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Bear: Neuroscience: Exploring the Brain 3e • Chapter 13: Spinal Control of Movement Christopher Reeve C1-C2 level injury 1 The Hierarchical Control of Movement Fig 10-2 Many components, but functions as a WHOLE • Motor Programs – Motor system: Muscles and neurons that control muscles – Role: Generation of coordinated movements – Parts of motor control • Spinal cord coordinated muscle contraction • Brain motor programs in spinal cord The Somatic Motor System • Types of Muscles – Smooth: digestive tract, arteries, related structures – Striated: Cardiac (heart – sort of) and skeletal (bulk of body muscle mass) 2 The Somatic Motor System • Somatic Musculature – Axial muscles: Trunk movement – Proximal muscles: Shoulder, elbow, pelvis, knee movement – Distal muscles: Hands, feet, digits (fingers and toes) movement The Somatic Motor System • The Lower Motor Neuron – Lower motor neuron: Innervated by ventral horn of spinal cord – Upper motor neuron: Supplies input to the spinal cord Spinal cord injuries •Motorneurons below the injury remain intact. •Motor cortex commands do not reach muscles and muscles atrophy. •Electrodes can artificially activate muscles and prevent atrophy UPPER MOTOR NEURON SYNDROME DAMAGE TO DESENDING PATHWAYS Damage to the pathways driving the motor neurons •Spasticity • TONE AND REFLEXES INCREASED Spastic cerebral palsy for example LOWER MOTOR NEURON SYNDROME - DAMAGE DIRECT TO MOTOR NEURONS Diseases or lesions at the level of the motorneuron or its axon •Atrophy- loss of muscle volume DECREASED TONE AND REFLEXES Poliomyelitis for example 3 The Somatic Motor System • Alpha Motor Neurons Final Common Pathway • Motor unit – one motor neuron and muscle fibers • Pool – single muscle – Two kinds of lower motor neurons • Alpha • Gamma (maintains muscle tension) Guillain Barre syndrome (ghee yan bah ray) • Syndrome not disease (unclear what disease) • • • • • • • Paralysis (can be total) Attacks Schwann cells, then axons Autoimmune Similar to MS in CNS 70% recovery! Why???? Following vaccine (rabies, swine flu) 1 case per million 1 death per 20 million (normal?) The Somatic Motor System • Graded Control of Muscle Contraction by Alpha Motor Neurons – Varying firing rate of motor neurons • (temporal summation) – Recruit additional synergistic motor units – Recruit smallest first, largest last (why small fine movements are not possible under great load) 4 The Somatic Motor System • THREE Inputs to Alpha Motor Neurons Feedback on muscle length (dorsal root ganglia) The Somatic Motor System • Types of Motor Units – Red muscle fibers: Large number of mitochondria and enzymes, slow to contract, can sustain contraction – White muscle fibers: Few mitochondria, anaerobic metabolism, contract and fatigue rapidly (but POWERFUL - escape) – Fast motor units: Rapidly fatiguing white fibers – Slow motor units: Slowly fatiguing red fibers The Somatic Motor System • PLASTIC • Neuromuscular Matchmaking – Experiment: John Eccles • Are muscle properties due to innervating nerve characteristics? • Alternate nerve input – Switch in muscle phenotype (physical characteristics) ACTIVITY – Hypertrophy: Exaggerated growth of muscle fibers – Atrophy: Degeneration of muscle fibers 5 FAST twitch (fatigue rapidly – white) SLOW twitch (fatigue slow – red) Together “calf muscle” charley horse Forced change in input – switch phenotype (physical characteristics) 30-60/sec bursts - 10-20/sec steady Excitation-Contraction Coupling • Muscle contraction – Alpha motor neurons release Ach • Innervate muscle fibers – ACh produces large EPSP in muscle fibers (via nicotinic Ach receptors – EPSP evokes action potential – Action potential (excitation) triggers Ca2+ release, leads to fiber contraction – Relaxation, Ca2+ levels lowered by organelle reuptake Excitation-Contraction Coupling • Muscle Fiber Structure Myofibrils contractile fibers Sarcoplasmic reticulum (endoplasmic) CALCIUM 6 Excitation-Contraction Coupling • The Molecular Basis of Muscle Contraction – Z lines: Division of myofibril into segments by disks – Sarcomere: Two Z lines and myofibril – CONTRACTILE UNIT Thin filaments ACTIN: Series of bristles Thick filaments MYOSIN: Between/among thin filaments Myosin sites blocked by troponin Sliding-filament model: Binding of Ca2+ to troponin causes myosin to bind to actin Myosin heads pivot, cause filaments to slide RELAXED/CONTRACTED twitch 7 Excitation-Contraction Coupling • Steps in Excitation-Contraction Coupling – Excitation: Action potential, ACh release, EPSP, action potential in muscle fiber, depolarization – Contraction: Ca2+, myosin binds actin, myosin pivots and disengages, cycle continues IF Ca2+ and ATP present – Relaxation: EPSP end, resting potential, Ca2+ by ATP driven pump, myosin binding actin covered Steps in Excitation-Contraction Coupling page 436 Rigor mortis • No ATP (myosin uses ATP to disengage) • Calcium pumps are driven by ATP • Somewhat permanent myosin/actin binding 8 Excitation-Contraction Coupling • Spinal Control of Motor Units – First source: Sensory feedback from muscle Excitation-Contraction Coupling • The Myotatic Reflex – Stretch reflex: Muscle pulled tendency to pull back – Feedback loop – Discharge rate of sensory axons: Related to muscle length – Monosynaptic (one synapse sensory/motor) – Example: knee-jerk reflex • The Myotatic Reflex (kneecap tendon stretches quad muscle, triggers contraction) WHY FASTER? 9 Monosynaptic reflexes bypass the brain???. Myasthenia Gravis • • • • • • • Severe muscle weakness (but fluctuates) 1 in 10,000 M & F Autoimmune disease (Thymus?) Attacks nicotinic Ach receptors Blocks and degrades receptors Treatment – inhibit AChE!! (carefully) Treatment – suppress immune system Duchenne Muscular Dystrophy • Genetic – Duchenne 1 in 3500 • ONLY males, so X-linked (single X is enough) X region codes for protein “dystrophin” In MD, no mRNA for this cytosketal protein Muscles tears WHY normal phenotype for early life? Could virus help????? (gene therapy) Could stem cells help? 10 Excitation-Contraction Coupling • Gamma Motor Neurons – Muscle spindle (stretch receptor) • Intrafusal fibers: gamma • Extrafusal fibers: alpha Excitation-Contraction Coupling • Gamma Loop – GAMMA LOOP • Gamma motor neuron intrafusal muscle fiber Ia afferent axon alpha motor neuron -> extrafusal muscle fiber Excitation-Contraction Coupling • Proprioception from Golgi Tendon Organs • Golgi tendon monitors muscle tension • (spindle muscle length) 11 Excitation-Contraction Coupling • Proprioception from the joints, too – Proprioceptive axons in collective joint tissues – Respond to angle, direction and velocity of movement in a joint – Information from joint receptors: Combined with muscle spindle, Golgi tendon organs, skin receptors Excitation-Contraction Coupling • Spinal Interneurons – Synaptic inputs • Primary sensory axons (Ia) • Descending axons from brain • Collaterals of lower motor neuron axons Excitation-Contraction Coupling • Inhibitory Input – Reciprocal inhibition: Contraction of one muscle set accompanied by relaxation of antagonist muscle • Example: Myotatic reflex 12 Excitation-Contraction Coupling • Excitatory Input from Interneurons too – Flexor reflex: – Complex reflex arc used to withdraw limb from aversive stimulus Excitation-Contraction Coupling • Excitatory Input Crossed-extensor reflex: Activation of extensor muscles and inhibition of flexors on opposite side Excitation-Contraction Coupling • The Generation of Spinal Motor Programs for Walking 13 Excitation-Contraction Coupling • The Generation of Spinal Motor Programs for Walking Central pattern generators – Membrane depolarizes – Na & Ca in (NMDA) – Ca activates K channels – K LEAVES cell – Membrane hyperpolarizes – Ca stops – K channels close – repeat • Spinal control of movement – Different levels of analysis – Sensation and movement linked • Direct feedback – Intricate network of circuits EXCITE CONTRACT RELAX UNDERSTAND steps on Pg 436 14 ventral and lateral tracts– motor - spinal cord to muscles cell bodies are in gray matter of spinal cord dorsal fibers – sensory, periphery to Spinal Cord cell bodies are outside -in dorsal root ganglia The Motor Neurons -and interneurons located in gray matter of ventral “horn” Lateral corticospinal tract synapses on motor neurons that move muscles in limbs and digits in contralateral side. Ventral corticospinal tract synapses on motor neurons of midline muscles (trunk) in ipsi side. 15