Neuromuscular aspects of Motor Control
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- 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
- 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
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Excitation Contraction Coupling
- Nerve AP
- Neuromuscular junction transmission
- AP transmitted along fibre and down T-tuble
- Ca released from sarcoplasmic reticulum and binds to tropin
- Cross-bridge generate force
- Ca pumped back to SR
- 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:
- muscle excitation - the proportion of muscle fibres in a muscle that are receiving AP from motor neurons.
- 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.
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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