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Neural and Sensory. Efferent innervation Neuromuscular junction a -, g -motorneurons Afferent innervation Muscle spindle Golgi tendon organ. Overview. Motor Unit Motorneuron + many muscle fibers Cholinergic synapse Spindle Enapsulated , specialized muscle-like cells
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Neural and Sensory • Efferent innervation • Neuromuscular junction • a-, g-motorneurons • Afferent innervation • Muscle spindle • Golgi tendon organ
Overview • Motor Unit • Motorneuron + many muscle fibers • Cholinergic synapse • Spindle • Enapsulated, specialized muscle-like cells • Combined length-velocity proprioception • Tendon organ • Nerve endings woven through collagen • Active tension mechanorecption
Dorsal/posterior Ventral/anterior MN density Efferent Innervation • a-motorneuron • 100 um cell body/soma • Ventral horn of spinal cord • Columnar organization by muscle • 8-18 um diameter axon • 60-120 m/s conduction velocity • Terminal branches within target muscle • Dendritic tree • 10,000+ inputs per MN • Excitatory glutamic synapses • Inhibitory glycinergic/GABAergic synapses Cervical Arm/hand Thoracic Lumbar Sacral Leg/foot
Neuromuscular junction • Motor endings • Synaptic vessicles • Mitochondria • Synaptic cleft • Primary • Secondary invaginations • Muscle • Post-synaptic density • Mitochondria deHarven & Coers, 1959
By SEM Sohn & al., 1999
Synaptic vessicle anchoring • SV anchored by “pegs and ladders” Harlow & al., 2000
Synaptic structure • Complementary • Presynaptic active zone • Postsynaptic density/receptor • ECM coordination • NerveagrinMuSK • MuscleLam 421/521VGCC • Electrical coordination • Nerve: ubiquitinylation preserves synaptotagmin • Muscle: depolarization degrades AChR
Presynaptic side • Docking site • SNAP-25 • Syntaxin (T-SNARE) • Synaptic vesicle • Synaptobrevin (V-SNARE) • Synaptotagmin • Docking trimer: t/v-SNARE+SNAP • Calcium sensitivity: synaptotagmin
Presynaptic active zone De Wit & al., 2009
Presynaptic active zone Garner 2002
Postsynaptic side • Post-synaptic density • MuSK/LRP4 receptor • Rapsyn/Dok-7 adapter • Src kinase • Acetylcholine receptor • Junctional folds • Sodium channels • Acetylcholinesterase
Post-synaptic density Shi & al., 2012
Afferent innervation • Afferent • Sensory • Dorsal root ganglion • Synapses in SC • Spindle • Muscle-likecells • Mechanosensitivenerve ending
Muscle Spindle • Capsule • Polar regions • g-motorneurons • Striated contractile proteins • Equatorial region • Primary afferent (Ia) • Secondary afferent (II) • Non-contractile • Nuclear bag & chain Barker, 1948
Spindle afferents Barker, 1948
Spindle efferents • Dynamic bag/bag1 • Usually only influences the primary ending • Increases velocity gain • Static bag/bag2 • Offset both primary & secondary • Reduces length sensitivity of primary • Chain • Take up slack • Increase length sensitivity of secondary
Spindle Mechanics • Equatorial stretch induces depolarization • Windings amplify length change • Stretch imposed from outside • Stretch imposed by contraction of poles • Accommodation • Viscoelastic creep/relaxation • Primary fast: velocity • Secondary slow: length
Stretch response Primary ending Secondary ending Muscle length Afferent firing rate Houk & al, 1981
Stretch Reflex • Ia afferent“homonymous” motorneuron • Servo-like length control • Tendon tap • Vibration illusions • IaIa interneuron • Inhibit antagonist(s) • Mechanical equivalence • Myotatic unit • IIdorsal horn interneurons • Distributed, slower excitation
Golgi tendon organ • Free nerve endings embedded in capsule at muscle-tendon interface • Group Ib Nitatori, 1988
GTO response • MTJ tension depolarizes afferent Muscle length GTO discharge rate Muscle force Crago & al., 1982
GTO Reflex • IbIb interneuron • Delayed homonymous inhibition • Distributed excitation/inhibition • “Interaction” torques • Locomotor pattern integration • Polysynaptic pathways are subject to extra modulation
Summary • a-motorneuron efferent forms coordinated NMJ • Presynaptic active zone for ACh release • Postsynaptic densities for ACh transduction • Monosynaptic stretch reflex • Combined length and velocity signal • Intrafusal fiber mechanics mimic extrafusal fibers • Polysynaptic GTO reflex • Force feedback • Distributed