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73 Cards in this Set

  • Front
  • Back
Feedback control of
motor behaviors
1. Utilize afferent information on moment to
moment basis

2. sensory signal compared with desired state (i.e., internal representation) –reference signal

3.Difference between actual and intended behavior –error signal

4. Negative/proportional feedback in closed loop
chain (e.g., thermostat)

5.Gain of feedback (ratio of input to output)
Feedforward control of
motor behaviors
Motor commands in advance to
“expected” perturbations
 Open loop control
 Feedback signals do not
contribute to second to
second variations in behavior
 Generation of an internal
model of desired state
 Still dependent on sensory
information, but a
“representation” of this
information
 raising a weight while
standing
 catching a ball
General Concepts Regarding Control of
Volitional Tasks
Motor actions have similar features regardless of conditions
(Donald Hebb 1950s)
 trajectory of reaching
 handwriting examples
General Concepts Regarding Control of
Volitional Tasks
Motor actions have similar features regardless of conditions
(Donald Hebb 1950s)
 trajectory of reaching
 handwriting examples
 Motor plan
 movement is planned by higher centers
 abstract form; not series of contractions/consequences
 Motor program - signal to specify
the intended movement given the
biomechanical constraints
Neural Basis of
Voluntary Movement
Sensorimotor cortex – skilled
movements

Basal Ganglia – initiation and
selection of motor programs

Cerebellum – coordination, timing,
learning
Sensorimotor cortex – skilled
movements
 Frontal lobe
– Primary motor cortex (M1,Area 4)
– Premotor cortex (PM, Area 6)
– Supplementary motor area (SMA,
Area 6)
 Parietal lobe
– Primary sensory cortex (S1, Area
1,2,3)
– Posterior partial cortex (Areas 5,
7)
Primary motor cortex
participates in specific
trajectory planning

delivers commands to lower
levels for initiation/
modulation of movement

Motor programs located here
or at lower levels (CPGs,
interneuron networks)
Primary somatosensory
cortex
provides sensory information
required for movement
planning and initiation

modulation of ongoing
movement
Primary cells
Layer 5 – pyramidal
cells (include large Betz cells),
distributed in specific cortical
areas
Cortical maps
(Penfield 1930-50s)
stimulated the primary
somatosensory cortex - asked the
patient about the sensation
Obtained movements with
stimulation in primary motor
cortex
The results led to a debate
about whether muscles or
movements were represented in
the cortex.

individual muscles are
represented in many locations.
Cortical control of movement
Specific cortical areas participate of specific movements/body parts -
homunculus (“little man”)

The area of cortex dedicated to processing information from a
particular body part reflects the degree of innervation of that body
part.

PM and SMA demonstrate similar topography
corticospinal tract (CST)
comprises
~ 1 million descending axons.
 At the junction of the medulla and
spinal cord, the CST divides into ventral
and lateral tracts.
 Ventral CST projects bilaterally to
motor neurons that innervate axial
muscles and to the intermediate
zone.
 Lateral CST (shown here) projects to
motor neurons of distal muscles and
interneurons in intermediate zone.
 Some axons make monosynaptic
connections onto motor neurons,
particularly those that control finger
movements.
 Other fibers synapse on interneurons
within the spinal cord.
activate cortical
neurons indirectly/non-invasively
Transcranial magnetic stimulation
(TMS) – also Transcranial
Electrical Stimulation (TES)
 Magnetic coil generates field
along cortical surface
 Faraday’s law – electrical
current generated in “wires”
(axons)
 Activate smaller cortical
cells which activate
pyramidal cells
(TES may bypass this, though
mechanisms controversial)
Multiple pathways
Multiple pathways to spinal
cord/brainstem encode
similar information
 Cortical tracts
 40% from M1 region
 S1 and PM areas contribute
significantly
 Typically to interneuron
pools – except to distal
motor pools
executing movements
 Corticospinal neurons make
direct excitatory connections
with only some motor neurons
 Direct connections for fine control
of the digits.
 After a bilateral resection of the
corticospinal tracts, monkeys unable
to grasp small objects with
thumb/index finger
Walking after bilateral resection
Walking after bilateral resection,
– unable to negotiate in feedforward
manner (obstacles, balance beam/ladder)
– Toe/ankle dorsiflexion problems, can’t
walk balance beam
discharge of action potentials
The discharge of action potentials by neurons in the primary motor
cortex (CM = corticomotoneuronal cell) depends on the motor task.
 Precision vs Power grip involve separate cortical/subcortical
circuits acting on similar IN/MN pools
Direction-specific encoding of cortical
cells
Population vectors encode both direction
and magnitude
Motor cortical cells also encode for force
Motor cortical cells alter firing
patterns according to extent of
muscle activity required to complete a
task
 Tonic and Phasic cortical cells may
participate
Premotor area
involved in goal-directed movements

Activity prior to visually-guided movements
Supplementary motor area
ensures correct sequencing of movement (order of
movement)
– biomechanical constraints
– task performed
– external conditions
 Activity prior to internally-guided movements
Posterior parietal cortex
encodes complex sensory information

internal sensory representation
Four “premotor” areas
 Lateral premotor areas
– Dorsal premotor
– Ventral premotor
 Medial premotor areas
– Supplementary motor area
– Cingulate area
Differences in neural substrates of
movement dependent on task
Alteration in Premotor (dorsal or ventral) and

Supplementary motor area during visually guided vs.
internally guided movements
 Similar in humans and non-human primates
Supplementary Motor Areas
-activated during sequential versus simple movements

-active during mental rehearsal
Ventral and Dorsal Premotor Areas
Different tasks utilize different pathways with external ques

- reaching-visual parietal dosal and PM areas
- grasping- visual parietal ventral and PM areas
Generic pathway for initiation of
volitional movements
Visual/Auditory association cortices and
Internal representation of body
and space (Posterior Parietal
Cortex)

Dorsolateral Prefrontal Cortex (DLPFC)

Premotor or Supplementary Motor Areas

Primary Motor Cortex
three
levels of control
spinal cord, brain stem, cerebral cortex
basal ganglia & cerebellum
1. Two systems of brain stem
neurons (medial and lateral
descending systems) receive input
from the cortex & subcortical
nuclei and project to the spinal
cord

2. The cerebellum and basal ganglia
provide feedback through the
thalamus (not shown) that regulate
cortical and brain stem motor
areas.
basal ganglia
involved in
motivation and initiation and
selection of motor programs.
cerebellum
is involved with
the timing and coordination of
movements (error detection) and
with the learning of motor skills.
Equilibrium
Vestibular apparatus
(inner ear, CN VIII)
-Semicircular canals –
directional rotational
accel/deceleration
-Utricle and saccule
Utricle and saccule
 Changes in rate of
linear movement

 Determining head
position in relation to
gravity
kinocilium and stereocilia
Neural signals generated in response to
mechanical deformation of hair cells
– Semicircular canals – cupula
– Utricle/saccule

Movement triggered by fluid
movement or particle movement
– Endolymph
– Otolith (Ca crystals embedded)
vestibular nuclei in brain
stem
Superior/medial from
semicircular and otolilths –
output to Medial
Longitudinal Fasciculs
(MLF)

Lateral (Dieter’s nucleus)
projects to lateral
vestibulospinal tract

Descending nucleus inputs
from otolisths – projects to
cerebellum/RF and SC
Extraocular muscles
Four are rectus muscles (straight)

Two are oblique: superior and inferior
Cranial nerve innervations:
Lateral rectus: VI (Abducens n.) – abducts eye outward

Medial, superior, inferior rectus & inf oblique: III (Oculomotor n.) –
able to look up and in if all work

Superior oblique: IV (Trochlear n.) – moves eye down and out
MLF
medial longitudinal
fasciculus
PPRF
paramedian pontine
reticular formation
Rostral interstitial nucleus –
anterior midbrain RF (MLF)
Vestibulo-ocular reflex
– keep the eyes still in space when the head
moves
Vestibulo-colic reflex
keeps the head still in space – or on a level
plane when you walk.
Vestibular-spinal reflex
adjusts posture for rapid changes in
position.
Lateral vestibular nucleus/spinal tract
•Descends the ipsilateral spinal
chord

•Terminates at all levels of the spine.

•Excitatory pathway activates
postural muscles (proximal to spine)
to correct for leftward listing
Medial vestibular nucleus/spinal tract
•Descends ipsilateral and
contralateral spinal cord but
asymmetrically

•Terminates in the cervical and
thoracic spine.

•Excitatory pathway activates
postural muscles (proximal to spine)
mainly in the neck.
Ascending projections to oculomotor nuclei
•Both direct contralateral and indirect ipsilateral control of extra ocular muscles

•Counters head movements to keep image stable on retina.

•Damage of vestibular nerve/nuclei causes nystagmus a oscillation of horizontal eye movement.
Gaze stabilization
 Vestibuloocular reflex (VOR) – stabilize eyes relative to external world
by compensating head movements

 Optokinetic nystagmus – elicted by moving object that produce illulsion or head movement (visual ocular response)
Gaze shifting
Vergence – aligns fovea of each eys with targets and different distances

Smooth pursuit – keep moving stimuli on fovea

Saccades – ballistic movements that abruptly change point of fixation
Saccades
MLF – allows
coordination of
conjugate eye
movements

PPRF –initiates fast
horizontal saccadic
movment

Rostral interstitial
nucleus – fast vertical
saccadic eye movements
Double vision
diplopia (divergent eye movement)
Misalignment:
strabismus (what is observed when shine a
light: not reflected in the same place on both eyes) – can be a cause of diplopia

 Cross eyed
 Gaze & movements not conjugate (together)
 Medial or lateral, fixed or not
 Many causes
– Weakness or paralysis of extrinsic muscle of eye
– Oculomotor nerve problem, other problems
Lazy eye
amblyopia

Cover/uncover test at 5 yo

If don’t patch good eye by 6, brain ignores lazy eye and visual

pathway degenerates: eye functionally blind
cerebellum
 Receives information about goals,
commands, and feedback signals
associated with the execution of
movement

 40X more axons project into it than
exit it (Greater number of neurons
than rest of CNS)

 Assists with spatial accuracy and
coordination of movement

 Largely motor function, some
cognitive

 Lateralization, topography of
organization

 Input and output: descending, spinal,
brainstem pathways
Vestibulocerebellum
Flocculonodular lobe (most primitive)

Receives vestibular, postural, ocular
information; controls balance and eye
movements
Spinocerebellum (Vermis)
 Intermed hemisphere and Vermis –
interposed and fastigial nuclei

proprioceptive/exteroceptive inputs

Governs “performance of movement”
Cerebro- (ponto-) cerebellum
(anterior/posterior lateral areas)
 Lateral hemisphere –Dentate nuclei

Projections from pontine nuclei

 Information of planned movement
Cerebellum cortical layers
Molecular layer
 Stellate and basket cells
 Axons of Granule cells
(parallel fibers)
 Terminals of climbing fibers
 Dendrites of Purkinje cells

Purkinje cell layer
(projection neurons to nuclei)

Granular layer
 Granule cells
 Inputs from mossy fibers
Mossy fibers
Originate from SC/BS nuclei

Carry sensory and some cortical
information

Mossy to granule (parallel
fibers) to Purkinje (generates
simple spikes)
Climbing fibers
from inferior olivary nucleus

Receives SC/cortical
information

Climbing to Purkinje
(generate complex spikes –
may depress parallel input)
Cerebellar outputs
cortical and brainstem
Role of Cerebellum
Role in Memory
 Place Cells
 Motor Learning?

Role in Error
correction
Transitions
immediate change in behavior

driven by prior experience and the ability to predict that new demands will exceed “current state” (feed-forward
strategies)
Adaptations
Gradual change in behavior that results from experience
(“feedback strategies”)

Driven by demands that exceed “current state”
Learning
Relatively permanent changes

Resulting from repeated exposure (adaptation may be a precursor)
Prism adaptation
Prisms inserted into eyeglasses
 Displace visual field
 Leads to initial errors in movement accuracy\

With extensive training, throwing with wedge prisms can become a skill.

Adaptations results in Learning – which allow faster
Transitions
Locomotor Adaptations
“Podokinetic system” - circuits
that allow directional control
of locomotor trajectory
through foot contact with the
floor (Weber et al. 1998.)
Walking on a circular treadmill
 Angular velocities (11-60º/s);
duration (7.5-60 minutes)

Stepping performed over
rotational axis or on perimeter

instructed to step in-place or walk
in a straight line on a circular
rotating treadmill
“Podokinetic” Adaptation (PKA)
 Subjects generate curved
walking trajectories with
blindfolds

Walking forward “in a straight
line”

Rotation while walking in place

Subjects unaware of trajectories
Termed...
podokinetic afterrotation (PKAR)

Recalibration of trunk and limb
position with vestibular
information

Transfers to overground backward
walking
How does PKAR occur?
Podokinetic stimulation

Overground transfer

General adaptation of
locomotor trajectory control
during stepping vs. specific
remodeling to adaptive
stimulus
Podokinetic stimulation
 Clockwise rotation, foot
turning relative to trunk

trunk must stabilize by
turning counterclockwise
Overground transfer
No PK stimulation

Negative aftereffects of
continued CCW rotation of
trunk on stable foot
Disruption of cerebellar circuits
Ataxia in humans

Surgical cooling of dentate
and interpositus nucleui

Disruption of the error
correction process?