ACTIVE SENSING Lecture 2: Motor systems Motor systems 1 Basic muscle operation and control Motor systems 2 Muscles have one trick: contraction flexion joint extension Motor systems 3 Muscles have one trick: contraction joint extension Motor systems 4 Muscles have one trick: contraction flexion joint extension Motor systems 5 Muscles have one trick: contraction flexion joint Motor systems 6 Muscles have one trick: contraction Extensor Fext Flexor Fflex flexion joint Torquenet = torqueflex - torqueext extension = Fflex dflex - Fext dext Motor systems 7 Elbow rotation Motor systems 8 Muscles Muscles consist of arrays of individual fibers, muscle fibers, covered by a membrane. Fibers are constituted from smaller contractile units called myofibrils. •Muscles have active state, when they contract (shorten) and inactive state when they lengthen. •Fibers that work together are synergistic. Antagonistic fibers work in opposition to each other. •Muscle fibers (motor units) differ in dynamic range: slow: slow, long duration, low tension fast fatigue-resistant: fast, medium duration, medium tension fast fatigable: very fast, short duration, high tension Motor systems Inactive state active state 9 Twitch, tetanic force, fatigability Slow Fast fatigue-resistant Fast fatigable Twitch Tetanic force Fatigability Motor systems 10 Motor Units Each muscle fiber is innervated by only a single axon, but a single axon innervates several muscle fibers. The motor neuron and the muscle fibers it innervates constitute a motor unit. Constituents of Motor Units Motor neurons Ventral horn of spinal cord The size of motor units varies with the precision of movement control: Leg: >1000 muscle fibers per unit Eye: <100 muscle fibers per unit Muscle fibers Motor systems 11 Excitation Contraction Coupling Phase 1: Firing of Motor Neuron Input to motorneuron Phase 2: Release of Neurotransmitter Motor systems 12 Excitation Contraction Coupling Phase 1: Firing of Motor Neuron Input to motorneuron Phase 2: Release of Neurotransmitter Phase 3: Muscle contraction Motor systems 13 Open-loop system Information flows in one direction (from neurons to muscles Input to motorneuron Motor systems 14 Open-loop system Information flows in one direction (from neurons to muscles Input to motorneuron Closed-loop system Information flows in a closed loop: from neurons to muscles and from muscles to neurons What kind of information ? Motor systems 15 Closed-loop system The direct feedback from muscles and joints is mediated by proprioceptive signals Proprioceptive receptor types Name: Muscle spindle receptors Sensitive to: muscle length Motor systems Golgi tendon organs Joint receptors muscle tension Flexion, extension 16 Proprioceptive receptor types Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Location: Fleshy part of the muscle Between muscle and tendon Joint capsule Parallel to muscle fibers Serial to muscle fibers Between bones Name: Motor systems 17 Proprioceptive receptor types Name: Sensitive to: Motor systems Muscle spindle receptors Golgi tendon organs Joint receptors muscle length muscle tension Flexion, extension 18 What proprioceptors encode? Motor systems 19 Proprioceptive receptor types Name: Sensitive to: Muscle spindle receptors Golgi tendon organs Joint receptors muscle length muscle tension Flexion, extension From Arthur Prochazka, University of Alberta Motor systems 20 Proprioceptive receptor types Name: Sensitive to: Muscle spindle receptors Golgi tendon organs Joint receptors muscle length muscle tension Flexion, extension force Encode: Motor systems f = k1 F 21 Proprioceptive receptor types Name: Sensitive to: Encode: Motor systems Muscle spindle receptors Golgi tendon organs Joint receptors muscle length muscle tension Flexion, extension Length + velocity force angle f = k1L + k2 V 0.6 f = k1 F f = k1 q 22 Proprioceptive receptor types Name: Sensitive to: Encode: Muscle spindle receptors Golgi tendon organs Joint receptors muscle length muscle tension Flexion, extension Length + velocity force angle f = k1L + k2 V 0.6 f = k1 F f = k1 q q Motor systems q q 23 PID control Proportional Integral Derivative Motor systems (to the controlled variable) (of the controlled variable) (of the controlled variable) Present q Past q Future q 24 Negative feedback loop Characteristic: The effect of a perturbation is in opposite direction Requirement: The cumulative sign along the loop is negative Function: Can keep stable fixed points Motor systems 25 Positive feedback loop Characteristic: The effect of a perturbation is in the same direction Requirement: The cumulative sign along the loop is positive Function: amplifies perturbations Motor systems 26 Reflexes probing closed-loop control Motor systems 27 The stretch reflex probes the control function of muscle spindles Motor systems 28 Is the loop positive or negative? The stroke stretches the spindle As a result the muscle contracts The result opposes the perturbation => negative FB loop Motor systems 29 Is the anatomical loop negative? Muscle spindle excites the motor neuron Motor neuron excites muscle fibers Muscle contraction suppresses spindle response Motor systems 30 What about the flexor muscles? Positive or negative loop? What is the underlying circuit? Motor systems 31 Pain reflex Positive or negative? What is the underlying circuit? Motor systems 32