reticular Formation: A complex network of nuclei in the core of the brain stem that contains,
among other things, motor programs that regulate complex species-common movements such as
swimming and running
o Comparison of the two Dorsolateral Motor Pathways and the Two Ventromedial Motor
Pathways

Dorsolateral Tracts
Ventromedial Tracts
Made up of two major tracts of nerves Made up of two major tracts of nerves
a. Corticospinal tract
a. Corticospinal tract
b. Corticorubrospinal tract
b. Cortico-brainstem-spinal
2. One tract has axons which synapse 2. One tract has axons which synapse
directly to the spinal cord() and the
directly to the spinal cord() and the
other has axons which synapse in the
other has axons which synapse in the
brain stem and then descend into the
brain stem and then descend into the
spinal cord().
spinal cord().

3. Descending motor tracts originate in 3.

the primary motor cortex and are
assumed to mediate voluntary
movement.
4. Dorsolateral tracts are less diffuse, 4.
ending on the side of the spinal cord
segment opposite to the side of the
brain where the signal originated.
Sometimes axons will synapse
directly onto a motor neuron.
5. Neurons project to distal muscles suh 5.
as leg and arm muscles.
6. Involved in controlling isolated
movements of the arms and legs.

Descending motor tracts originste in

the primary motor cortex and are
assumed to mediate voluntary
movement.
Ventromedial tracts are much more
diffuse, with many axons innervating
interneurons on both sides of spinal
gray matter in multiple segments.

Neurons project to proximal muscles,

such as the muscles in the chest and
abdomen (trunk)
6. Involved in the maintenance and
control of posture and whole-body
movements.

8.7: Sensorimotor Spinal Circuits

Motor circuits of the spinal cord show a lot of complexity independent of cortical neuron complexity.
Muscles
o A muscle is a mass of fibers bound together in a tendon.
o Motor Units: A single motor neuron and all of the skeletal muscle fibers that are innervated by
it.
All the fibers contract when the motor neuron fires
The number of fibers per motor unit is variable
Motor units that require finer motor control (ie. Fingers) have fewer fibers per motor
neuron
Motor units that require gross motor control (ie. Thigh) have a greater number of fibers
per motor neuron.
Smallest unit of motor activity
Hundreds of thousands of thread-like membranes bound together in a tough membrane
comprise the skeletal muscle which is attached to the bone by a tendon.
o Motor End-Plate: The receptive area on a muscle fiber at a neuromuscular junction.
Motor neurons release acetylcholine at neuromuscular junctions, and causes the muscle to
contract.
Contraction is the only method muscles have of generating force.
Force generated by muscles can, there for, only ever be in one direction.
o Motor Pool: All of the motor neurons that innervate the fibers of a given muscle.
o Skeletal muscles are often considered to be one of two basic types
Fast muscle fibers are able to contract and relax quickly. These fibers are capable of
generating strong force but tire quickly because they do not have many blood vessels. This
lack of vascularization gives these muscles a pale colour.
Slow muscle fibers are not able to contract and relax quickly, but they are able to remain
contracted for longer because they contain more blood vessels. This vascularization gives
these muscles a red colour.
o Each muscle has both fast and slow muscle fibers.
o Fast muscle fibers help produce quick movements such as jumping while slow muscle fibers

help with regulated movements such as walking

o Because muscles can only exert force in a single direction, joints which can move in more then
one direction must be controlled by more then one muscle.
Flexors: Muscles that act to bend or flex a joint
Extensors: Muscles that act to straighten or extend a joint.
Synergistic Muscles: Pairs of muscles whose contraction produces a movement in the same
direction.
Any two muscles whose contraction produces the same movement
Antagonistic Muscles: Pairs of muscles that act in opposition.
Any two muscles that act in opposition to each other
Ie. Biceps and triceps
o Muscles are elastic, they are not inflexible and cable like.
o Isometric Contraction: Contraction of a muscle that increases the force of its pull but does not
shorten the muscle.
Contraction of a muscle that does not result in the two end bones becoming closer
o Dynamic Contraction: Contraction of a muscle that causes the muscle to shorten.
Contraction of a muscle that results in the two end bones becoming closer together than
before the muscle contracted.
o Muscular tension can be increased by increasing the number of neurons in the motor pool that
are firing, by increasing the firing rate of the neurons already in the motor pool or a
combination of both.
Receptor Organ of Tendons and Muscles
o Activity in the skeletal muscles is monitored by the Golgi tendon organs, which are embedded
in tendons and by muscle spindles which are embedded in the muscle.
o Golgi tendon organs and muscle spindles respond to different aspects of skeletal muscle tension
due to their differing locations.
o Golgi Tendon Organs: Receptors that are embedded in tendons and and are sensitive to the
amount of tension in the skeletal muscles to which their tendons are attached.
Respond to the amount of pull the muscle is exerting on the tendon
Unable to respond to changes in muscle length
Serve a protection function against over straining our muscles. When tension in a contracted
muscle is so extreme that it risks damage, the Golgi tendon excites inhibitory interneurons
which promote relaxation of the muscle.
o Muscle Spindles: Receptors that are embedded in skeletal muscle tissue and are sensitive to
changes in muscle length.
Respond to to changes in the length of the muscle
Unable to respond to changes in muscle tension
Intrafusal muscle within each muscle spindle innervated by its own intrafusal motor neuron
Intrafusal Muscle: A thread like muscle that adjusts the tension on a muscle spindle
Intrafusal Motor Neuron: A motor neuron that innervates an intrafusal muscle.
Without intrafusal motor input, a muscle spindle would fall slack each time its skeletal
muscle contracted. If the intrafusal neuron were slack, it would be unable to do its job
responding to changes in muscle length.
The intrafusal muscle solves this problem by becoming shorter along with the skeletal
muscle and remaining taught enough to respond to slight changes in length.

o Skeletal Muscle (Extrafusal Motor Neuron): Striated muscle that is attached to the skeleton
and is usually under voluntary control.
Stretch Reflex
o Patellar Tendon Reflex: The stretch reflex that is elicited when the patelar tendon is struck.
Monosynaptic
Patella means knee, and the patellar tendon reflex refers specifically to the reflex which is
tested by striking below the knee with a rubber-headed mallet.
o Stretch Reflex: A reflective counteracting reaction to an unanticipated external stretching force
on a muscle
Monosynaptic
Serves to maintain limb stability
The sudden stretch of the muscle stretches its muscle spindle
The muscle spindle stretching causes the spindle receptors to also stretch
Spindke receptors initiate action potentials which carry excitatory signals to the ventral horn
of the spinal cord
The ventral horn of the spinal cord sends the action potentials back to the muscle whose
stretch originally caused the excitement.
The arrival of the potential back at the starting point causes the muscle to contract in
reaction to the initial stretch.
The stretch reflex is designed to protect the body against its position being altered
unintentionally by an external force.
When an external force is applied to the body, the stretch reflex compensates so that the
position of the body does not change much in response to force applied.
o Spindle Afferent Neurons: Neurons that carry signals from muscle spindles into the spinal
cord via the dorsal root.
Withdrawal Reflex
o Withdrawal Reflex: The reflexive withdrawal of a limb when it comes into contact with a
painful stimuli.
Not monosynaptic
When a painful stimulus is experienced, the initial reflex is to pull away
First response is recorded in the motor neurons of the flexor muscles, which takes
approximately 1.6 Ms
The shortest route in the withdrawal reflex circuit involves one interneuron.
Reciprocal Innervation
o Reciprocal Innervation: The principal of spinal cord circuitry that causes a muscle to
automatically relax when a muscle that is antagonistic to it contracts.
Antagonistic muscles interact so that movements are smooth
Ie. Flexors are inhibitted when extensors are excited, etc...
In the withdrawal reflex, when the painful signal reaches the dorsal horn of the spinal cord
(ie. From a finger), both excitatory and inhibitory interneurons fire so that one muscle
contracts and its antagonist muscle relaxes.
Motion of many muscles are coordinated by the spinal cord
Movement is quickest when there is simultaneous excitation of all agonists and complete
inhibition of all antagonists, but most movements are not produced that way
Most muscles are always contracted to some degree and movements are produced by

adjusting the level of relative contraction between agonists and antagonists

o Cocontraction: The simultaneous contraction of two antagonistic muscles.
Movement produced by cocontraction is smooth and can be stopped at any time by a slight
adjustment in contraction of the antagonistic muscles
Cocontraction insulates us from the effects of unexpected external forces.
Recurrent Collateral Inhibition
o Recurrent Collateral Inhibition: The inhibition of a neuron that is produced by its own
activity via a collateral branch of its axon and an inhibitory interneuron.
After a muscle has contracted, a feedback loop occurs through Renshaw cells which allows
the muscle fibers to rest.
Each motor neuron has a small branch that departs from the main axon before exiting the
spinal cord.
This branch synapses with an interneuron that inhibits the firing of that neuron
The small interneuron that mediates the firing-based inhibition are called Renshaw cells.
Because of the process of recurrent collateral inhibition, every time a motor neuron fires, it
also briefly inhibits further firing and delegates some of the muscular responsability for the
contraction to other motor neurins in the motor pool.
Walking: A Complex Sensorimotor Reflex
o Walking is a complex reflex in some species
o Walking involves integrating information from the vestibular system, visual system, auditory
system and somatosensory system.
o Walking involves coordinating movement in the arms, legs, trunk, feet etc...
o Movement of walking must also be adaptable to changes in terrain, commands from the brain,
external forces etc...
o In some animals, walking is partially controlled by the spinal cord.
in cats whose brains are not connected to the spinal cord, suspending them over a treadmill
that simulates the responses their spine would get if the cat were walking and actually
produces walking in the cat.
In humans, the descending motor pathways seem to play a greater role in walking than in
other animals.
8.8: Central Sensorimotor Programs.
All but the highest levels of the sensorimotor system have complex movement pathways 'wired or
'programmed' into them.
Higher levels of the sensorimotor system activate lower levels to produce these complex movements.
Basal ganglia and cerebellum serve to coordinate the series of movements that has already been
learned.
Higher levels of the sensorimotor system retain the ability to intervene and change motion, but most of
the individual movements performed are done without consciously controlling each one.
Central Sensorimotor Programs: Patterns of activity that are programmed into the sensorimotor
system.
Central Sensorimotor Programs are Capable of Motor Equivalence
o Movements can be accomplished in different ways, using differing muscles
o Since different muscles can accomplish the same task, central sensorimotor programs must be
stored at a higher level than the muscles themselves
o Some sensorimotor programs are stored by the secondary motor cortex
o Motor Equivalence: The ability of the sensorimotor system to carry out the same basic

movement in a variety of ways using different muscles.

Motor equivalence illustrates the plastic properties of the brain
The fact that you can sign your name in pen with your hand and on a beach with your toe,
yet the name still looks similar suggests that the central sensorimotor program for signatures
is not stored in either your hand or your foot, but rather somewhere higher than both which
can be adapted to use many different muscle groups in many different situations.
Sensory Information that Controls Central Sensorimotor Programs is not Necessarily Conscious
o There is evidence that patients respond to visual stimuli of which they have no conscious
awareness
o There is also evidence that patients had trouble interacting with objects they did perceive
consciously
o Ebbinghaus Illusion: Two disks of equivalent size are placed on a table, one surrounded in a
circle by smaller disks and the other surrounded in a circle by larger disks. Though the brain
consciously perceives the two central disks as different sizes, when the hand reaches out to grab
the central disk, the finger and thumb open to the correct width, regardless if conscious
perception.
The Ebbinghaus illusion demonstrated that conscious perception and visually guided
movement are processed through different streams.
o Central sensorimotor programs are capable of using feedback without engaging or alerting
higher brain centers
o Central sensorimotor programs are organized into a hierarchy, with each level receiving
feedback information from lower levels only as necessary.
Central Sensorimotor Programs can Develop Without Practise
o Programs for many species-typical behaviours are developed in those animals with no practise
o Fentress study - showed that mice raised without forelimbs still produced cleaning behaviours
that were coordinated between shoulder, eye and tongue.
The study also showed the importance of sensory feedback since mice that were unable to
feel their limb against their tongue would take extensive cleaning breaks to lick other mice
or even the floor.
Practise can Create Sensorimotor Programs
o Practising a behaviour or movement can generate new programs or modify existing
sensorimotor programs
o Response Chunking
Response Chunking Hypothesis: The idea that practise combines the central sensorimotor
programs that control individual responses into programs that control sequences of
responses.
An important aspect of chunking is that lower order chunks (sequences which have already
been formed from individual actions) can be combined into a higher order chunk, ie. a
sequence of sequences.
o Shifting Control to the Lower Levels
When a central sensorimotor program is learned, control is shifted from the higher levels of
the hierarchy to lower levels.
This shift from higher to lower control has two advantages
Transfer of control to the lower levels frees up resources in the higher levels to focus on
details and adapting to the situation.
o An example of this is a dancer who has learned the steps to the dance very well and
so can focus on the interpretation of the piece instead of the individual movements

of the body.
Other advantage of this transfer is efficiency and increased speed
o Components of the lower levels are able to act simultaneously without interfering
with each other or waiting for instructions from higher levels
Functional Brain Imaging of Sensorimotor Learning
o FMRI studies in humans have largely confirmed what has been revealed about our brains
through the use of invasive procedures on animals
o Tasks that are being learned create a high level of activity in the primary motor and
somatosensory cortices, premotor cortex, posterior parietal cortex, dorsolateral prefrontal cortex
and cerebellum. Tasks being learned create an average level of activity in the supplementary
motor cortex.
o Tasks that are well-learned produce a high level of activity in the supplementary motor area, as
well as primary and secondary motor cortices. Well-learned tasks also produce average levels of
activity in the premotor cortex, posterior parietal cortex and the cerebellum.
o Posterior parietal cortex was active during both learning a new sequence and performing a well
known sequence, but it was more active during the learning phase.
Supports the hypothesis that the posterior parietal cortex is responsible for integrating
sensory information into the control of movement.
Hearing the tones is important, but more so during the learning phase when the participant
must actively attend to the tones in order to know if he or she performed the action
correctly.
o Dorsolateral prefrontal cortex was activated during the performance of the newly learned
sequence but not the well learned one.
Supports the hypothesis that the dorsolateral prefrontal cortex is active largely when the
action is guided under conscious control, which is typically the case with early stage motor
learning.
o Areas of the secondary motor cortex responded differently to the newly learned and well
learned conditions.
Contralateral premotor cortex was more active when participants performed the newly
learned sequence
Supplementary motor area was more active during the performance of the well-learned
sequence of digits.
Supports hypothesis that premotor cortex plays a more dominant role when an action is
being guided by sensory feedback, as is the case in early stage motor learning.
Supplementary motor area plays a more prominent role when an action is performed
without much need for sensory guidance, which is the case for well-practiced motor
sequences.
o Contralateral primary motor and somatsensory cortices were equally active during the
performance of both well learned and newly learned tasks.
Consistent with the research design that both groups would be performing a variation of the
same task. (Finger pressing variations)
o Contralateral basal ganglia was equally active during the performance of both well learned and
newly learned tasks
o Both sides of the cerebellum were activated during both well learned and newly learned actions,
but was more active when participants were performing newly learned sequences
Supports the hypothesis that the cerebellum plays a prominent role in motor learning