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motor unit
single alpha motor neuron and the muscle fibers it innervates
function of motor units
Coordinate contraction of different muscle groups
location of motor neuron bodies
spinal cord ventral horn
motor neuron pool definition
all the motor neurons innervating a single muscle
alpha motor neurons
Large neurons that innervate extrafusal fibers to generate muscle contraction
gamma motor neurons
Small neurons that innervate intrafusal fibers in muscle spindles
lower motor neuron innervation of distal muscles
lateral ventral horn; regulate control of limbs
Lateral ventral horn much larger in the cervical and lumbar enlargement
lower motor neuron innervation of proximal muscles
medial ventral horn; involved in postural control
the size principle of motor unit recruitment
as input strength on alpha motor neurons increases, smaller motor neurons are recruited and fire action potentials before larger motor neurons
Slow motor units (S)
Small α-MNs innervate small, weak muscle fibers very resistant to fatigue are recruited by low amount of synaptic input
Important for postural control and standing
Fast fatigue-resistant motor units (FR)
Intermediate α-MNs innervate motor units with twice the force of slow fibers
somewhat resistant to fatigue
Recruited by moderate synaptic input
Fast fatigable motor units (FF)
Largest α-MNs innervate motor units with high force, but fatigue rapidly
Recruited by high levels of input
Important for running, jumping
inputs that lower MNs integrate
Supraspinal long-range neuronal projections, local intraspinal projections, and sensory projections
locations of supraspinal long-range neuronal projections
Motor cortex and brainstem
locations of local intraspinal projections
Interneurons within spinal cord
contralateral/commisural, intersegmental/propriospinal, and intrasegmental
locations of sensory projections
DRG sensory neurons; deep tendon and other reflexes
function of intrafusal fiber contraction
innervated by gamma MNs; keep muscle spindle tensed over wide range of muscle lengths
muscle spindle gain control
Allows spindle to accurately convey stretch information at any muscle length
alpha gamma coactivation
Most motor commands activate both alpha and gamma motor neurons
Central Pattern Generators
Spinal/brainstem networks of interconnected excitatory and inhibitory neurons that produce oscillating, rhythmic output
play an important part in breathing and repetitive locomotive responses
upper motor neurons
Glutamatergic pyramidal neuron bodies in layer 5 of M1; One of largest somal sizes in the brain
Send topographically organized axonal projections to subcortical sites
Upper MNs controlling proximal and axial muscles
Descend bilaterally in medial tracts, innervate multiple segments, and aid in posture and balance
Upper MNs controlling distal muscles
Descend contralaterally in lateral tracts, innervate few segments, and aid in voluntary, skilled movements
location of upper MN projections
descend through internal capsule, with collateral branches in striatum, midbrain, thalamus
corticobulbar tract
innervates targets in pons and medulla to control face movements
lateral corticospinal tract
most of corticospinal axons cross to the contralateral side at the medullary/pyramidal decussation
ventral corticospinal tract
some axons go down ipsilateral side
muscle field
2-4 muscles activated by single upper MN
Concurrent activation of “ensembles” of upper motor neurons encode behaviorally relevant coordinated movements
Upper MN lesions
Weakness; mild/no atrophy; hyperactive deep reflexes after initial period of spinal shock; spasticity; Babinski’s sign and clonus; widespread distribution of impairment in body regions; impairment of fine voluntary movements
lower MN lesions
Weakness or paralysis; severe atrophy; Hypoactive superficial and deep reflexes; initial signs and symptoms persist; fasciculations and fibrillations; geographic distribution of impairment; impairments of reflexive and gross and/or fine voluntary movements
Basal Ganglia Motor Functions
motor initiation; activated by decision to move; involved in starting selected motor programs; inhibits motor programs that aren’t selected
basal ganglia anatomy
Dorsal striatum, ventral striatum, globus pallidus, subthalamic nucleus (STN), and substantia nigra
components of dorsal striatum
Caudate and putamen
components of ventral striatum
Nucleus accumbens
components of globus pallidus
external segment (GPe) and internal segment (GPi)
components of substantia nigra
pars reticulata (SNr) and pars compacta (SNc)
BG input
Cortico-striatal projection and SNc
Cortico-striatal Projection of BG
from motor planning areas in frontal cortex; glutamatergic
SNc Projection of BG
dopaminergic
Medium Spiny Neurons
receive glutamatergic input on dendritic spines from cortical neurons and dopaminergic input from the SNc; send GABAergic projections to GPe, GPi, and SNr
BG Output
GP and SNr
GP projections
External segment to STN; internal segment to VA/VL thalamus
SNr projections
superior colliculus
BG body movement loop
Motor/premotor/somatosensory cortex → putamen → lateral GPi → Ventral lateral and ventral anterior nuclei
BG oculomotor loop
Posterior parietal/prefrontal cortex → caudate → GPi/SNr → Mediodorsal and ventral anterior nuclei
BG prefrontal loop
Dorsolateral PFC → Anterior caudate → GPi/SNr → Mediodorsal and ventral anterior nuclei
BG limbic loop
Amygdal/hippocampus/orbitofrontal/anterior cingulate/temporal cortex → ventral striatum → ventral pallidum → Mediodorsal nucleus
disinhibitory circuit of BG direct pathway
GP is tonically active
Activation of MSNs releases tonic inhibition of thalamus → upper MN activation and movement initiation
BG direct pathway movement facilitation
MSN → GPi → Thalamus → Cortex
facilitates movement initiation
indirect BG pathway
MSN → inhibits GPe → excites STN → excites GPi → inhibits thalamus → inhibits movement
function of indirect BG pathway
Antagonizes direct pathway; suppresses movement
focused selection hypothesis
Direct pathway promotes intended motor program initiation
Indirect pathway inhibits broad set of unintended motor programs
modulatory effect of D1
D1 receptors activate cAMP and enhance direct pathway activity
Modulatory effects of D2
D2 receptors decrease cAMP and inhibit indirect pathway activity
GPi function
inhibits thalamus
GPe function
inhibits GPi and STN
STN function
excites GPi
VA/VL complex of thalamus function
excites motor cortex, premotor cortex, and supplementary motor areas
direct pathway with dopamine input
glutamate from cerebral cortex and D1 excite MSNs → increased MSN activity → increased GPi inhibition → disinhibition of VA/VL → increased activity in VA/VL complex → increased frontal cortex activity
indirect pathway with dopamine input
D2 receptors inhibit MSNs → decreased GPe inhibition → decreased STN inhibition → decreased GPi activation → decreased thalamic inhibition → increased movement
hypokinetic direct and indirect pathways of BG
degenerated SN → decreased dopamine → increased GPe → increased STN inhibition → increased activation of GPi → increased thalamic inhibition → reduced excitation of frontal cortex
decreased inhibition of GPi by D1 → increased thalamic inhibition → reduced excitation of frontal cortex
hyperkinetic direct and indirect pathways of BG
MSNs of indirect pathway degenerate → decreased GPe inhibition → increased STN inhibition → increased GPi inhibition by GPe and decreased GPi activation by STN → decreased tonic inhibition of thalamus → increased excitation of frontal cortex
function of cerebellum
coordination on ongoing movement and modifies descending motor commands
cerebrocerebellum
guides fine motor control movements such as upper limb movements and fine motor control of hands
spinocerebellum
controls medial muscles, coordinating limb/trunk movements, adjusting posture, and error correction of movement
vestibulocerebellum
maintaining balance, spatial orientation, and visual tracking
contralateral cerebellar input
Pons, Inferior Olive, and Middle Cerebellar Peduncle
ipsilateral cerebellar input
Spinocerebellar Tract, accessory Cuneate Nucleus, and vestibular nuclei
cerebellar output to contralateral forebrain via cerebrocerebellum
Cerebrocerebellum → dentate nucleus → premotor cortex (motor planning)
cerebellar output to contralateral forebrain via spinocerebellum
Spino cerebellum → interposed and fastigial nuclei → motor cortex and brainstem (motor execution)
cerebellar output to contralateral forebrain via vestibulocerebellum
Vestibulocerebellum → Vestibular nuclei → Lower motor neurons in spinal cord and brainstem (balance and VOR)
cerebellar output to ipsilateral brainstem via superior colliculus
Cerebellar cortex → fastigial DCN → inferior cerebellar peduncle → Superior colliculus → Reticular formation → anterior medial white matter of spinal cord → lower motor neurons in medial ventral horn
cerebellar output to ipsilateral brainstem via reticular formation
Cerebellar cortex → fastigial DCN → inferior cerebellar peduncle → reticular formation → anterior medial white matter of spinal cord → lower motor neurons in medial ventral horn
cerebellar output to ipsilateral brainstem via vestibular nuclei
Cerebellar cortex → inferior cerebellar peduncle → vestibular nuclei → anterior medial white matter of spinal cord → lower motor neurons in medial ventral horn
cerebellar output to contralateral brainstem
contralateral cerebellar cortex → dentate DCN → superior cerebellar peduncle → crosses midline → Superior colliculus → Reticular formation → anterior medial white matter of spinal cord → lower motor neurons in medial ventral horn
purkinje cell
GABAergic; large soma and extensive spines; provides output from cerebellar cortex to deep nuclei
granule cell
Glutamatergic; small, many neuronal subtypes
mossy fiber
Afferent axons to cerebellum from all sources except for inferior olivary nuclei; enter via the inferior middle cerebellar peduncles
parallel fiber
bifurcated axons of cerebellar granule cells that extend along molecular layer of cerebellar cortex where they synapse on dendritic spines of Purkinje cells.
climbing fiber
Axons that originate in inferior olivary nuclei, ascend through inferior cerebellar peduncle, and make terminal arborizations that invest proximal dendritic trees of Purkinje cells
cerebellar mossy fiber input
Pontine nuclei send mossy fiber projections, Excitatory synapse on granule cells, Granule cell parallel fibers synapse on Purkinje cell (PC) dendritic spines, and trigger simple spikes
cerebellar climbing fiber input
Inferior olivary neurons send climbing fiber projections, excitatory synapse on PC dendrites, and trigger complex spikes that modify parallel fiber-PC efficacy. complex spikes first fire heavily then produce an AP at 1-2hz for short time
cerebellar loops as a comparator
If cortical inhibitory loop and deep excitatory loop activity balanced, then motor commands maintained
purkinje cell LTD
If error detected then climbing fibers trigger complex spikes, climbing fiber activity causes Ca2+ influx into Purkinje cell leading to LTD of parallel fiber synapses, and weakening of cortical inhibitory loop enhances DCN output, which is relayed to motor cortex to alter motor output
cerebellar ataxia
involves irregular movements that lack coordination; failure in correcting errors during a motor command
dermatome
DRG sensory neurons
where somatosensory reception begins
Meissner corpuscles
detects motion across skin; fast adapting
Merkel cells
detects points, edges, curves; slow adapting
Pacinian corpuscle
detects skin vibration; fast adapting; large receptive field
qualities of rapidly adapting receptors
qualities of slowly adapting receptors
qualities affecting sensory perception
mechanosensory transduction
Muscle Spindle
Golgi Tendon Organs
Ruffini Corpuscle
axon of gamma motor neuron
group Ia afferent axons