Muscle fatigue is an exercise-induced reduction in maximal voluntary muscle force. It may arise not only because of peripheral changes at the level of the muscle, but also because the central nervous system fails to drive the motoneurons adequately. Evidence for “central” fatigue and the neural mechanisms underlying it are reviewed, together with its terminology and the methods used to reveal it. Much data suggest that voluntary activation of human motoneurons and muscle fibers is suboptimal and thus maximal voluntary force is commonly less than true maximal force. Hence, maximal voluntary strength can often be below true maximal muscle force. The technique of twitch interpolation has helped to reveal the changes in drive to motoneurons during fatigue. Voluntary activation usually diminishes during maximal voluntary isometric tasks, that is central fatigue develops, and motor unit firing rates decline. Transcranial magnetic stimulation over the motor cortex during fatiguing exercise has revealed focal cha...

Spinal and Supraspinal Factors in Human Muscle Fatigue

This is a review of the proprioceptive senses generated as a result of our own actions. They include the senses of position and movement of our limbs and trunk, the sense of effort, the sense of force, and the sense of heaviness. Receptors involved in proprioception are located in skin, muscles, and joints. Information about limb position and movement is not generated by individual receptors, but by populations of afferents. Afferent signals generated during a movement are processed to code for endpoint position of a limb. The afferent input is referred to a central body map to determine the location of the limbs in space. Experimental phantom limbs, produced by blocking peripheral nerves, have shown that motor areas in the brain are able to generate conscious sensations of limb displacement and movement in the absence of any sensory input. In the normal limb tendon organs and possibly also muscle spindles contribute to the senses of force and heaviness. Exercise can disturb proprioception, and this has implications for musculoskeletal injuries. Proprioceptive senses, particularly of limb position and movement, deteriorate with age and are associated with an increased risk of falls in the elderly. The more recent information available on proprioception has given a better understanding of the mechanisms underlying these senses as well as providing new insight into a range of clinical conditions.

The Proprioceptive Senses: Their Roles in Signaling Body Shape, Body Position and Movement, and Muscle Force

There is dual tactile innervation of the human hairy skin: in addition to fast-conducting myelinated afferent fibers, there is a system of slow-conducting unmyelinated (C) afferents that respond to light touch. In a unique patient lacking large myelinated afferents, we found that activation of C tactile (CT) afferents produced a faint sensation of pleasant touch. Functional magnetic resonance imaging (fMRI) analysis during CT stimulation showed activation of the insular region, but not of somatosensory areas S1 and S2. These findings identify CT as a system for limbic touch that may underlie emotional, hormonal and affiliative responses to caress-like, skin-to-skin contact between individuals.

Unmyelinated tactile afferents signal touch and project to insular cortex

A new view of pain as a homeostatic emotion

The characteristics of vibration-induced illusory joint movements were studied in healthy human subjects. Unseen by the subject, constant frequency vibration trains applied to the distal tendon of the Triceps or Biceps induced an almost constant velocity illusory movement of the elbow whose direction corresponded to that of a joint rotation stretching the vibrated muscle. Vibration trains of the same duration and frequency applied alternatively to the Biceps and Triceps evoked alternating flexion-extension illusory movements. During successive application of vibration trains at frequencies from 10 to 120 Hz, the perceived velocity of the illustory movements increased progressively from 10 to 70-80 Hz, then decreased from 80 to 120 Hz. The maximal perceived velocity was three times higher during alternating vibration of the Biceps and Triceps than during single muscle stimulation. Unit activity from 15 muscle spindle primary endings and five secondary endings located in Tibialis anterior and Extensor digitorum longus muscles were recorded using microneurography in order to study their responses to tendon vibration and passive and active movements of the ankle. Primary endings were all activated by low amplitude tendon vibration (0.2-0.5 mm) previously used to induce illusory movements of the elbow. The discharge of some was phase-locked with the vibration cycle up to 120 Hz, while others responded one-to-one to the vibration cycle up to 30-50 Hz, then fired in a sub-harmonic manner at higher frequencies. Secondary endings were much less sensitive to low amplitude tendon vibration. Primary and secondary ending responses to ramp and sinusoidal movements of the ankle joint were compared. During the movement, the primary ending discharge frequency was almost constant, while the secondary ending activity progressively increased. During ankle movements the primary ending discharge appeared mainly related to velocity, while some secondary activities seemed related to both movement velocity and joint angle position. Muscle spindle sensory ending responses to active and passive ankle movements stretching the receptor-bearing muscle (plantar flexion) were qualitatively and quantitatively similar. During passive reverse movements (dorsiflexion) most of the sensory endings stopped firing when their muscle shortened. Active muscle shortening (isotonic contraction) modulated differently the muscle spindle sensory ending discharge, which could stop completely, decrease or some times increase during active ankle dorsiflexion. During isometric contraction most of the muscle spindle sensory endings were activated. The characteristics of the vibration-induced illusory movements and the muscle spindle responses to tendon vibration and to active and passive joint movements strengthened the possibility of the contribution of primary endings to kinesthesia, as suggested by several previous works...

Kinaesthetic role of muscle afferents in man, studied by tendon vibration and microneurography

Electromyograms (EMGs) were recorded from surface electrodes over the sternomastoid muscles and averaged in response to brief (0.1 ms) clicks played through headphones. In normal subjects, clicks 85 to 100 dB above our reference (45 dB SPL: close to perceptual threshold for normal subjects for such clicks) evoked reproducible changes in the averaged EMG beginning at a mean latency of 8.2 ms. The earliest potential change, a biphasic positive-negativity (p13-n23), occurred in all subjects and the response recorded from over the muscle on each side was predominantly generated by afferents originating from the ipsilateral ear. Later potentials (n34, p44), present in most but not all subjects, were generated bilaterally after unilateral ear stimulation. The amplitude of the averaged responses increased in direct proportion to the mean level of tonic muscle activation during the recording period. The p13-n23 response was abolished in patients who had undergone selective section of the vestibular nerve but was preserved in subjects with severe sensorineural hearing loss. It is proposed that the p13-n23 response is generated by activation of vestibular afferents, possibly those arising from the saccule, and transmitted via a rapidly conducting oligosynaptic pathway to anterior neck muscles. Conversely, the n34 and p44 potentials do not depend on the integrity of the vestibular nerve and probably originate from cochlear afferents.

https://jnnp.bmj.com/content/jnnp/57/2/190.full.pdf

Myogenic potentials generated by a click-evoked vestibulocollic reflex.

In awake human subjects, neural responses in cutaneous nerves to electrical stimulation were recorded with intrafascicular tungsten micro-electrodes. Changes in the activity of individual fibre groups during blocking procedures were recorded and correlated with simultaneous alterations in the perception of standardized stimuli. Light touch sensibility in hairy skin was mediated by A-beta-gamma fibres, cold and pinprick by A-delta fibres and warmth and dull pain by C fibres.

Fibre function and perception during cutaneous nerve block.

In awake human subjects, neural responses in radial nerves to electrical stimulation were recorded with intrafascicular tungsten microelectrodes. Changes in the activity of individual fibre groups during blocking procedures were recorded and correlated with simultaneous alterations in the perception of standardized stimuli. Light touch sensibility in hairy skin appeared to depend on the integrity of A-beta-gamma fibres, cold and pinprick on A-delta fibres, and warmth and dull pain on C fibres.

https://jnnp.bmj.com/content/jnnp/38/9/865.full.pdf

The effects of a voluntary contraction on the H reflexes of various muscles were quantified to determine whether the reflex responses were sufficiently reproducible to be used in diagnostic studies. During a voluntary contraction, H reflexes could be recorded reliably from tibialis anterior and abductor pollicis brevis, but accurate identification of the onset of the H wave from the on-going background EMG required duplicate averages of multiple responses. During a contraction the H reflex could be obtained at lower stimulus intensities in the forearm flexor muscles than when relaxed, and a more clear separation of the H wave from the M wave was possible. The background contraction abolished the attenuation of reflex amplitude with increasing stimulus repetition rates, such that repetition rates of up to 4 Hz could be used without significant loss of reflex amplitude. There were only small and usually insignificant differences in the latency of the H reflex or its variability when elicited with the forearm muscles relaxed and when flexor carpi radialis was contracting steadily. The reflex latencies of abductor pollicis brevis, tibialis anterior and soleus were compared with F wave latencies for these muscles. The minimal F wave latencies were shorter than the H reflex latencies for abductor pollicis brevis (mean 2.2 ms) and tibialis anterior (mean 1.0 ms) but not for soleus. Comparison of the spread of F wave latencies (F max-F min) suggests that, for soleus, F waves are recorded only from the faster conducting motor units in the pool, presumably those less readily recruited in the H reflex. It was calculated that the distribution of motor conduction velocities responsible for the F waves of abductor pollicis brevis was 8.8 m.s-1. This value underestimates the likely distribution of motor conduction velocities for the thenar muscle by as much as 50%, consistent with the view that F waves rarely occur in slowly conducting motor units, the units of lowest threshold in reflex studies. It is concluded that, for many motoneuron pools, the H reflex and the F wave appear preferentially in different motoneurons, low and high threshold, respectively, and that reflex studies can provide information not available from somatosensory evoked potentials or F wave studies.

The effects of voluntary contraction on the H reflex of human limb muscles.

1. In seven human subjects who were standing without support the sural nerves were stimulated electrically using trains of non-painful stimuli (five pulses at 300 Hz), designed to activate afferents from cutaneous mechanoreceptors. The reflex effects of the stimulus train on different muscles of the ipsilateral and contralateral legs were sought in post-stimulus averages of rectified EMG. Changes in the pattern of reflex influence were investigated when the subjects maintained different postures. 2. Clear reflex responses were seen in ipsilateral tibialis anterior, soleus, biceps femoris and vastus lateralis, but only when the muscles were actively contracting. In each muscle, inhibition was the dominant reflex response within the first 100 ms. In four of the seven subjects, reflex changes were detectable in the contralateral tibialis anterior and soleus, the peak-to-peak modulation within the first 200 ms being 25-50% of that for the homologous ipsilateral muscle. 3. When subjects attempted to stand on a tilted platform, an unstable platform or on one leg with the other flexed, different combinations of muscles were active, involving both flexors and extensors or predominantly flexors or predominantly extensors. In each posture the reflex effects were demonstrable only in the active muscles. 4. With ipsilateral tibialis anterior, there were task-dependent changes in the short-latency components of the EMG response, approximately 60 ms and 80 ms after the stimulus. When seated performing voluntary contractions these components were difficult to define, and when standing on a platform tilted toe-up they were small. When the ipsilateral leg was flexed or when standing on an unstable base, these early components were more prominent in each subject. With contralateral tibialis anterior, the dominant reflex pattern was inhibition when seated and contracting voluntarily, and facilitation during bipedal stance tilted toe-up. These changes in reflex pattern could not be explained by different levels of background contraction. 5. It is concluded that cutaneous mechanoreceptors of the foot have widespread reflex actions on muscles throughout both limbs, particularly the ipsilateral limb, and that the reflex pattern in different muscles and within a single muscle may change dependent on the task that the subject is undertaking. These task-dependent changes indicate plasticity in the expression of cutaneous reflex activity, affecting both short-latency spinal as well as long-latency pathways.

N F Skuse

Papers

Myogenic potentials generated by a click-evoked vestibulocollic reflex.

Fibre function and perception during cutaneous nerve block.

Fibre function and perception during cutaneous nerve block.

The effects of voluntary contraction on the H reflex of human limb muscles.

Task-dependent changes in the responses to low-threshold cutaneous afferent volleys in the human lower limb.