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physiology section 2 lecture 4
physiology section 2 lecture 4
95
Biology
Graduate
09/25/2013

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Term
what does a golgi tendon organ sense?
Definition
senses force from muscle firing. ues Ib afferent neurons
Term
The Ib axon (afferent) from
the golgi tendon organ
contacts an interneuron in
the spinal cord; the
interneuron is known as the
Ib inhibitory interneuron. Why?
Definition
to prevent the muscle from ripping off the bone (sort of-it can be modulated)
Term
what same input to your foot at rest can cause two different responses from the Ib golgi afferent neuron. what is this and why does this happen?
Definition
at rest, you get an inhibitory hyperpolarization, but during walking, that same input causes a depolarization. you can switch from inhibitory to excitatory. may not be hard wired
Term
what are some mechanical proprioceptive pathways? what type of fibers do they use?
Definition
(group II, III)
 Plantar skin during stance
phase of walking
 Tactile inputs on fingertips
during grasp - if anesthetized, you
Term
what are some noxious proprioceptive pathways, and what type of fibers do they use?
Definition
(group III-IV)
 flexion withdrawal reflex - stimulates flexion of one hip, but extension of the other to keep you upright (stepping on a tac)
 local sign - pulls away from the noxious stimulus (anywhere else)
Term
stumbling corrective response
Definition
in cats: A mild mechanical stimulus
applied to the dorsal part of the paw during the swing phase produces excitation of flexor motor neurons and inhibition of extensor motor neurons, leading to rapid
flexion of the paw away from the stimulus and elevation of the leg in an attempt to step over the object. Because this corrective response is readily observed in
spinal cats, it must be produced to a large extent by circuits entirely contained within the spinal cord.
Term
which ones are hard wired, and which aren't?
Definition
flexor withdrawal appears to be hard wired, but local sign and stumbling corrective response appear to be learned.
Term
Reflexes modulate and even reverse action in different
conditions
Can not be explained by simple
habituation
Network of cells
Definition
Term
Decerebrate
transection at the brainstem (midbrain) (a-a’)
Definition
all you have is brain stem and spinal cord
Term
pinal
transection at lower thoracic level (b-b’)
Definition
just lumbo sacral reflexes (no higher spinal cord, brain, or brainstem
Term
deafferented
transection of the dorsal roots
Definition
no afferent inputs, no reflexes
Term
Immobilized
removes movement related feedback by paralyzing the muscles
(fictive locomotion)
Definition
you can record from the nerves, but no muscle contraction. removes muscle feedback.
Term
Neonatal (and/or) in vitro preparation.
Definition
good way to compare eNg to eMg.
Term
what is the definition of a central pattern generator?
Definition
circuit with intrinsic
ability to generate rhythmic,
coordinated motor behaviors
Term
describe Graham Brown’s half-center model (1911)
Definition
comprises one set of
neurons that project to motor neurons innervating extensor
muscles and another set that projects to motor neurons
innervating flexor muscles.
 Two centers inhibit each other reciprocally so that when one
half-center is active, the other is inhibited.
Term
What is the most studied/detailed Central Pattern
Generator?
Definition
lobster stomatogastric network.Total of 33 neurons control all feeding behaviors
 Approximately 50% of
all possible connections
are present
 Anatomy ≠Physiology
 Modulated by
 sensory information
(food in gut)
 descending modulation
(hunger/satiety of
animal)
Term
how was physical therapy influenced by early study of central pattern generators ?
Definition
it was heavily influenced by reflexes, so we used to do a LOT of reflex testing.
Term
decerebrate animal experiment. what did this show?
Definition
no cerebrum, limbs are straight out and has lots of tone. when you then knock out the dorsal roots, the animal falls flat, so reflexes were thought to be extremely important.
Term
we moved from this half center model to a modular network model. how does this model work?
Definition
the modular network model says that each muscle group has its motorneurons, inhibitory interneurons, and excitatory interneurons, all of which are highly interconnected to other muscle groups.
Term
what is nialamide
Definition
it is a norepinephrine reuptake inhibitor.
Term
what is 4 aminopyridine?
Definition
cuts down K+ leak channels (increases excitability.)
Term
by
electrical stimulation of high
threshold cutaneous & muscle
afferents (FRAs) in spinalized
cats treated with L-Dopa and
nialamide
Definition
with just one input, you get a rhythmic pattern of output (first flexor fires, then extensor, and back again!) reciprocal
Term
what are some neurotransmitters that are used to initiate or inhibit locomotion?
Definition
Excitatory (primarily glutamate)
Specific agonists elicit rhythmic activity in virtually
all systems –inconsistent with locomotion
Inhibitory (antagonists)
Strychnine (glycine) and biccuculine (GABAA)
Term
Both NE and 5HT
increase the
excitability of the motor neurons and can lead to what?
Definition
spontaneous generation of locomotion (walking half rat)
Term
5HT and NMDA trigger locomotor patterns similar to regular real locomotion. what does this mean?
Definition
this means you do not NEED sensory input to get locomotion
Term
has this rhythmic generator been activated in humans with a complete spinal injury?
Definition
epidural electrodes at
T11-L1 (Dimitrijevic et al.
1998)
 Variable “pattern”
generation with variable
intensity stimulation
Term
why can’t really identify CPGs in humans?
Definition
Unable to isolate spinal cord networks from both descending and
afferent sources
 Does it really matter – do humans demonstrate automaticity
related to stepping behaviors?? YES
Term
what is the Mesencephalic Locomotor Region (MLR)?
Definition
its in the midbrain (brainstem) closely related to the reticular system. it has Direct projections from cortex, basal ganglia,
thalamus.
Term
how are actions exerted through the Mesencephalic Locomotor Region?
Definition
medial reticular formation (reticulospinal
pathway)
– chemical and electrical activation
– modulates with ipsilateral swing phase
– posture/muscle tone (thermostat-like behavior)
ventrolateral/ventromedial spinal pathways
– reduction in postural tone/locomotor ability with
ventral spinal hemisection (Brustein and Rossignol
1998)
– Some recovery can still occur
Term
what is the significance of the experiment where EMG activity was recorded from a walking decerebratecat with
hindlimb muscles that were de-afferented. (LG = lateral
gastrocnemius, EDB = extensor digitorum brevis, IP = iliopsoas, ST
= semitendinosus
Definition
descending pathways can modulate this. you do NOT need afferent pathways
Term
how can you walk without afferent input or descending commands?
Definition
from CPG's. we saw emg activity in decerebrated + deafferented cats. no descending commands and no sensory, but the motor units were still firing.
Term
what does this experiment show? hronic spinal cats
transected after birth
(Forssberg et al. 1980)
 Treadmill walking
(motorized)
 EMG recorded from
hindlimb muscles
Definition
you don't need the brain, you can just have sensory input and get learning.
Term
how is Load through stance-phase limb afferent input important?
Definition
Too little – reduced Ia, Ib,
cutaneous input
 Too much – 1) can’t bear weight
and 2) Doesn’t allow swing
initiation in late stance

sensory information reinforces the stance phase.
Term
how is Hip extension in late stance important?
Definition
Stretch of sartorius muscle
activates whole limb flexion
 Moving limb back increases
stretch earlier . .
 Moving limb forward reduces
swing initiation (Lam and Pearson
2001) Ia and group II inputs. this is spinally mediated!
Term
Hip flexion in swing triggers ??
Definition
opposite limb extension
Term
Behavior of limb dependent on
multiple afferent inputs. what are some examples?
Definition
 IF hip flexed and extensor
muscle force high . .
THEN prolong stance phase
duration
Term
IF hip extended and extensor
muscle force low?
Definition
then initiate swing phase
Term
Complete injury –
NE alpha 2
agonists
in cats triggers
locomotion 8 days
post-SCI
Incomplete injury –
specific NE alpha 1 and
5HT agonist facilitate
weightbearing
Definition
Term
Different CPGs
for different
tasks or available
neural circuits
used selectively??
Definition
we have shared networks and you can select what you need based on inputs and CPGs
Term
what is a synergy?
Definition
roup of muscles activated
in a fixed balance
 Across single or multiple joints
 Synchronous (simultaneous) (walking, standing) vs
 Temporal (time-varying) synergies
Term
why do we have synergies?
Definition
Provides variety of movement strategies based on
“task-relevant subspace of control variables”
 Simplified control of particular biomechanical
features of the limb (global limb vs single joint
 Primitive solution to motor coordination
Term
how do synergies function?
Definition
Selection of interneuronal subgroups
Term
how does the telencephelon modulate locomotion?
Definition
Motor cortex stimulation initiates locomotion
 80% of corticospinal fibers modulate with locomotor
activity (increased to flexors; Drew et al. 2002)
 Contribution of corticospinal tracts
bilateral pyramidotomy – temporary alteration in gross
walking pattern (depends on lesion size)
dorsolateral spinal hemisection (Jiang and Drew 1996)
– toe drag; no ladder or beam walking
– inability to modulate stepping responses to first obstacle (this is due to coritcal control of distal flexors)
 Humans: Motor Evoked Potential modulation greater to TA than MG
(Schubert et al. 1997; Capaday et al. 1999

extensors are more spinally mediated, flexors are more cortically mediated.
Term
how does the diencephalon modulate locomotion
Definition
Behavioral context for locomotion (Sinnamon
1993)
Exploratory system
– basal ganglia, hippocampus - inhibitory control
Appetite system
– lateral hypothalamus, perifornix
– “brings in contact with incentive and consummative
stimuli”
Defense system
– Medial hypothalamus, periaqueductal gray
– “increase distance between threatening/painful stimuli”
Direct projections from these centers to
brainstem locomotor regions
Term
how does stimulation of the mesencephalic locomotor region initiate CPGs?
Definition
the signal goes from MLR --> medial reticular formation --> ventrolateral or ventromedial spinal pathways -->CPG's
Term
how does feedback control motor behaviors?
Definition
Utilize afferent
information on moment to
moment basis
sensory signal
compared with desired
state (i.e., internal
representation) –
reference signal
Difference between
actual and intended
behavior–error signal
 Negative/proportional
feedback in closed loop
chain(e.g., thermostat)
 Gain of feedback (ratio of
input to output)
Term
how does feedforward control motor behaviors??
Definition
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
Term
how do we know that Motor actions have similar features regardless of conditions ?
(Donald Hebb 1950s
Definition
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
Term
what are the parts of the sensorimotor cortex?
Definition
 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 parietal cortex
Term
what do basal ganglia do?
Definition
initiation and
selection of motor programs
Term
what does the cerebellum do?
Definition
coordination, timing, learning
Term
what is the role of the primary motor cortex?
Definition
articipates 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
Term
what is the role of the Primary somatosensory
cortex
Definition
rovides sensory information
required for movement
planning and initiation
 modulation of ongoing
movement
Term
how does the cortex control movement?
Definition
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 musclesor
movementswere represented in
the cortex. (its both, moreso movements)
individual muscles are
represented in many locations.
Term
pyramidal decussation happens where?
Definition
at the base of the medulla for cortex control of motor signals
Term
what is the descending pathway of modulation?
Definition
The 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.
VentralCST projects bilaterally to
motor neurons that innervate axial
muscles and to the intermediate
zone.
LateralCST (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
Term
wrist and fingers are the last thing to come back after cortical damage. why?
Definition
there are many interneurons which can be modulated for a muscle like the bicep or pec major if you lose it. the fingers, however, are probably just controlled by 1 part of the motor cortex with little internuerons?
Term
What is transcranial magnetic stimulation
(TMS) – also Transcranial
Electrical Stimulation (TES)
Definition
(magnetic fields induce currents in wireds - faraday's law). Magnetic coil generates field
along cortical surface

Activate smaller cortical
cells which activate
pyramidal cells
(TES may bypass this, though
mechanisms controversial)
Term
how much of the coritcal tracts are from M1 region?
Definition
Cortical tracts
 40% from M1 region
 S1 and PM areas contribute
significantly
 Typically to interneuron
pools – except to distal
motor pools
Term
Corticospinal neurons make
direct excitatory connections
with only some motor neurons. which ones?

After a bilateral resection of the
corticospinal tracts, monkeys unable
to grasp small objects with
thumb/index finger
Definition
Direct connections for fine control
of the digits.
Term
how is Walking affected after a bilateral resection of corticospinal tracts?
Definition
unable to negotiate in feedforward
manner (obstacles, balance beam/ladder)
– Toe/ankle dorsiflexion problems, can’t walk balance beam
Term
The discharge of action potentials by neurons in the primary motor
cortex (CM = corticomotoneuronal cell) depends on the motor task. how does this affect precision grip vs power grip?
Definition
Precision vs Power grip involve separate cortical/subcortical
circuits acting on similar IN/MN pools
Term
do cortical cells encode direction-specifically?
Definition
yes. they are Population vectors that encode both direction
and magnitude
Term
do motor cortical cells also encode for force?
Definition
yes. Motor cortical cells alter firing
patterns according to extent of
muscle activity required to complete a
task
 Tonic and Phasic cortical cells may
participate
Term
how is Direction-specific information and force specific information encoded in cortical cells
Definition
when you load AGAINST the preferred direction, cortical cells fire very hard.

when you load WITH the preferred direction, the cells don't fire as hard
Term
what does the Premotor area do?
Definition
 involved in goal-directed movements
 Activity prior to visually-guided movements
Term
what does the Supplementary motor area do?
Definition
 ensures correct sequencing of movement (order of
movement)
– biomechanical constraints
– task performed
– external conditions
 Activity prior to internally-guided movements
Term
what does the Posterior parietal cortex do?
Definition
 encodes complex sensory information
 internal sensory representation
Term
what are the four premotor areas?
Definition
 Lateral premotor areas
– Dorsal premotor
– Ventral premotor
 Medial premotor areas
– Supplementary motor area
–Cingulate area
Term
visually guided inputs go where?
Definition
premotor areas
Term
what is the difference in activation of the Premotor (dorsal or ventral) and
Supplementary motor area during visually guided vs.
internally guided movements
Definition
for a visually guided movement, you have a normal distribution of activation of the two areas.

For an internally guided movement (eyes closed!) you have less activation of the premotor area, and more activation of the supplementary motor area
Term
when is the supplemetary motor areas active?
Definition
active during
sequential (vs just
simple movements)
 Active during mental
rehearsal
Term
when are the ventral and dorsal premotor areas active?
Definition
Different tasks utilize
different pathways w/
external cues
 Reaching (visual-parietal-dorsal
PM areas)
 Grasping (visual-parietal-ventral
PM areas)
 Activity related to association
of stimuli to learned
movements
 Oftentimes activity without
movement (mirror neurons)
Term
how do the basal ganglia and cerebellum modulatie descending motor patterns?
Definition
Two systems of brain stem
neurons (medial and lateral
descending systems) receive input
from the cortex & subcortical
nuclei and project to the spinal
cord.
The cerebellumand basal ganglia
provide feedback through the
thalamus(not shown) that regulate
cortical and brain stem motor
areas.
The basal gangliaare involved in
motivation and initiation and
selection of motor programs.
The cerebellumis involved with
the timing and coordination of
movements (error detection) and
with the learning of motor skills
Term
is cerebellum ipsalateral or contralaterally controlling?
Definition
same saide, ipsalateral
Term
descending modulation by the cerebellum.
Definition
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 pathway
Term
what is the vestibulocerebellum? what is its function?
Definition
flocculonodular lobe (most primitive)
 Receives vestibular, postural, ocular
information; controls balance and eye
movements
Term
what is the spinocerebellum (vermis)? what is its function?
Definition
intermed hemisphere and Vermis –
interposed and fastigial nuclei
 proprioceptive/exteroceptive inputs
 Governs “performance of movement
Term
what is the cerebrocerebellum? what is its function?
Definition
Lateral hemisphere –Dentate nuclei
 Projections from pontine nuclei
 Information of planned movement
Term
what are the layers of the cerebellar cortex?
Definition
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
Term
mossy fibers synapse with which cerebellar cells?
Definition
granule cells
Term
granule cells go into which cerebellar cells?
Definition
parallel fibers
Term
parallel fibers synapse with the which cerebellar cells?
Definition
purkinje cells
Term
where do mossy fibers originate?
Definition
pontine nuclei
Term
where do climbing fibers originate?
Definition
inferior olivary nucleus.
Term
what is the relay center of the brain?
Definition
the thalamus
Term
role of cerebellum - 1st movement vs 2nd movement.
Definition
we get refinement of the motor signal based on error signal from the cerebellum
Term
role of cerebellum - 1st movement vs 2nd movement.
Definition
we get refinement of the motor signal based on error signal from the cerebellum
Term
what is a transition?
Definition
immediate change in behavior
 driven by prior experience and the ability to predict that
new demands will exceed “current state” (feed-forward
strategies)
Term
what is an adaptation?
Definition
radual change in behavior that results from experience
(“feedback strategies”)
 Driven by demands that exceed “current state"
Term
what is learning?
Definition
Relatively permanent changes
 Resulting from repeated exposure (adaptation may be a
precursor)
Term
prism adaptation research
Definition
Prisms inserted into eyeglasses
Displace visual field
Leads to initial errors in
movement accuracy, but cerebellum comparator functions lead to adaptations and then learning.
Term
With extensive training, throwing
with wedge prisms can become a
skill. Adaptations result in Learning–which allow faster Transitions
Definition
Term
locomotor adaptations - podokinetic system
Definition
after walking on a circular treadmill for a while, if you get off and close your eyes and walk, you walk in circles, and you have no idea you are walking in circles
Term
how does podokinetic adaptation occur?
Definition
there are both vestibular and cerebellar contributions to the adaptation response.
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