Neuromuscular aspects of Motor Control

1

  • Motor Unit - a motor neuron and muscle fibres innervate
  • Motor Neuron Pool - all alpha motor neurons innervating a muscle
  • Input from cortico-spinal neurons can trigger AP in axon of motor neuron if postsynaptic potential reaches threshold

Synaptic Integration - Post-synaptic potentials

  • Inhibitory - hyperpolarises motor neuron membrane = AP less likely
  • Excitatory - depolarises motor neuron membrane = AP more likely
  • Spatial and temporal summation of post synaptic potentials influence AP triggering

Facilitation and Inhibition can be pre-synaptic
Pre and post excitatory and inhibitory processes occur via:

  • Ionotropic receptors - directly gate ion channels to rapidly influence membrane potentials
  • Metabotropic receptors - indirectly gate ion channels through 2nd messengers for slower neuromodulation

Example of Neuromodulation - Persistant Inward Current

  • Combined activation of ionotropic channels and release of monoanimes facilitate persistant inward currents and self-sustained firing of AP

Why Modulate the Excitability of Motor Neurons?

  • State of high arousal = want motor neurons to be highly responsive
  • When balancing we want less excitable motor neurons = less disruption
  • muscle neurons synapse with muscle fibres at neuromuscular junction
  • When an endplate potential reaches synapse then an AP may be generated in muscle fibre = contraction

Measuring Signal

  1. Electromyography (EMG) - records AP of all/parts of muscle, uses 2 electrodes on skin (surface) or inserted in muscle (intramuscular)
    Limitations of surface:
  • Muscles close together = record other muscles
  • If muscle is deep = only records top muscles
    Strengths of intramuscular:
  • Can identify single neuron

Size Principle Innervation Number

  • Higher recruitment threshold = higher innovation no. fibres being recruited (type 2) - not consistent/linear relationship

Motor Unit Number
No. motor units within a muscle:

  • varies between muscles
  • Muscle force can be increased through increased stimulation frequency
  • Involuntary contractions - discharge rate

Motor Unit Number Estimation (MUNE)

  • Compound muscle AP - M-wave response to max stimulation of nerve
  • Single motor unit AP - single motor unit response to stimulation
  • MUNE = CMAP/S-MUP

A more inhibited motor neuron pool:

  • Fewer neurons likely to discharge less frequently
  • muscle activation less for given amount of voluntary central drive
    A more excited motor neuron pool:
  • More motor neurons likely to discharge and frequently
  • Muscle activation greater for given amount of voluntary central drive

Evaluating Excitability of Neuron Pool

H-Reflex

  • Only in limited no.muscles more when relaxed
  • Influenced by presynaptic inhibition from afferent neurons

CMEPs

  • Cervicomedullary motor evoked potentials - stimulates cortico-spinal neurons directly
  • Possible during contractions at rest
  • Painful
  • Hard to evoke response in some p's

Neural Response to Fatigue - Fatigue and Force Output

  • Repeated contraction reduce the force output for a given level of central drive

Neural Response to Sustained Submax Contraction (EMG)

  • Additional recruitment to maintain force output as other motor units fatigue
  • Motor units recruits at lower force threshold as fatigue sets

Discharge rate coding and fatigue

  • Motor units recruited initially reduce their discharge rate with fatigue
  • Relaxation rates of muscle slow with fatigue
  • 'muscle wisdom' - decreasing discharge rate takes advanatge of slower relaxation rates to maintain force output economically (Enoka)

Reduced Discharge Rates Controlled by Inhibition of Motor Neurons

  • Reduction in CMEPs
  • Indicates inhibition of motor neuron pool
  • likely driving reduction in discharge rate

2

Excitation Contraction Coupling

  1. Nerve AP
  2. Neuromuscular junction transmission
  3. AP transmitted along fibre and down T-tuble
  4. Ca released from sarcoplasmic reticulum and binds to tropin
  5. Cross-bridge generate force
  6. Ca pumped back to SR
  7. Cross-bridges detach (relax)
  • Each step in excitation-contraction coupling is rate limited.
  • e.g. excitation (nerve impulse and Ca release) takes ~5-10milliseconds
  • e.g. contraction and relaxation ~100 milliseconds
  • Therefore, mechanical force generation lags behind excitation process

Time Delay Occurs:

  1. muscle excitation - the proportion of muscle fibres in a muscle that are receiving AP from motor neurons.
  2. Muscle activation - the proportion of available actin and myosin cross bridges that are bound (generating tension)
  • Tdeact = time constant describing lag between excitation of a muscle turning off and deactivation of muscle
    -Tact = time constant describing lag between excitation of a muscle and activation

Muscle Twitch vs Tetanus

  • Higher rates of stimulation cause summation of force and hence a tetanus (fused force)

Active Tension in Skeletal Muscle

  • The number of active actin and myosin cross-bridges determines the amount of active force production.
  • Sarcomeres sit in series and distribute strain (length change) across muscle fibres
  • Resting fibre lengths of a muscle relates to how many sarcomeres are in series in muscle fibres

Muscle Architecture

Muscle Architecture - fibres in series vs in parallel

  • In series: length changes sum, force equal
  • In parallel: length changes equal, forces sum

Muscle Architecture - resting length

  • Resting fibre length influences active force production by the whole muscle
  • Active and passive force-length relations vary between muscles
    Muscles with Long Resting Fibre Lengths:
  • Have more sarcomeres
  • maintain greater force production throughout their ROM
  • Well suited to producing large length changes and mechanical work
    work = force x displacement

Muscles with long fibres and large PCSA

  • Metabolically expensive
  • A shorter muscle with same no. fibres in parallel (PCSA) can produce same force by activating a smaller vol of muscle = economical.

Muscle Architecture - pennation angle

  • Pennation allows a greater no. fibres to be packed into a vol of muscle.
  • Fibre length shortens
  • Results in greater PCSA without having to increase muscle vol (+mass)

Muscle structure and FV relationships

  • Shorter fibres have lower absolute max shortening speed than longer fibres.
  • However, they may produce more force (greater PCSA)
    Force-length-vel and tension
  • length and vel influence force

Tendon Architecture influences stiffness

  • Increasing tendon stiffness/width increases stiffness
  • Increase tendon length will decrease stiffness/increase compliance

Structure-function relationship (benefits of a long compliant achillies tendon)

  • Energy storage and return- reduces work muscle fibres have to do
  • Allows muscles to work on a economical part of their force-vel relationship
  • can protect muscle fibres from rapid stretch
    Cons of long compliant achillies tendon
  • hard to generate external work

Structure-function relationship - proximal - to - distal leg muscle

  • proximal muscles have long muscle fibres that facilitate external work production
  • Distal muscles have longer compliant tendons that help improve steady running economy.
  • Hip extensors have large flat tendons, making them stiff.

3

Length-tension relationship

  • Muscle fibre length changes with joint angle
  • length-tension predicts max force generated
    Muscle Moment Arms
  • May change with joint angle

Torque Output and Muscle Design

  • Muscle force output and moment arm relationships both contribute to resultant output by a muscle at a given joint angle.
    Muscle Moment arms - linear to ang kinematics
  • change in muscle length (k) = theta x r1

Moment arm vs Muscle length

  • Longer fibres required for longer moment arm
  • Longer muscle fibres permit greater ROM
  • Combination allows large ang displacements over which forces can be produce
    -Long fibred muscles must have a large vol to have PCSA
    Summary of Short Moment Arms
  • small moment arms help muscle remain on a favourable part of the force-vel relationship, while producing fast joint rotations.
  • Require more force for given torque
  • Narrows force-angle relationship

Moment Arms and Effective Mechanical Advantage (EMA)

  • Mechanical advantage - ratio of input to output moment arms
    EMA = External load (output force)/muscle moment arm (input force)

Bi-Articular Muscle Function

  • Bi-articular muscles act across 2 joint
  • This can be seen as counterintuitive for some movements
  • Using bi-articular muscles, power can be transferred from large hip muscle to the knee and ankle

Elastic Power amp and stretch-shortening cycles

  • Jump height = take off vel squared/2g
  • Increased take off vel requires a large impulse force in a short time (force x time)

Power Amplification

  • Muscle shortens to stretch tendon while joint at low vel
  • Joint movement is then powered by rapid release of energy stored in tendon
  • Tendons aren't constrained by force-vel relationship = release energy faster than muscle can generate work

Power Amplification: where's the 'catch'?

  • Some insects have a physical 'catch' mechanism - a latch, it can be used to block motion
  • Frogs manipulate the muscles mechanical advantage - R/r
  • High value early = tendon stretch
  • Low value late = tendon recoil

What is a stretch-shortening cycle?

  • Muscle actively lengthen then shortens
  • eccentric-concentric sequence