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## Overall organization of neural structures that control movement
Lower motor
system
State of muscle
contraction/relaxation
Execute
movement
Output
system
Upper motor
system
Gating
Motor
learning
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<div><img src="tmp/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.
Note:
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## 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
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## Overview of descending motor control
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<div><img src="tmp/Neuroscience5e-Fig-17.01-1R_abcbb34.jpg" height="100px"><figcaption></figcaption></div>
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somatotopic organization of the ventral horn in the cervical enlargement. Locations of descending projections from the motor cortex in the lateral white matter and from the brainstem in the anterior-medial white matter are shown.
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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.
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## Overview of descending motor control
Neuroscience 5e Fig. 17.1
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<div><img src="tmp/Neuroscience5e-Fig-17.01-2R_08a492a.jpg" height="100px"><figcaption></figcaption></div>
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med ventral horn has lower motor neurosn for posteure balance and orienting movements of head and neck during shits of visual gaze. receipve descending input from the pathways orginating mainly in the brainstem, course through the anterior medial white matter of the spional cord and terminate bilaterally.
lateral ventral horn contains lower motor neurons that mediate skilled voluntary movements of the distal extremities. Receive descending projection from the contralateral motor cortex via lateral division of the corticospinal tract.
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## 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.
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<div><img src="tmp/Neuroscience5e-Fig-17.11-1R_649492a.jpg" height="100px"><figcaption></figcaption></div>
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info from semicircular canals in inner ear. balance. feedback postural control
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## 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.
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<div><img src="tmp/Neuroscience5e-Fig-17.11-2R_f9fc798.jpg" height="100px"><figcaption></figcaption></div>
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feedforward postural control. stabilization during ongoing movements.
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## Location of the reticular formation in relation to some other major landmarks
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<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.
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<div><img src="tmp/neuro4e-fig-17-02-3r_5f38a2d.jpg" height="100px"><figcaption></figcaption></div>
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tectospinal tract.
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## The medial descending motor pathways
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<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
Severed reticulospinal tract will allow biceps to fire but
will not allow the gastrocnemius to fire for posture.
EMG= electromyography. Measure muscle APs
Neuroscience 5e Fig. 17.13
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## Feedforward and feedback mechanisms of postural control
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<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
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<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.
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<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
[http://www.pbs.org/wgbh/aso/tryit/brain/probe.html](http://www.pbs.org/wgbh/aso/tryit/brain/probe.html)
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<div><img src="tmp/5892_CorticalTopographybox!_ecb699f.png" height="100px"><figcaption></figcaption></div>
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<div><img src="tmp/image_d314743.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
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<div><img src="tmp/image1_3f82ace.png" height="100px"><figcaption></figcaption></div>
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## Pathways from the motor cortex to the spinal cord
Indirect pathway: postural
adjustments, especially for
axial and proximal muscles.
Corticospinal tract
(direct pathway)
Corticobulbar tract (indirect pathway. “bulbar” == brainstem nuclei)
Neuroscience 5e Fig. 17.5
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90% of corticaospinal axons at caudal end of medulla cross (decussate, lateral corticospinal tract). 10% remain ipsilaterally (ventral corticospinal tract).
most corticobulbar inputs (except lower face and tongue) terminate bilaterally.
maps: muscle, movement sequences, intention?
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## The corticospinal and corticobulbar tracts
Neuroscience 5e Fig. 17.4
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<div><img src="tmp/Neuroscience5e-Fig-17_42dcdad.jpg" height="100px"><figcaption></figcaption></div>
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Corticobulbar is blue, corticospinal in red. Note that corticospinal cross the midline in the caudal medulla. Corticobulbar is for facial muscles.
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## 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
Neuroscience 5e Box 17A
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<div><img src="tmp/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?
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<div><img src="tmp/Neuroscience5e-Box-17B-0_a572ded.jpg" height="100px"><figcaption></figcaption></div>
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Topographic distribution of microstimulation sites that evoke behavorially relevant movements in a macaque monkey.
Shaded region in map of stimulation sites indicates cortex folded into the anterior bank of the central sulcus.
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## Activity of single upper motor neurons is correlated with muscle movements
Neuroscience 5e Fig. 17.6
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left illustrates spike triggered averaging method for correlating muscle activity with the discharges of single upper motor neurons.
right shows the response of a thumb muscle by a fixed latency to the single spike discharge of a pyramidal tract neuron. This can be used to determine all muscles influenced by a given motor neuron.
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## Purposeful movements resulting from prolonged microstimulation of the primary motor cortex
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<div><img src="tmp/Neuroscience5e-Fig-17.07-0_b1923da.jpg" height="100px"><figcaption></figcaption></div>
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Coordinated movements of hand and mouth after stimulation near the middle of the precentral gyrus towards head (like for eating).
Coordinated movements of hand towards belly as if inspecting an object. Notice clustering of centralized trajectories after many trials instead of just random movements.
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## Directional tuning of an upper motor neuron in the primary motor cortex
Monkey trained to move joystick in response to light
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## Directional tuning of an upper motor neuron in the primary motor cortex
Activity of a single neuron recorded in motor cortex
is dependent on the direction of the future movement.
Neuroscience 5e Fig. 17.8
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<div><img src="tmp/Neuroscience5e-Fig-17.08-2R_256673f.jpg" height="100px"><figcaption></figcaption></div>
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Notice that the neuron is broadly tuned, even with this colored shading.
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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)
Neuroscience 5e Fig. 17.8
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<div><img src="tmp/Neuroscience5e-Fig-17.08-3R_4ac9fb7.jpg" height="100px"><figcaption></figcaption></div>
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<div><img src="tmp/Neuroscience5e-Fig-17.08-4R_1e1c58e.jpg" height="100px"><figcaption></figcaption></div>
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Summing response from a bunch of neurons shows that the direction is better encoded from an ensemble or population of neurons— so that different movement directions/sequences are represented by overlapping and distributed populations of neurons giving rise a series of neuronal population vectors rep all the different directions.
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## Section of pyramidal tracts in monkeys produces loss of independent digit control
Intact (normal)
After section of
corticospinal fibers
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<div><img src="tmp/image_a144227.jpg" height="100px"><figcaption></figcaption></div>
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corticalspinal, lateral dorsal input for control of distal/fine movements of the fingers.
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## Primary motor cortex and the premotor area in human
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## Primary motor cortex and the premotor area in macaque monkey
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<div><img src="tmp/Neuroscience5e-Fig-17.09-0_1d7aeaa.jpg" height="100px"><figcaption></figcaption></div>
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Divisions of the motor cortex in the macaque monkey brain.
lateral premotor and supplementary motor areas are involved in selecting and organizing purposeful movements of the limbs and face.
the frontal eye fields organize voluntary gaze shifts. The cingulate motor areas are involved in expression of emotional somatic behavior.
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## 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.
Note:
thes neurons encode intention to perform a movement rather than just the movement itself.
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## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
[https://www.youtube.com/watch?v=RuK2Y8JojN8](https://www.youtube.com/watch?v=RuK2Y8JojN8)
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<div><img src="tmp/Neuroscience5e-Fig-17.10-1R_46b3807.jpg" height="100px"><figcaption></figcaption></div>
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Indeed a nice way to understand this is by examining portions of the lateral premotor cortex that contain so called mirror neurons that have been focus of a bit of attention over recent years.
peristimulus response histograms
passive observation of human hand interacting with (placing food on) tray and also during motor monkeys own movement to retrieve food
based on Giacomo Rizzolatti et al, 1996
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## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
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<div><img src="tmp/Neuroscience5e-Fig-17.10-2R_cf5432f.jpg" height="100px"><figcaption></figcaption></div>
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does not respond when pliers are used to interact with food.
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## Mirror motor neuron activity in a ventral-anterior sector of the lateral premotor cortex
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<div><img src="tmp/Neuroscience5e-Fig-17.10-3R_10c4215.jpg" height="100px"><figcaption></figcaption></div>
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Also fires when the behavior is executed behind a visual barrier.
Suggests that parts of the premotor cortex play a role in encoding the actions of others.
Studies of this mirror neuron system is an active area of neurosci research and some hypotheses anticipate that this connections in the mirror neuron system could be disrupted in neurodevelopmental disorders such as autism or schizophrenia— but it is still important to note that these are active investigations and hypotheses still be tested.
http://nautil.us/blog/mirror-neurons-are-essential-but-not-in-the-way-you-think
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## 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
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<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.
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## Planning movement sequence without moving activates supplemental motor area (medial premotor area)
Mental rehearsal of finger sequence
Motor cortex
Sensory cortex
Repeated simple finger flexion
Repeating sequence finger-thumb apposition
Supplementary
motor area
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<div><img src="tmp/image1_52fff81.jpg" height="100px"><figcaption></figcaption></div>
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First neuroimaging data
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## Activation of motor areas depend different on behavioral context
Primary motor cortex
Lateral premotor area
Medial motor area
1st key touch
1st key touch
1st key touch
Visual
Cue
Learned
Sequence
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<div><img src="tmp/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.
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## 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)
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## The Babinski sign
[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)
Neuroscience 5e Fig. 17.16
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<div><img src="tmp/Neuroscience5e-Fig-17.16-0_08731e1.jpg" height="100px"><figcaption></figcaption></div>
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normal response on left. Following damage to descending corticospinal pathways stroking sole give abnormal fanning of toes.
Also common in infants during maturation of the descending corticospinal pathways
spinal shock and decr activity deprived of input from motor cortex and brainstem
after several days recovery begins (not fully understood) and includes
-babinski sign
-spasticity (decerebrate rigidity). Cause by removal of suprresive infl by cortex on postural centre of vesitbulaer nuclei and reticular formation.. Rep abnormal incr in the gain of th spinal corste strech reflesxes.
-loss of ability of fine movements.
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## Signs of motor neuron lesions
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<div><img src="tmp/Neuroscience5e-Tab-17.01-0_1f5d45b.jpg" height="100px"><figcaption></figcaption></div>
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## 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
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