lecture14 fin, 15 init

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ackman678
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---
## Central pattern generators organize the cycle of locomotion for terrestrial mammals
## Central pattern generators organize the cycle of activity for locomotion
<figure><img src="figs/Neuroscience5e-Fig-16.15-0_312da6c.jpg" height="400px"><figcaption>Neuroscience 5e Fig. 16.15</figcaption></figure>
@@ -627,14 +627,25 @@ Note:
---
## Central pattern generators organize the cycle of locomotion for terrestrial mammals
## Central pattern generators organize the cycle of activity for locomotion
<div><iframe src="https://www.youtube.com/embed/wPiLLplofYw" width="420" height="315"></iframe><figcaption>Locomotion in decerebrate cat</figcaption></div>
Note:
S Grillner, P Zangger. On the central generation of locomotion in the low spinal cat. Experimental Brain Research, 1979
---
## Central pattern generator model circuit
## Central pattern generator circuit model for swimming
<div><figcaption class="big">Lamprey</figcaption><img src="figs/Lamprey_anatomy_5f701d1.png" height="100px"><figcaption>[CC0](https://en.wikipedia.org/wiki/Lamprey)</figcaption></div>
<div><figcaption class="big">Simplified lamprey CPG model. E, excitatory; I, inhibitory; M, motor neuron
</figcaption><img src="figs/lamprey-spinal-cpg-model_a1862d9.svg" height="300px"><figcaption>J. Ackman, [CC0](https://wiki.creativecommons.org/wiki/CC0). Based on Grillner et al., *Brain Res Rev* 2008</figcaption></div>
<!--
<figure>
<figcaption>Interlimb coupling (C) with mutually inhibitory connections.
E, extensor. F, flexor. Arrows, excitatory. Closed circles, inhibitory
@@ -642,10 +653,20 @@ E, extensor. F, flexor. Arrows, excitatory. Closed circles, inhibitory
<img src="figs/Ting-JNeurophys1998-Fig8_7b01f81.jpg" height="400px">
<figcaption>Ting et al., *J Neurophysiol* 1998, Fig. 8</figcaption></figure>
-->
<!-- <figure><img src="figs/cpg-model-circuit_ce4156c.png" height="400px"><figcaption></figcaption></figure> -->
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TODO: add midline, interneuron labels to svg file.
TODO: add neurons for more complete model representation
TODO: make model with flexor/extensors for vertebrate locomotion
lamprey
: ancient vertebrate
: jawless fish
: good model for cpg spinal cord circuits, well worked out
simple network of neurons that could result in alternating flexor and extensor muscle movements for locomotion and be basis of a central pattern generator circuit.
spinal locomotor and brainstem respiratory CPGs (Yuste et al, Nat Rev Neurosci 2005)

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## Overall organization of neural structures that control movement
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Lower motor
system
@@ -38,7 +28,7 @@ learning
2016-03-01 09:46:39
<div><img src="figs/Neuroscience5e-Fig-16.01-0_f1cc94b.jpg" height="100px"><figcaption></figcaption></div>
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## Upper motor control
* Axons from the upper motor neurons descend to influence the local circuits in the brainstem and spinal cord that organize movements.
* Upper motor pathways include several brainstem centers and a number of cortical areas in the frontal lobe.
* Brainstem centers are especially important for postural control.
* Motor and premotor cortex are responsible for the planning and precise control of complex sequences of voluntary movements.
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@@ -65,16 +52,14 @@ Note:
## Arrangement of motor neurons and local circuit interneurons within the spinal cord
* Medial ventral horn: motor neuron pools that innervate axial muscles and proximal limb muscles
* Lateral ventral horn: motor neurons that innervate distal limb muscles.
* Local circuit interneurons lie in the intermediate zone of the spinal cord grey matter.
Neuroscience 5e Fig. 16.3
<div><img src="figs/neuro4e-fig-16-03-0_82eb56b.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/neuro4e-fig-16-03-0_82eb56b.jpg" height="100px"><figcaption></figcaption></div>
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## Overview of descending motor control
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.01-1R_abcbb34.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.01-1R_abcbb34.jpg" height="100px"><figcaption></figcaption></div>
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## Arrangement of motor neurons and local circuit interneurons within the spinal cord
* Medial intermediate zone local circuit neurons project to medial ventral horn motor neurons.
* Medial local circuit neurons have axons that may project to targets along the entire length of the cord, and also cross the midline to innervate contralateral side.
* Lateral regions of the intermediate zone contain local neurons that synapse with motor neurons in the lateral ventral horn.
* Lateral circuit neurons project over a smaller area and do not cross the midline.
* Allows distal regions to act independently of each other.
<div><img src="figs/neuro4e-fig-16-04-0_328e2b3.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/neuro4e-fig-16-04-0_328e2b3.jpg" height="100px"><figcaption></figcaption></div>
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## Overview of descending motor control
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Neuroscience 5e Fig. 17.1
<div><img src="figs/Neuroscience5e-Fig-17.01-2R_08a492a.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.01-2R_08a492a.jpg" height="100px"><figcaption></figcaption></div>
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@@ -159,16 +120,12 @@ lateral ventral horn contains lower motor neurons that mediate skilled voluntary
## Medial brainstem pathways modulate theaction of motor neurons in the ventromedial area
* Vestibular nuclei:
* Receive information from inner ear
* Project to medial regions of spinal gray matter.
* Controls axial muscles and proximal limbs.
* Called the vestibulospinal tract.
<div><img src="figs/Neuroscience5e-Fig-17.11-1R_649492a.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.11-1R_649492a.jpg" height="100px"><figcaption></figcaption></div>
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@@ -179,16 +136,12 @@ info from semicircular canals in inner ear. balance. feedback postural control
## Medial brainstem pathways modulate theaction of motor neurons in the ventromedial area
* Reticular formation:
* Receives input from higher motor cortex.
* Complex network of circuits located in the core of the brainstem-from midbrain to medulla.
* Important for posture.
* Called the reticulospinal tract.
<div><img src="figs/Neuroscience5e-Fig-17.11-2R_f9fc798.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.11-2R_f9fc798.jpg" height="100px"><figcaption></figcaption></div>
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@@ -198,17 +151,7 @@ feedforward postural control. stabilization during ongoing movements.
## Location of the reticular formation in relation to some other major landmarks
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/neuro4e-fig-17-03-0_18e53fa.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/neuro4e-fig-17-03-0_18e53fa.jpg" height="100px"><figcaption></figcaption></div>
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## Medial brainstem pathways modulate theaction of motor neurons in the ventromedial area
* Superior colliculus
* Projects to medial cell groups in the cervical cord
* Influences neck muscles (colliculospinal tract)
* But major output of superior colliculus to spinal cord mediated by reticular formation. Axial musculature control of neck and performing orienting movements of head and eye movements.
<div><img src="figs/neuro4e-fig-17-02-3r_5f38a2d.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/neuro4e-fig-17-02-3r_5f38a2d.jpg" height="100px"><figcaption></figcaption></div>
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@@ -236,19 +176,9 @@ tectospinal tract.
## The medial descending motor pathways
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/image_d920d94.pdf" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/image_d920d94.pdf" height="100px"><figcaption></figcaption></div>
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## Feedforward processing
* Able to predict changes in posture, and generate an appropriate stabilizing response.
* Some muscles fire in anticipation of a need for postural adjustment.
* Reticulospinal tract important for this process. If it is severed in a cat, no change in compensatory muscles occur during the process.
* Stimulate motor cortex in the right place can induce paw lifting, and several limb muscles to fire. Inhibition of the reticulospinal tract will allow the paw to move but will prevent the movement of other limbs.
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## Anticipatory maintenance of body posture
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Severed reticulospinal tract will allow biceps to fire but
will not allow the gastrocnemius to fire for posture.
@@ -294,7 +211,7 @@ Neuroscience 5e Fig. 17.13
<div><img src="figs/Neuroscience5e-Fig-17.13-0_5df4fb2.jpg" height="100px"><figcaption></figcaption></div>
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## Feedforward and feedback mechanisms of postural control
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.14-0_52ca14e.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.14-0_52ca14e.jpg" height="100px"><figcaption></figcaption></div>
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## Primary motor cortex and premotor cortex are in the frontal lobe
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.02-0_d55a793.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.02-0_d55a793.jpg" height="100px"><figcaption></figcaption></div>
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## Primary motor cortex
* Located in the precentral gyrus
* Receives inputs from S1, posterior parietal structures (incorporates multiple sensory modalities, used for planning).
* Controls contralateral side of the body
* Topographic organization- body represented across the medial-lateral axis. More space given to areas of fine motor control. Multiple neurons can get the same muscle to fire- not located in exact same place in cortex.
<div><img src="figs/Neuroscience5e-Fig-17.05-0_ed37921.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.05-0_ed37921.jpg" height="100px"><figcaption></figcaption></div>
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## Somatotopic representation across S1 and M1
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
[http://www.pbs.org/wgbh/aso/tryit/brain/probe.html](http://www.pbs.org/wgbh/aso/tryit/brain/probe.html)
<div><img src="figs/5892_CorticalTopographybox!_ecb699f.png" height="100px"><figcaption></figcaption></div>
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## Motor cortex
* Located in the frontal lobe
* Several adjacent and interconnected areas
* Primary motor cortex located in the precentral gyrus
* Gets input from sensory cortex, basal ganglion and cerebellum
* Has 6 layers, layer V is the output layer (pyramidal cells, including the large Betz cells consisting of about 5% of projection to spinal cord and concerned with fine distal movements)
* Primary pathway- the corticospinal tract. Axons cross in the caudal medulla, and innervate in lateral ventral horns
<div><img src="figs/image1_3f82ace.png" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/image1_3f82ace.png" height="100px"><figcaption></figcaption></div>
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@@ -408,16 +287,6 @@ Note:
## Pathways from the motor cortex to the spinal cord
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Indirect pathway: postural
adjustments, especially for
@@ -438,7 +307,7 @@ Neuroscience 5e Fig. 17.5
<div><img src="figs/Neuroscience5e-Fig-17.15-0_f03ec91.jpg" height="100px"><figcaption></figcaption></div>
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## The corticospinal and corticobulbar tracts
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Neuroscience 5e Fig. 17.4
<div><img src="figs/Neuroscience5e-Fig-17_42dcdad.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17_42dcdad.jpg" height="100px"><figcaption></figcaption></div>
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@@ -485,11 +344,8 @@ Corticobulbar is blue, corticospinal in red. Note that corticospinal cross the
## Facial pathway
* The primary pathway to facial muscles is the corticobulbar pathway.
* Projection from motor cortex to motor nuclei in brainstem that control facial muscles.
* Some of these projections are bilateral and some only contralateral.
* Important for diagnosis where motor damage occurs after a stroke.
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## Patterns of facial weakness and their importance for localizing neurological injury
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Neuroscience 5e Box 17A
<div><img src="figs/Neuroscience5e-Box-17A-0_47bce5b.jpg" height="100px"><figcaption></figcaption></div>
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## Motor fields
* A stimulation of a neuron in the primary motor cortex will get multiple muscles to fire, and will inhibit other muscles.
* Stimulating any of multiple upper neurons can get the same muscle to fire.
* The “receptive field” of a upper motor neuron has to do with organized movements rather than specific muscle groups.
* Upper motor neurons therefore act upon more than one motor pool.
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## What do motor maps represent?
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Box-17B-0_a572ded.jpg" height="100px"><figcaption></figcaption></div>
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## Activity of single upper motor neurons is correlated with muscle movements
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Neuroscience 5e Fig. 17.6
<div><img src="figs/Neuroscience5e-Fig-17.06-2R_34ac4f3.jpg" height="100px"><figcaption></figcaption></div>
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## Purposeful movements resulting from prolonged microstimulation of the primary motor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.07-0_b1923da.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.07-0_b1923da.jpg" height="100px"><figcaption></figcaption></div>
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## Directional tuning of an upper motor neuron in the primary motor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Monkey trained to move joystick in response to light
<div><img src="figs/Neuroscience5e-Fig-17.08-1R_3cd5f1a.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.08-1R_3cd5f1a.jpg" height="100px"><figcaption></figcaption></div>
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## Directional tuning of an upper motor neuron in the primary motor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Activity of a single neuron recorded in motor cortex
is dependent on the direction of the future movement.
@@ -666,7 +459,7 @@ Neuroscience 5e Fig. 17.8
<div><img src="figs/Neuroscience5e-Fig-17.08-2R_256673f.jpg" height="100px"><figcaption></figcaption></div>
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## Directional tuning of an upper motor neuron in the primary motor cortex
* Individual neurons are tuned too broadly to accurately predict direction of movement
* By comparing populations of neurons, one can calculate a direction.
* Can use the activity of motor cortex to control robots.
[https://www.youtube.com/watch?v=7kctOHnrvuM](https://www.youtube.com/watch?v=7kctOHnrvuM)
@@ -688,9 +479,9 @@ Neuroscience 5e Fig. 17.8
<div><img src="figs/Neuroscience5e-Fig-17.08-3R_4ac9fb7.jpg" height="100px"><figcaption></figcaption></div>
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## Section of pyramidal tracts in monkeys produces loss of independent digit control
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Intact (normal)
After section of
corticospinal fibers
<div><img src="figs/image_a144227.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/image_a144227.jpg" height="100px"><figcaption></figcaption></div>
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@@ -726,16 +507,6 @@ corticalspinal, lateral dorsal input for control of distal/fine movements of the
## Primary motor cortex and the premotor area in human
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
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## Primary motor cortex and the premotor area in macaque monkey
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.09-0_1d7aeaa.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.09-0_1d7aeaa.jpg" height="100px"><figcaption></figcaption></div>
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@@ -777,15 +538,10 @@ the frontal eye fields organize voluntary gaze shifts. The cingulate motor areas
## The premotor cortex
* Lies adjacent (rostral) to the primary motor cortex
* Makes extensive reciprocal connections with the primary motor cortex
* Projects directly to spinal cord (30% of axons in the corticospinal tract).
* Lateral premotor cortex- has neurons that are tuned to a particular direction of movement (like primary motor cortex) but differs in that they fire earlier than neurons in the primary motor cortex. This is especially important in conditional motor tasks, that pair a movement with a visual cue.
* During the pairing of a visual cue with a motor task, the neurons will fire before any initiation of the task. This is used for intentions.
* Lesions in monkey prevent vision conditioned tasks, although vision is fine and the task can be done in other ways.
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@@ -796,19 +552,9 @@ thes neurons encode intention to perform a movement rather than just the movemen
## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
[https://www.youtube.com/watch?v=RuK2Y8JojN8](https://www.youtube.com/watch?v=RuK2Y8JojN8)
<div><img src="figs/Neuroscience5e-Fig-17.10-1R_46b3807.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.10-1R_46b3807.jpg" height="100px"><figcaption></figcaption></div>
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@@ -832,17 +578,7 @@ based on Giacomo Rizzolatti et al, 1996
## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.10-2R_cf5432f.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.10-2R_cf5432f.jpg" height="100px"><figcaption></figcaption></div>
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@@ -856,17 +592,7 @@ does not respond when pliers are used to interact with food.
## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Fig-17.10-3R_10c4215.jpg" height="100px"><figcaption></figcaption></div>
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@@ -893,10 +619,9 @@ http://nautil.us/blog/mirror-neurons-are-essential-but-not-in-the-way-you-think
## Premotor cortex two-hand coordination
* The monkey has learned the task: push the object through the hole and catch it with the other hand
* With damage to premotor cortex, cannot coordinate two hands to do the task
<div><img src="figs/image2_08b969e.png" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/image2_08b969e.png" height="100px"><figcaption></figcaption></div>
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## Medial premotor cortex
* Mediates the selection of movements.
* Specified by internal rather than external cues.
* Important for selecting movements based on memory, not in response to cues.
* Cells will fire when just thinking about an event.
Note:
@@ -922,16 +644,6 @@ Note:
## Planning movement sequence without moving activates supplemental motor area (medial premotor area)
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
Mental rehearsal of finger sequence
Motor cortex
@@ -946,7 +658,7 @@ Supplementary
motor area
<div><img src="figs/image1_52fff81.jpg" height="100px"><figcaption></figcaption></div>
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## Activation of motor areas depend different on behavioral context
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
@@ -990,7 +692,7 @@ Learned
Sequence
<div><img src="figs/image2_132bba7.jpg" height="100px"><figcaption></figcaption></div>
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## Effects of damage to the cerebral cortex
* By investigating patients with various types of brain damage we can see how the various components of motor performance may be affected. Examples:
* Lesions to primary motor cortex (e.g. from a stroke) result in loss of voluntary movements on the contralateral (opposite) side of the body.
* Apraxia is the specific loss of the ability to plan and correctly perform co-ordinated motor skills, mainly as a result of damage to the supplementary motor area. Speech disorders result from damage to motor cortex.
* Patients can move muscles, and walk on command but can no longer link gestures to a coherent act, or to recognize the appropriate use of an object even though they can recognize what an object is.
Note:
@@ -1017,9 +716,7 @@ Note:
## Damage to cortex: alien limb syndrome
* A disorder in which person feels unable to control movements of a body part, believes that the limb is alien, or believes that the body part has its own personality
* It is typically associated with lesions in the supplementary motor area or those affecting blood flow to the anterior regions of the corpus callosum and the anterior cingulate
* Man who simultaneously tried to strangle and save his wife from himself.
[https://www.youtube.com/watch?v=dIBBDuQrd-I](https://www.youtube.com/watch?v=dIBBDuQrd-I)
@@ -1032,16 +729,6 @@ Note:
## The Babinski sign
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
[https://www.youtube.com/watch?v=ZFu7bdbnZx8](https://www.youtube.com/watch?v=ZFu7bdbnZx8)
[https://www.youtube.com/watch?v=oI_ONptx2Ns](https://www.youtube.com/watch?v=oI_ONptx2Ns)
@@ -1050,7 +737,7 @@ Neuroscience 5e Fig. 17.16
<div><img src="figs/Neuroscience5e-Fig-17.16-0_08731e1.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Fig-17.16-0_08731e1.jpg" height="100px"><figcaption></figcaption></div>
Note:
@@ -1086,17 +773,7 @@ after several days recovery begins (not fully understood) and includes
## Signs of motor neuron lesions
* Body Level One
* Body Level Two
* Body Level Three
* Body Level Four
* Body Level Five
<div><img src="figs/Neuroscience5e-Tab-17.01-0_1f5d45b.jpg" height="100px"><figcaption></figcaption></div>
<div><img src="tmp/Neuroscience5e-Tab-17.01-0_1f5d45b.jpg" height="100px"><figcaption></figcaption></div>
Note:
@@ -1107,25 +784,16 @@ Note:
## Principles
* Motor
* Output to muscles via ventral root
* Two main pathways:
* 1. Ventromedial system for balance, posture and controlling head & eye movements. Important for muscles of legs & trunk needed for walking.
* 2. Dorsolateral system for controlling movements of upper limbs & extremities such as fingers and toes as well as movement of facial muscles.
*
* Sensory
* Input to primary somatosensory area via dorsal root
* Two main pathways:
* 1. Dorsal spinothalamic tract for proprioception (body awareness and position in space) and haptic feedback (sensation of fine touch and pressure) crosses in medulla
* 2. Ventral spinothalamic tract for nocioceptive information crosses over in spinal cord
Note: