Brian McKeon

The afferent volleys responsible for spinal proprioceptive reflexes in man

1. To define the neural volleys responsible for the Achilles tendon jerk and the H reflex, muscle afferent activity was recorded using micro‐electrodes inserted percutaneously into appropriate fascicles of the tibial nerve in the popliteal fossa.

Interactions between cutaneous and muscle afferent projections to cerebral cortex in man

In order to demonstrate interactions between cutaneous and muscle afferent volleys in the ascending somatosensory pathways, different nerves of the lower limb were stimulated together in a conditioning-test paradigm, the changes in the earliest component of the cerebral potential evoked by the test stimulus being taken to indicate such an interaction. It was first confirmed that the cerebral potential evoked by stimulation of the posterior tibial nerve at the ankle is derived from muscle afferents in the mixed nerve and has shorter latencies than the cerebral potential evoked by purely cutaneous volleys in the sural nerve (see Burke et al. 1981). Complete suppression of the cerebral potential evoked by stimulation of muscle or cutaneous afferents was produced by conditioning volleys in a different nerve or in a different fascicle of the same nerve. The major factors determining the degree of suppression were found to be the relative sizes of the conditioning and test volleys and their timing, rather than whether the volleys were of cutaneous or muscular origin. It is concluded that the transmission of cutaneous or muscle afferent volleys to cortex can be profoundly altered in normal subjects by conditioning activity. The possibility that normal background afferent activity can similarly modify afferent transmission has implications for diagnostic studies, particularly when they are performed under non-standard conditions, such as in the operating theatre or intensive care unit. It is also concluded that, although a subject may perceive cutaneous paraesthesiae when the posterior tibial nerve is stimulated at the ankle, there may be no cutaneous component to the evoked cerebral potential.

Anticipation and fusimotor activity in preparation for a voluntary contraction.

1. Multi‐unit and single‐unit recordings were made of muscle spindle afferent activity from the pretibial muscles of human subjects who were initially relaxed. The muscles were subjected to a stretching perturbation of 1 s duration, occurring irregularly, on average once every 5 s. In test sequences, an auditory or visual warning was provided 1 . 06 s before some of the perturbations. Subjects were required to oppose every perturbation by contracting the receptor‐bearing muscle as rapidly as possible. 2. Following the warning all subjects sometimes tensed the receptor‐bearing muscle unintentionally in preparation for the perturbation. In these contractions, the discharge of a spindle ending accelerated only if the contraction strength exceeded the endings threshold for activation, established in control voluntary contractions performed under isometric conditions. 3. When the receptor‐bearing muscle did not contract in the interval between warning and perturbation, there was no detectable change in the multi‐unit recordings of spindle activity or in recordings from twelve of thirteen single spindle afferents. The thirteenth spindle afferent discharged prior to the perturbation in the absence of detectable e.m.g. in response to (only) three of twenty‐three warning stimuli. However, this ending had been so responsive during isometric voluntary contractions that a contraction level at which it did not respond could not be established, and it is suggested that the findings with this ending resulted from its low threshold rather than from selective activation of the fusimotor system. 4. When subjects were warned of the perturbations, the dynamic response of spindle endings to the perturbations was not increased in size or altered in latency. 5. The motor response to perturbations without warning generally contained only long‐latency (volitional) e.m.g. activity occurring 107‐‐200 ms after the onset of the perturbation. When a warning was given, short‐latency (reflex) e.m.g. activity was also recorded, beginning 46‐‐76 ms after the onset of the perturbation. 6. It is concluded that anticipation of the need to contract a muscle does not result in selective activation of fusimotor neurones in preparation for the contraction. The change in stretch reflex gain that occurs as a result of ‘anticipation’ occurs through a central process which does not involve the fusimotor system.

Convergence in the somatosensory pathway between cutaneous afferents from the index and middle fingers in man.

SummaryAverage short-latency cerebral potentials were recorded from the parietal scalp to mechanical stimulation of the index and middle fingers and to electrical stimulation of the digital nerves in normal subjects. The early components of the cerebral potential, representing the arrival of the afferent volley at the sensorimotor cortex, were studied during stimulation of the fingers separately and together. When strong or moderate stimuli were used there was a suppressive interaction between the afferent input from the two fingers with either electrical or mechanical stimulation. During simultaneous stimulation of both fingers the size of the early component of the cerebral potential was less than predicted by simple addition of the potentials produced by stimulation of the fingers individually.When very weak stimuli, close to the level necessary for detection by the subject, the input from the two fingers produced additive or facilitatory interactions in the early components of the cerebral potential. These results suggest that there is convergence between the afferent inputs from the index and middle fingers along the somatosensory pathway. At levels of stimulation comparable to those which produced facilitation in the electrophysiological studies, simultaneous stimulation to both fingers was detected significantly more frequently than would be expected from the detection of stimulation to individual fingers.

Coupling between human muscle spindle endings and motor units assessed using spike-triggered averaging

The electromyographic activity (EMG) generated by voluntary contraction of a muscle was averaged using the potentials from 18 identified muscle spindle afferents as a trigger. In post-spike averages of 1000-10,000 sweeps, no evidence of reflex excitation of the homonymous motoneurone pool was detected. In pre-spike averages there was no evidence of a motor-unit EMG potential that was closely correlated to the trigger spike. A single spindle afferent has only a weak reflex effect on an active motoneurone pool and must be part of a synchronized volley to affect motoneurone discharge significantly. No evidence was found for spindle activation via beta-motoneurones in weak voluntary contractions.

The relationship between the size of a muscle afferent volley and the cerebral potential it produces.

This study examined the relationship between the size of an afferent neural input produced by electrical stimulation of the posterior tibial nerve at the ankle and the size of the early components of the evoked cerebral potential. For five of six subjects the first peak of the afferent neural volley recorded in the popliteal fossa was uncontaminated by either motor efferents or cutaneous afferents. This was established by measuring the conduction times of motor fibres in the posterior tibial nerve and cutaneous fibres in the sural and posterior tibial nerves over the ankle to popliteal fossa segment. It is likely therefore that the first peak of the afferent volley contained predominantly, if not exclusively, activity in rapidly conducting afferents from the small muscles of the foot. The size of the two earliest components of the cerebral potential did not increase in direct proportion to the size of the afferent volley which produced it. The early components of the cerebral potential reached a maximum when the responsible muscle afferent volley was less than 50% of its maximum.

Reproducibility of the visual evoked potential using a light-emitting diode stimulator.

The intersubject and intrasubject reproducibility of the VEP was evaluated using two different methods of producing pattern reversal--a mirror/projector system and a light-emitting diode (LED) system. Intersubject reproducibility was determined in 100 normal subjects (50 males, 50 females). Ten subjects were studied on ten different occasions over 11 months to establish intrasubject reproducibility. The two methods gave comparable results. It is concluded that the mirror/projector system has no advantages over the LED system, which is cheaper, more robust and more convenient. Most of the intersubject variability was found to be due to subject variables and it is concluded that technical improvements are unlikely to reduce further the variability of the test. A variability in the latency of P100 of up to 7.7 ms was recorded in the serial studies on the ten subjects, indicating that in longitudinal studies on patients, changes in latency must exceed 9-10 ms to be significant. With both stimulus systems, there were significant sex-related differences in latency (P50 and P100 but not N70) and amplitude (N70-P100). The extent of the difference was such that the upper limit of normal latency for P100 (2.5-3 SD from the mean) was 4.2-4.7 ms longer for males than females-a value which exceeds 1 SD. Separate control values for males and females are advisable whichever method is used to produce pattern reversal.

Identification of muscle spindle afferents during in vivo recordings in man

A preamplifier for use in microelectrode recordings from human muscle afferents is described. Electrical stimuli can be delivered through the microelectrode to produce the twitch contractions necessary for afferent identification while still recording the activity of the afferent fibre.