Chapter 8: THE SENSORIMOTOR SYSTEM

Three Principles of Sensorimotor Function

The Sensorimotor System is Hierarchically Organized
• Advantage: higher levels of the hierarchy are left free to perform more complex functions
• Parallel hierarchical systems: signals flow between levels over multiple paths
• Functional segregation: each level of the sensorimotor and company hierarchies tends to be composed of different units
(neural structures or departments), each of which performs a different function
• Information mainly flows down
Motor Output is Guided by Sensory Input
• Sensory feedback: plays an important role in directing the continuation of the responses that produced it
o Ballistic movements: brief, all-or-none, high-speed movements (e.g. swatting a fly); not influenced by sensory feedback
• Many adjustments in motor output that occur in response to sensory feedback are controlled unconsciously by the lower
levels of the sensorimotor hierarchy with- out the involvement of the higher levels
Learning Changes the Nature and Locus of Sensorimotor Control
• Initial stages of motor learning: each individual response is performed under much
conscious control
• After much practice, individual responses become organized into continuous
integrated sequences of action that low smoothly and are adjusted by sensory
feedback without conscious regulation
• Transfer of control to lower levels of the CNS characterize most sensorimotor
learning
General Model of Sensorimotor System Function
• Neural structures that play important roles in the control of voluntary control
o Begins at the level of association cortex à traces major motor signals as they
descend the sensorimotor hierarchy to the skeletal muscles that ultimately
perform the movements

Sensorimotor Association Cortex

Posterior Parietal Association Cortex
• Portion of parietal neocortex posterior to the primary somatosensory cortex
• Plays an important role in integrating these two kinds of information, in directing
behavior by providing spatial information, and in directing attention
• Receives information from 3 sensory systems that play roles in the localization of
the body and external objects in space: visual system, auditory system, and
somatosensory system
• Much of the output goes to areas of motor cortex (in the frontal cortex): to the
dorsolateral prefrontal association cortex, to various areas of secondary motor
cortex, and to the frontal eye field (in the prefrontal cortex; controls eye
movements)
• Desmurget et al. (2009): applied electrical stimulation to the inferior portions of the posterior parietal cortexes of
conscious neurosurgical patients
o Low current levels: experienced an intention to perform particular action
o High levels: felt that they had actually performed it (action didn’t actually occur)
• Damage to posterior parietal cortex can produce variety of deficits (e.g. in the perception and memory of spatial
relationships, in accurate reaching and grasping); most striking: apraxia and contralateral neglect
• Apraxia: disorder of voluntary movement that is not attributable to a simple motor deficit (e.g., not to paralysis or
weakness) or to any deficit in comprehension or motivation
o Have difficulty making specific movements when they are requested to do so, particularly when the movements are out
of context; can often readily perform the very same movements under natual conditions when they are not thinking
about what they are doing
o Caused by unilateral damage to the left posterior parietal cortex or its connections
• Contralateral neglect: disturbance of a patient’s ability to respond to stimuli on the side of the body opposite to the side of
a brain lesion in the absence of simple sensory or motor deficits
 o Behave as if the left side of their world does not exist, and they often fail to appreciate that they have a problem
(egocentric left)
o Object0based contralateral neglect: failure to respond to the left side of objects even when the objects are presented
horizontally or upside down
o Disturbance is associated with large lesions of the right posterior parietal cortex
o 2 types of evidence suggest that information about objects that are not noticed by patients with contralateral neglect
may be unconsciously perceived
§ When objects were repeatedly presented at the same spot to the left of the patients, they tended to look to the
same spot on future trials, although they were unaware of the objects
§ Patients could more readily identify fragmented (partial) drawings viewed to their right if complete versions of the
drawings had previously been presented to the left, where they were not consciously perceived
Dorsolateral Prefrontal Association Cortex
• Receives projections from the posterior parietal cortex; sends projections to areas
of secondary motor cortex, to primary motor cortex, and to the frontal eye field
• Activity of neurons depend on the characteristics of objects, locations of objects,
or a combination of both
• Activity of some neurons is related to the response rather than to the object
o Begin to fire before the response and continue to fire until the response is
complete
• Decisions to initiate voluntary movements may be made in this area of cortex, but
these decisions depend on critical interactions with posterior parietal cortex and
other areas of frontal cortex

Secondary Motor Cortex

• Receive much of their input from association cortex and send much of their output to primary motor cortex
• Supplementary motor area: wraps over the top of the frontal lobe and extends down its medial surface into the
longitudinal fissure
• Premotor cortex: runs in a strip from the supplementary motor area to the lateral fissure
• Both are clearly visible on the lateral surface of the frontal lobe, anterior to the primary motor cortex
Identifying the Areas of Secondary Motor Cortex
• 8 areas of secondary motor cortex in each hemisphere:
o 3 supplementary motor areas (SMA, preSMA, and supplementary eye field
o 2 premotor areas (dorsal and ventral)
o 3 cingulate motor areas (in the cortex of the cingulate gyrus)
• Involved in the programming of specific patterns of movements after taking
general instructions from dorsolateral prefrontal cortex
Mirror Neurons
• Neurons that fire when an individual performs a particular goal-directed hand
movement or when they observe the same goal-directed movement performed
by another
• Discovered in the early 1990s by Giacomo Rizzolatti
o Studying macaque monkey ventral premotor neurons
• Provide a possible mechanism for social cognition (knowledge of the perceptions, ideas, and intentions of others); neurons
respond to the understanding of the purpose of an action
o Mapping the actions of others onto one’s own action repertoire would facilitate social understanding, cooperation, and
imitation
• Found in several areas of the macaque monkey frontal and parietal cortex

Primary Motor Cortex

• Located in the precentral gyrus of the frontal lobe
• Major point of convergence of cortical sensorimotor signals; major point of departure of sensorimotor signals from the
cerebral cortex
Conventional View of Primary Motor Cortex Function
• 1937: Penfield and Boldrey mapped the primary motor cortex of conscious human patients during neurosurgery; applied
brief, low-intensity electrical stimulations to various points on the cortical surface and
noting which part of the body moved in response to each stimulation
o Found that stimulation of each particular cortical site activated a particular
contralateral muscle and produced a simple movement
• Primary motor cortex is organized somatotopically (according to a map of the body)
o Motor homunculus: somatic map of the human primary motor cortex
• Each site in the primary motor cortex receives sensory feedback from recep- tors in
the muscles and joints that the site influences
o Exception: general pattern of feedback in monkeys; have at least 2 different hand
areas in the primary motor cortex of each hemisphere, and one receives input
from receptors in the skin rather than from receptors in the muscles and joints
• Stereognosis: process of identifying objects by touch; depends on a complex interplay between
motor responses and somatosensory stimulation produced by them
• Function of primary motor neurons: encode the direction of movement; each has a different
preferred location
Current View of Primary Motor Cortex Function
• Investigators used longer bursts of current (e.g. 0.5 to 1 second) to map the primary motor cortex
o Elicited complex natural-looking response sequences
• Crude somatotopic organization: stimulation in the face area tended to elicit face movements
o Elicited responses were complex species-typical movements that involved several body parts
• Sites that moved a particular body part overlapped greatly with sites that moved other body parts
• Motor neurons are coded to particular angles (end point) of movement (vs. movement to a particular direction)
• Same stimulation of motor cortex can produce opposite movements depending on the starting position, but the end
position of the movements remains the same
• Implications of findings:
o Signals from every site in the primary motor cortex diverge greatly, so each particu- lar site has the ability to get a body
part (e.g., an arm) to a target location regardless of the starting position
o Sensorimotor system is inherently plastic
o Primary motor complex contains an action map
Effects of primary motor cortex lesions
• Large lesions to the primary motor cortex may disrupt a patient’s ability to move one body part (e.g., one finger)
independently of others, may produce astereognosia (deficits in stereognosis), and may reduce the speed, accuracy, and
force of a patient’s movements
o These lesions do not eliminate voluntary movement; due to the parallel pathways that descend directly from secondary
and association motor areas to subcortical motor circuits without passing through primary motor cortex

Cerebellum and Basal Ganglia

• Neither is a major part of the pathway by which signals descend through the sensorimotor hierarchy
• Both interact with different levels of the sensorimotor hierarchy and, in so doing, coordinate and modulate its activities
Cerebellum
• Constitutes only 10% of the mass of the brain; contains more than half of the brain’s neurons
• Receives information from primary and secondary motor cortex, information about descending motor signals from brain-
stem motor nuclei, and feed- back from motor responses via the somatosensory and vestibular systems
o Compares these 3 sources of input and correct ongoing movements that deviate from their intended course
• Plays a major role in motor learning, esp. in the learning of sequences of movements in which timing is a critical factor
• Damage à lose ability to control precisely the direction, force, velocity, and amplitude of movements and the ability to
adapt patterns of motor output to changing conditions; disturbances in posture, balance, gait, speech, control of eye
movement, learning new motor sequences
o Diverse sensory, cognitive, and emotional deficits
Basal Ganglia
• Complex heterogeneous collection of interconnected nuclei
• Contribute few fibers to descending motor pathways; instead, they are part of neural loops that receive cortical input from
various cortical areas and transmit it back to the cortex via the thalamus (loops carry signals to and from the motor areas
of the cortex)
• Involved in a variety of cognitive functions
• Participate in habit learning (type of motor learning that is usually acquired gradually, trial by trial)

Descending Motor Pathways

Dorsolateral Corticospinal Tract and Dorsolateral Corticorubrospinal Tract
• Dorsolateral corticospinal tract (direct)
o Group of axons descends from the primary motor cortex through the medullary pyramids (2 bulges on the ventral
surface of the medulla) à decussates à descend in the contralateral dorsolateral spinal white matter
o Betz cells: extremely large pyramidal neurons of the primary motor cortex
o Most of the neurons synapse on small interneurons of the spinal gray matter, which synapse on the motor neurons of
distal muscles of the wrist, hands, fingers, and toes
• Dorsolateral coritorubrospinal tract (indirect)
o Descends from from the primary motor cortex synapses in the red nucleus of the midbrain à decussate and descend
through the medulla, where some terminate in the nuclei of the cranial nerves that control the muscles of the face, or;
descend in the dorsolateral portion of the spinal cord
o Synapse on the interneurons that in turn synapse on motor neurons that project to the distal muscles of the arms and
legs
Ventromedial Corticospinal Tract and Ventromedial Corticobrainstem-spinal Tract
• Ventromedial corticospinal tract (direct)
o Axons descend ipsilaterally from the primary motor cortex à ventromedial areas of the spinal white matter à
branches diffusely and innervates the interneuron circuits in several different spinal segments on both sides of the
spinal gray matter
• Ventromedial corticobrainstem-spinal tract (indirect)
o Comprises of motor cortex neurons that feed into a complex network of brain stem structures
o Axons descend bilaterally in the ventromedial portion of the spinal cord
o Each side arries signals from both hemispheres, and each neuron synapses on the interneurons of several different
spinal cord segments that control the proximal muscles of the trunk and limbs
• 4 brain structures that interact with the ventromedial corticobrainstem-spinal tract
o Tectum: receives auditory and visual information about spatial location
o Vestibular nucleus: receives information about balance from receptors in the semicircular canals of the inner ear
o Reticular formation: contains motor programs that regulate complex species-typical movements (e.g. walking)
o Motor nuclei of the cranial nerves: control the muscles of the face
Comparison of the 2 Dorsolateral Motor Pathways and the 2 Ventromedial Motor Pathways
• Each is composed of two major tracts, one whose axons descend directly to the spinal cord and another whose axons
synapse in the brain stem on neurons that in turn descend to the spinal cord
• 2 ventromedial tracts are much more diffuse
o Axons innervate interneurons on both sides of the spinal gray matter and in several different segments
o Dorsolateral: terminate in the contralateral half of one spinal cord segment, or directly on a motor neuron
• Motor neurons activated by 2 ventromedial tracts project to proximal muscles of the trunk and limbs (e.g., shoulder
muscles); motor neurons activated by the two dorsolateral tracts project to distal muscles (e.g., finger muscles)
• 2 ventromedial tracts are involved in the control of posture and whole-body movements (e.g., walking and climbing) and
that they can exert control over the limb movements involved in such activities
• Dorsolateral tracts control the movements of the limbs
o Only the corticospinal division is capable of mediating independent movements of the digit

Sensorimotor Spinal Circuits

Muscles
• Motor units: smallest units of motor activity
o Each motor unit comprises a single motor neuron and all of the individual skeletal muscle fibers that it innervates
o Motor neuron fires à muscle fibers of the unit contract together (to generate force)
o Units with the fewest fibers permit the highest degree of selective motor control
• A skeletal muscle comprises hundreds of thousands of threadlike muscle fibers bound together in a tough membrane and
attached to a bone by a tendon
• Acetylcholine (released by motor neurons at the neuromuscular junction) activates the motor end-plate on each muscle
fiber and causes the fiber to contract
• Motor pool; all motor neurons that innervate the fibers of a single muscle
• Fast muscle fibers: contract and relax quickly; poorly vascularized (have few blood vessels)
• Slow muscle fibers: slower and weaker; capable of more sustained contraction because they are more richly vascularized
• Flexors: act to bend or flex a joint; elbow joint flexion: biceps contract, triceps extend
• Extensors: act to straighten or extend it; elbow joint extension: biceps extend, triceps contract
• Synergistic: two muscles whose contraction produces the same movement
• Antagonistic muscles
• Isometric contraction: activation of a muscle can increase the tension that it exerts on
two bones without shortening and pulling them together
• Dynamic contraction: can shorten and pull them together
• Tension in a muscle can be increased by increasing the number of neurons in its motor
pool that are firing, by increasing the firing rates of those already firing, or more
commonly by a combination of these two changes
Receptor Organs of Tendons and Muscles
• Activity of skeletal muscles is monitored by:
o Golgi tendon organs: embedded in the tendons (connect each skeletal muscle to
bone); respond to increases in muscle tendon (i.e. to the pull of the muscle on the
tendon); insensitive to changes in muscle length; provide CNS with information
about muscle tension
§ When the contraction of a muscle is so extreme that there is a risk of damage,
Golgi tendon organs excite inhibitory interneurons in the spinal cord that cause
the muscle to relax
o Muscle spindles: embedded in the muscle tissue itself; respond to changes in muscle
length; do not respond to changes in muscle tension
• Muscle-spindle feedback circuit
o Each muscle spindle has its own threadlike intrafusal muscle, which is innervated by
its own intrafusal motor neuron
§ Without its intrafusal motor input, a muscle spindle would fall slack each time its
skeletal muscle (extrafusal muscle) contracted à muscle spindle can’t respond
to changes in extrafusal muscle length
§ Intrafusal motor neuron shortens the intrafusal muscle each time the extrafusal
muscle becomes shorter, thus keeping enough tension on the middle, stretch-
sensitive portion of the muscle spindle to keep it responsive to slight changes in
the length of the extrafusal muscle
Stretch Reflex
• Reflex elicited by a sudden external stretching force on a muscle
• Patellar tendon reflex: kind of a leg extension
o By a sharp blow to the tendon of completely relaxed muscle à to make reflex
readily observable
• Sudden stretch of the thigh muscle stretches its muscle-spindle muscle-spindle stretch
receptors à initiate action potentials carried from the stretch receptors into the spinal
cord by spindle afferent neurons via the dorsal root à excites motor neurons in the
ventral horn of the spinal cord à sends action potensials back to the muscle whose
stretch originally excited them à compensatory muscle contraction and a sudden leg
extension
• Function: keep external forces from altering the intended position of the body (e.g.
produces immediate compensatory contraction of the muscle that counteracts the
force to keep you from spilling coffee)
• How stretch reflex maintains limb stability
o Shows the important role played by sensory feedback in the regulation of motor
output
o Shows the ability of lower circuits in the motor hierarchy to take care of “business
details” without the involvement of higher levels
Withdrawal Reflex
• Not somosynpatic
• When a painful stimulus is applied to the hand, the first responses are recorded in the motor neurons of the arm flexor
muscles about 1.6 milliseconds later, about the time it takes a neural signal to cross two synapses
• Shortest route in the withdrawal-reflex circuit involves one interneuron
• Other responses are recorded in the motor neurons of the arm flexor muscles after the initiall volley
o Triggered by signals that have traveled over multisynaptic pathways
Reciprocal Innervation
• Important principle of spinal cord circuity
• Antagonistic muscles are innervated in a way that permits a smooth, unimpeded motor response: When one is contracted,
the other relaxes
• News of a sudden painful event in the hand arrives in the dorsal horn of the spinal cord and has two effects:
o Signals excite both excitatory and inhibitory neurons
§ Excitatory interneurons à excite motor neurons of elbow flexor
§ Inhibitory neurons à inhibit motor neurons of the elbow extensor
• Activities of agonists and antagonists are automatically coordinated by the internal circuitry of the spinal cord
• Movements are quickest when there is simultaneous excitation of all agonists and complete inhibition of all antagonists
• Most muscles are always contracted to some degree, and movements are produced by adjustment in the level of relative
cocontraction between antagonists
• Movements produced by cocontraction are smooth, and they can be stopped with precision by a slight increase in the
contraction of the antagonistic muscles
o Cocontraction insulates us from the effects of unexpected external forces
Recurrent Collateral Inhibition
• Muscle fibers and motor neurons that innervate them need a break
(managed by inhibitory neurons in the spinal cord)
• Each motor neuron branches just before it leaves the spinal cord, and the
branch synapses on a small inhibitory interneuron, which inhibits the very
motor neuron from which it receives its input
• Renshaw cells: small inhibitory interneurons that mediate recurrent
collateral inhibition
• Each time a motor neuron fires, it momentarily inhibits itself and shifts the
responsibility for the contraction of a particular muscle to other members
of the muscle’s motor pool
Walking: A Complex Sensorimotor Reflex
• Must integrate different kinds of information from different systems à
must produce an integrated series of movements that involves the muscles
of the trunk, legs, feet, and upper arms
• Must be plastic: able to adjust its output immediately to changes in the
slope of the terrain, to instructions from the brain, or to sudden external forces
• Spinal cord can control walking

Central Sensorimotor Programs and Learning

A Hierarchy of Central Sensorimotor Programs
• All but the highest levels of the sensorimotor system have certain patterns of activity programmed into them, and complex
movements are produced by activating the appropriate combinations of these programs
• Once activated, each level of the sensorimotor system is capable of operating on the basis of current sensory feedback
without the direct control of higher levels
o Most of the individual responses that you make are performed without direct cortical involvement, and you are often
barely aware of them
Characteristics of Central Sensorimotor Programs
Central sensorimotor programs are capable to motor equivalence
• Some basic movements can be carried out in different ways involving different muscles (e.g. sign name with fingers; write
name with toe on sand)
• Illustrated inherent plasticity of the sensorimotor system; specific central SM programs are not stored in neural circuits,
general programs are stored higher in your sensorimotor hierarchy and then are adapted to the situation as required
Sensory information that controls central sensorimotor programs is not necessarily conscious
• E.g. neural mechanisms of conscious visual perception (ventral stream) are not necessarily the same as those that mediate
the visual control of behavior (dorsal stream)
• Evidence for the separation of conscious perception and sensory control: Haffendedn & Goodal (1998) used the
Ebbinghaus illusion
o When participants were asked to indicate the size of each central disk with their right thumb and pointing finger, they
judged the disk on the left to be bigger than the one on the right
o When asked to reach out and pick up the disks with the same two digits, the preparatory gap between the digits was a
function of the actual size of each disk rather than its perceived size
Central sensorimotor programs can develop without practice
• Fentress (1973): adult mice raised from birth without fore- limbs still made the patterns of shoulder movements typical of
grooming in their species—and that these movements were well coordinated with normal tongue, head, and eye
movements
Practice can create central sensorimotor programs
• 2 kinds of processes that influence the learning of central sensorimotor programs: response chunking and shifting control
to lower levels of the sensorimotor system
• Response chunking: practice combines the central sensorimotor programs that control individual responses into programs
that control sequences (chunks) of behavior
o Chunks can themselves be combined into higher-order chunks; these chunks can be combined so that it can be typed as
a unit
• Shifting control to lower levels has 2 advantages
o Frees up the higher levels of the system to deal with more esoteric aspects of performance
o Permits great speed because different circuits at the lower levels of the hierarchy can act simultaneously, without
interfering with one another
Functional Brain Imaging of Sensorimotor Learning
• PET study of Jenkins et al (1994): recorded PET activity from human volunteers who performed two different sequences of
key presses
o Presses were performed with the right hand, one every 3 seconds, and tones indicated when to press and whether or
not a press was correct
o Results: involvement of the cortical sensorimotor areas; involvement of association areas and the cerebellum
diminished when sequences were well practiced