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Three types of movements (antialphabetic):
- [blank_start]Voluntary[blank_end] movements
- [blank_start]Rhythmic[blank_end] movements
- [blank_start]Reflexes[blank_end]
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Voluntary
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Rhythmic
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Reflexes
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The visuomotor delay is approximately [blank_start]200[blank_end] ms.
The proprioceptive delay is approximately [blank_start]50[blank_end] ms.
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200
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250
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300
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150
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50
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30
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80
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150
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Three things that make motor control difficult (alphabetic):
- [blank_start]environmental[blank_end] uncertainty
- [blank_start]motor[blank_end] noise
- [blank_start]sensory[blank_end] noise
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environmental
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motor
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sensory
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The human body has "only" 600 muscles. Nonetheless motor control is very difficult, since the number of possible movement patterns increases [blank_start]exponentially[blank_end] with the [blank_start]degrees of freedom[blank_end].
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exponentially
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degrees of freedom
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The principle of [blank_start]redundancy[blank_end] means that there is a [blank_start]gap[blank_end] between high- and low-level specification. Any high level task can be achieved in [blank_start]infinitely[blank_end] many low level ways.
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redundancy
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gap
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infinitely
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[blank_start]Fitt's[blank_end] law describes the [blank_start]speed-accuracy[blank_end] trade-off.
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In simple grasping tasks the paths of hand movements are usually [blank_start]straight[blank_end] and the speed profiles are [blank_start]bell-shaped[blank_end].
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The two-thirds power law:
[blank_start]angular[blank_end] velocity ~ [blank_start]curvature[blank_end]^(2/3)
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Three levels of analysis:
(1) [blank_start]Computational[blank_end]: What is the [blank_start]problem[blank_end] the brain is trying to solve?
(2) [blank_start]Algorithmic[blank_end]: What is the [blank_start]strategy[blank_end] to solve this problem?
(3) [blank_start]Neuronal[blank_end]: How is it done by the nervous system?
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Computational
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Algorithmic
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Neuronal
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problem
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strategy
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The forward model [blank_start]predicts behaviour[blank_end] based on [blank_start]motor commands[blank_end].
The inverse model [blank_start]calculates motor commands[blank_end] from [blank_start]desired behaviour[blank_end].
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Feedforward ([blank_start]open[blank_end]-loop) control:
Because of the delays in the sensorimotor system, open-loop control is used for [blank_start]short[blank_end] movements.
Disadvantages:
- [blank_start]Inaccuracies[blank_end] cannot be corrected
- [blank_start]Unexpected changes[blank_end] will not be taken into account
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short
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long
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open
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closed
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Inaccuracies
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Delays
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Noise
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Unexpected changes
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Inaccuracies
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Noise
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In order for feedforward control to be sufficient, the [blank_start]inverse[blank_end] model would have to be perfect.
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Feedback (closed-loop) control can compensate for errors due to
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In an experiment, subjects are given 400g blocks and practice to lift them.
After training the weight is surprisingly increased to 800g.
The experiment shows the subjects use
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The [blank_start]gain[blank_end] factor is the amount by which the [blank_start]corrective[blank_end] motor command is increased or decreased per unit of error.
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[blank_start]High[blank_end] gain and [blank_start]high[blank_end] sensorimotor delay can lead to [blank_start]overcompensation[blank_end] and [blank_start]instabilities[blank_end].
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High
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Low
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high
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low
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overcompensation
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too little compensation
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instabilities
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a stable motor command
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An [blank_start]efference[blank_end] copy is an internal copy of an outflowing ([blank_start]efferent[blank_end]), movement-producing signal generated by the motor system.
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efference
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afference
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efferent
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afferent
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Two strategies to compensate for sensory delays:
(1) [blank_start]Intermittency[blank_end] of movements
(2) [blank_start]Prediction[blank_end]
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The experiment where you move your eyeball with your hand and the world moves shows that the brain uses [blank_start]motor commands to the eye[blank_end] in order to predict the eye's position.
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What's the problem with using a forward model in feedback control to predict the position of a limb.
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Errors accumulate over time.
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Delay leads to instabilites.
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Actually an inverse model would have to be used.
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Which are the main structures of the CNS involved in motor control?
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A motor unit consists of [blank_start]a spinal motor neuron[blank_end] and [blank_start]the muscle fibres that it innervates.[blank_end]
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a spinal motor neuron
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a sensory interneuron
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a pyramidal motor neuron
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the muscle fibres that it innervates.
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the motor neurons it projects to.
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the disc in the spinal cord it projects
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A muscle is typically innervated by [blank_start]a few hundred[blank_end] spinal motor neurons.
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a few hundred
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a few thousand
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about 50
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about 50.000
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The [blank_start]innervation number[blank_end] tells how many muscle fibres a motor neuron innervates.
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The innervation number
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indicates the increment in force when the motor unit is activated.
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can be very different for the same muscle.
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is the same for all neurons that project to one muscle
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indicates the speed by which the muscle is activated
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The muscle force is controlled by
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recruitment (how many motor neurons of the unit fire)
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the discharge rate of motor units
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the innervation number of the motor unit
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the axon diameter of the motor neuron
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Motor units differ in [blank_start]speed[blank_end] and maximum [blank_start]force[blank_end].
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Recruitment:
[blank_start]Small[blank_end] motor neurons are recruited first because of the [blank_start]high[blank_end] resistance within their axon.
They have a [blank_start]lower[blank_end] innervation number and therefore produce [blank_start]less[blank_end] force.
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Small
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Large
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high
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low
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lower
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higher
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less
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more
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Spinal reflexes are [blank_start]involuntary[blank_end] and triggered by [blank_start]external stimuli[blank_end].
Despite classical theory they can be quite flexible.
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involuntary
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external stimuli
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The withdrawal from a painful stimulus is called [blank_start]flexion-withdrawal[blank_end] reflex.
The [blank_start]flexor[blank_end] muscle is activated and the [blank_start]extensor[blank_end] muscle is inhibited.
This is called [blank_start]reciprocal innervation[blank_end].
The opposite effect happens on the [blank_start]contralateral[blank_end] limb.
The force [blank_start]does[blank_end] (does or does not) depend on the stimulus intensity.
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flexion-withdrawal
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flexor
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extensor
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reciprocal innervation
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contralateral
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does
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The stretch reflex is a [blank_start]muscle contraction[blank_end] in response to stretching within the muscle.
The antagonist muscle is inhibited, this is called [blank_start]reciprocal innervation[blank_end].
The stretch reflex increases the [blank_start]stiffness[blank_end] of the muscle.
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stiffness
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muscle contraction
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reciprocal innervation
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The areas of the cerebral cortex that are used in motor control:
- [blank_start]Primary motor cortex[blank_end] (M1)
- [blank_start]Premotor cortex[blank_end] (PM)
- [blank_start]Supplementary motor area[blank_end] (SMA)
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Primary motor cortex
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Premotor cortex
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Supplementary motor area
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The motor cortices receive input from
Frage 34
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The motor homunculus is most detailed for the [blank_start]primary motor cortex[blank_end].
It disappears if you [blank_start]zoom on[blank_end].
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primary motor cortex
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premotor cortex
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supplementary motor area
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zoom in
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zoom out
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The motor pathway to lateral motorneurons runs [blank_start]contralateral[blank_end] and controls mainly [blank_start]distal muscles[blank_end].
It includes the [blank_start]corticospinal tract[blank_end].
The patway to medial motorneurons runs [blank_start]ipsilateral[blank_end] and controls mainly [blank_start]stance and posture[blank_end].
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contralateral
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ipsilateral
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distal muscles
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nearby muscles
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corticospinal tract
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medial tract
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ipsilateral
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contralateral
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stance and posture
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fine movements
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unconscious movements
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The corticospinal tract
The connections come from the cortical layer [blank_start]V[blank_end].
Many of these connections terminate on spinal [blank_start]interneurons[blank_end].
Only [blank_start]M1[blank_end] contains neurons projecting directly to spinal motor neurons. These cortical neurons are called [blank_start]corticomotorneurons[blank_end]. Those neurons have only one synapse.
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V
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IV
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III
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II
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I
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interneurons
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medial neurons
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projection neurons
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M1
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PM
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SMA
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corticomotorneurons
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complex neurons
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The axons of corticomotorneurons terminate on [blank_start]spinal motor neurons[blank_end].
The can also influence other muscles via spinal [blank_start]interneurons[blank_end].
Most of the time the excite agonist muscles and inhibit antagonist ones.
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spinal motor neurons
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the muscle fiber
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interneurons
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projection neurons
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receptors
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Reading information about the direction of a movement from several motor neurons is done via [blank_start]population codes[blank_end].
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Neuronal tuning with respect to movement means
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that firing rates of cortical motor neurons correlate with parameters of movement, e.g. direction, velocity, acceleration, ...
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that reinforcement learning can be applied to fine-tune execution of motor tasks.
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that firing rates of cortical motor neurons are highly intercorrelated ("tuning each other")
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Different digits of the hand are controlled by sharply separated areas in M1.
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Four symptoms of cerebellar disorders:
[blank_start]Hypotonia[blank_end]: A diminished resistance to passive limb displacements.
[blank_start]Astasia-abasia[blank_end]: An inability to stand or walk.
[blank_start]Ataxia[blank_end]: The abnormal execution of multijointed voluntary movements, characterized by lack of coordination.
[blank_start]Action tremor[blank_end]: A form of tremor at the end of a movement, when the patient attempts to stop the movement by using antagonist muscles.
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Hypotonia
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Astasia-abasia
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Ataxia
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Action tremor
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The basal ganglia are involved in
Frage 43
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Relation between function and areas of the brain (hypothesis)
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basal ganglia
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motor cortex
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parietal cortex
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cerebellum
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thalamus
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prefrontal cortex
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medulla oblongata
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The motor control loop
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controller
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state estimation
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efference copy
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sensory feedback
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forward model
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inverse model
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gain
Frage 45
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Comparator
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Sensed state
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Input processing
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Sensor
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Forward model
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Inverse model
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Estimated state
Frage 46
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dynamic
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predictive
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sensory
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correction
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estimation
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gain
Frage 47
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Desired state
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Estimated state
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Feedforward controller
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Inverse controller
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Motor command
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State estimation
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Actuator
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Optimal feedback control
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position
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weight
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motor command
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The forward-model is used at the [blank_start]beginning[blank_end] of movements.
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There are slow- and fast-[blank_start]twitch[blank_end] muscle fibers.
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[blank_start]Supraspinal[blank_end] centers play an important role in modulating and adapting spinal reflexes, even to the extent of reversing movements when appropriate.
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Stretch reflex:
In addition to the muscle that is stretched [blank_start]synergistic[blank_end] muscles are contracted.
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M1: Brodman [blank_start]4[blank_end]
PM & SMA: Brodman [blank_start]6[blank_end]
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