Frederick W.J. Cody

A saccular origin of frequency tuning in myogenic vestibular evoked potentials?: implications for human responses to loud sounds

Previous research has indicated that an early component of click-evoked myogenic potentials in the sternocleidomastoid muscle is vestibularly mediated, since it can be obtained in subjects with loss of cochlear function, but is absent in subjects with loss of vestibular function (Colebatch et al., 1994). We report here the results of an experiment to investigate whether this response shows any tuning properties. In a sample of 11 subjects, we obtained acoustically evoked EMG from the sternocleidomastoid muscle in response to 110 dB SPL 10 ms tone pips with frequencies of 100 Hz, 200 Hz, 400 Hz, 800 Hz, 1600 Hz and 3200 Hz. The results of this experiment indicate that this response does indeed have a well-defined frequency tuning which may be modelled as a resonance with a maximum response at frequencies between 300-350 Hz. The possible saccular origin of the tuning response and the consequences that this may have in human responses to loud sounds is discussed. Also discussed are the consequences of particular electrode arrangements in relation to the innervation and anatomy of sternocleidomastoid.

Cortical control of human soleus muscle during volitional and postural activities studied using focal magnetic stimulation

The surface-recorded electromyographic (EMG) responses evoked in the ankle musculature by focal, transcranial, magnetic stimulation of the motor cortex were studied in healthy human subjects. Such soleus evoked motor responses (EMRs) were characterised over a wide range of background levels of motor activity and using different stimulus intensities. EMRs were recorded during predominantly (1) volitional and (2) postural tasks. In the former task subjects were seated and voluntarily produced prescribed levels of soleus activation by reference to a visual monitor of EMG. In the latter task subjects assumed standing postures without EMG feedback. Comparison of the EMRs of soleus, traditionally considered a slow anti-gravity extensor muscle, during these tasks was used to evaluate its cortical control in primarily volitional versus primarily postural activities. The form of soleus EMRs produced by single magnetic cortical stimuli comprised an initial (approx. 30 ms) increase and subsequent (approx. 50 ms) depression of EMG. Cortical stimulation could elicit substantial excitatory soleus EMG responses; for example, responses evoked by mild, magnetic stimuli (125% threshold for inducing a response in the relaxed muscle) as subjects exerted full voluntary plantarflexor effort averaged almost 20% of the maximum M-wave which could be elicited by an electrical stimulus to the posterior tibial nerve. Excitatory EMRs could be elicited in the voluntarily relaxed soleus muscle of the majority of subjects during sitting. The amplitude of soleus responses, induced by threshold stimuli for the relaxed state or approximately 125% threshold intensity, increased approximately linearly with background EMG over a wide range of volitional contraction levels. By contrast, there was no systematic change in the latency of excitatory soleus EMRs with increasing voluntary effort. The excitatory responses evoked in the voluntarily relaxed soleus of seated subjects by magnetic stimulation were regularly facilitated by incremental, voluntary contraction of the contralateral ankle extensors in a graded manner. However, such facilitation of responses was not observed when subjects voluntarily activated the muscle in which EMRs were elicited. The pattern of the responses elicited in soleus by magnetic stimulation during the postural task generally resembled that found during the volitional task. The amplitudes of excitatory soleus EMRs at a given stimulus intensity, obtained when subjects stood quietly, leaned forwards or stood on their toes to produce differing levels of ankle extensor contraction, increased with background EMG. Overall, the relationship between the size of cortically evoked soleus responses and the tonic level of motor activity, observed in individual subjects at matched stimulus intensities, did not consistently differ between postural and volitional tasks. The present results suggest that the motor cortex is potentially capable of exerting rapid regulation of the soleus muscle, and presumably other ankle extensors, not only when the muscle participates in volitional tasks but also when it is engaged in postural maintenance.

Tactile spatial acuity varies with site and axis in the human upper limb

Historically, beginning with Webers [E.H. Weber, On the sensitivity of the tactile senses, in: H.E. Ross, D.J. Murray (Eds. and Trans.), E.H. Weber on the Tactile Senses, Erlbaum (UK) Taylor & Francis, Hove, 1996 (Original work published in 1834), pp. 21-136] classical studies, regional variations in the accuracy of localisation of tactile stimuli applied to a limb have been recognised. However, important questions remain concerning both the map of localisation resolution and its neuroscientific basis since methodological confounds have militated against an unambiguous, unified interpretation of the diverse findings. To test the hypotheses that localisation precision on the upper limb varies with site (hand, wrist, forearm) and limb axis (transverse, longitudinal), regional differences in locognosic acuity were quantified in psychophysical experiments. Participants identified the perceived direction (e.g. medial or lateral) relative to a central reference locus of brief tactile test stimuli applied to a cruciform array of loci. Acuity was greater in the transverse than longitudinal axis. This effect probably arises from the asymmetry of receptive fields of upper limb first-order sensory units and their higher-order projection neurons. Additionally, acuity was greater on the dorsal surface at the wrist than either the hand or forearm sites, in the longitudinal axis, supporting an enhancement of resolution at joints (anchor points). This effect may contribute to improved proprioceptive guidance of active wrist movements.

Proprioceptive guidance of human voluntary wrist movements studied using muscle vibration.

1. The alterations in voluntary wrist extension and flexion movement trajectories induced by application of vibration to the tendon of flexor carpi radialis throughout the course of the movement, together with the associated EMG patterns, have been studied in normal human subjects. Both extension and flexion movements were routinely of a target amplitude of 30 deg and made against a torque load of 0.32 N m. Flexor tendon vibration consistently produced undershooting of voluntary extension movements. In contrast, voluntary flexion movements were relatively unaffected. 2. The degree of vibration‐induced undershooting of 1 s voluntary extension movements was graded according to the amplitude (0.75, 1.0 and 1.5 mm) of flexor tendon vibration. 3. As flexor vibration was initiated progressively later (at greater angular thresholds) during the course of 1 s voluntary extension movements, and the period of vibration was proportionately reduced, so the degree of vibration‐induced undershooting showed a corresponding decline. 4. Varying the torque loads (0.32, 0.65 and 0.97 N m) against which 1 s extension movements were made, and thereby the strength of voluntary extensor contraction, produced no systematic changes in the degree of flexor vibration‐induced undershooting. 5. Analysis of EMG patterns recorded from wrist flexor and extensor muscles indicated that vibration‐induced undershooting of extension movements resulted largely from a reduction in activity in the prime‐mover rather than increased antagonist activity. The earliest reductions in extensor EMG commenced some 40 ms after the onset of vibration, i.e. well before voluntary reaction time; these initial responses were considered to be ‘automatic’ in nature. 6. These results support the view that the central nervous system utilizes proprioceptive information in the continuous regulation of moderately slow voluntary wrist movements. Proprioceptive sensory input from the passively lengthening antagonist muscle, presumably arising mainly from muscle spindle I a afferents, appears to be particularly important and to act mainly in the reciprocal control of the prime‐mover.

Patterns of activity evoked in cerebellar interpositus nuclear neurones by natural somatosensory stimuli in awake cats

1. Stable extracellular unitary recordings were made from 138 cerebellar interpositus nuclear neurones (IPNs) in awake cats. Mean background discharge, in animals in a state of relaxed wakefulness and in the absence of overt movement, was 41·0±2·6 impulses/sec (mean± s.e.m).

Effect of click trains on duration estimates by people with Parkinson's disease.

Patients with a diagnosis of Parkinsons disease and age- and IQ-matched controls estimated the duration of short 500-Hz tones (325–1,225 ms), on trials where the tone was either preceded by 3 s of 5-Hz clicks, or presented without clicks. The click manipulation had been shown in earlier studies with student participants to make verbal estimates longer. Patients were tested both on and off their dopaminergic medication, and controls were also tested in two sessions. Verbal estimates were markedly and significantly longer on trials with clicks than on those without clicks for both the patients and the controls, but there were no significant performance differences between patients or controls, nor between the on and off medication sessions in the patients. The study shows that a manipulation of subjective time, which has had small but consistent effects in student participants, also affects timing in patients and adds to a growing body of evidence that timing in patients with Parkinsons disease may in many cases have the same characteristics as those of neurologically intact control groups.

Vibration-evoked reciprocal inhibition between human wrist muscles

SummaryReciprocal inhibition of the voluntarily contracting wrist extensor (extensor carpi radialis, ECR) evoked by proprioceptive afferent input from the flexor (flexor carpi radialis, FCR), was studied in healthy human subjects. Vibration of the FCR tendon was used to elicit Ia-dominated afferent discharge whilst inhibition of ECR was assessed as the reduction in asynchronous, on-going EMG. A small early phase of inhibition (I1) was evident in 25% of trials. The latency (ca. 25 ms) of this component suggested that it was mediated by an Ia oligosynaptic, possibly ‘classical’ disynaptic, inhibitory pathway. A later and apparently separate phase of reduced activity (12, ca. 40 ms) was, however, far more consistently observed (96% of trials) and of greater magnitude. The 12 component was usually followed, some 20 ms later, by a phase of elevated activity (El, 72% trials). Reductions in simultaneously recorded net extensor torque commenced at about 60 ms following the onset of flexor tendon vibration, i.e. some 20 ms after the main I2 EMG component. These mechanical responses must have almost exclusively resulted from reciprocal inhibition of extensor EMG since vibration of the relaxed FCR evoked minimal excitatory flexor activity. The reflex pattern, in any individual subject, was relatively unaffected by altering the duration of the vibration train between one and nineteen cycles (125 Hz). This suggests that the entire response complex resulted largely from the initial afferent volley. The sizes of both the I1 and I2 reductions in ECR activity increased with increasing voluntary extensor contraction so that their depths remained constant proportions of background EMG. Very similar results were obtained when reciprocal inhibition of FCR was produced by vibration of the belly of ECR. Thus, reciprocal inhibition between wrist muscles is mainly expressed as a rather stereotyped, short duration reduction in EMG whose depth is determined by the pre-existing level of motor activity. Some functional implications of this form of reflex behaviour are discussed.

Electromyographic reflexes evoked in human wrist flexors by tendon extension and by displacement of the wrist joint.

1. The electromyographic (EMG) reflexes evoked in the wrist flexor muscle, flexor carpi radialis (FCR), by percutaneous extension of its tendon and by forcible extension of the wrist joint have been studied. Reflexes were elicited during steadily maintained voluntary flexor contraction of 10% of each subjects maximum. 2. Tendon extension, using ‘ramp and hold’ displacements, evoked fairly prolonged (ca 50 ms) increases in EMG activity. These responses were usually subdivided into two main excitatory peaks of respectively short (SL, ca 20 ms) and long (LL, ca 45 ms) latency. This pattern contrasted with that observed following brief tendon taps when only a single, SL peak was elicited. 3. ‘Stretch’ reflexes evoked by ‘ramp and hold’ wrist extensions, as has been noted by numerous earlier investigators, were also protracted and comprised two main excitatory components. These responses resembled those produced by tendon extension both in their general form and in their behaviour upon altering the velocity of mechanical stimuli. Quantitatively, however, two main differences were evident. The reflexes evoked by wrist extension, including their SL and LL peaks, were generally somewhat larger. Additionally, when parameters of the two modes of stimulation were adjusted to elicit SL responses of equivalent amplitude, the LL responses elicited by tendon extension were regularly smaller and of shorter duration than those elicited by wrist extension. 4. Termination of the two forms of mechanical stimulation, by releasing tendon or wrist extension, each elicited a SL reduction in EMG activity. Such troughs were more pronounced and more consistently observed upon release of wrist extension. 5. Neither local anaesthesia of the skin overlying the flexor tendons at the wrist nor ischaemia of the hand and lower forearm produced any systematic modification of reflex response patterns. 6. It is concluded that intramuscular receptors (presumably muscle spindles) in FCR mediate both the SL and LL reflexes evoked in this muscle by extension of its tendon. Intramuscular receptors also seem certain to be very largely responsible for the EMG responses generated in this muscle by wrist extension.

Tactile spatial acuity is reduced by skin stretch at the human wrist

The skin is an elastic organ that is continuously distorted as our limbs move. The hypothesis that the precision of human tactile localisation is reduced when the skin is stretched, with concurrent expansion of receptive fields (RFs) was tested. Locognosic acuity over the dorsal wrist area was quantified during application of background stretch by (a) Wrist-Bend (skin stretch combined with non-cutaneous proprioceptor activation) and (b) Skin-Pull (skin stretch alone). Participants identified the perceived direction (distal or proximal) of brief test stimuli, applied along a 7-point linear array, relative to a central reference locus. Performance was significantly reduced during the large amplitude compared to the small amplitude of tonic skin stretch, but there was no effect of stretch mode (Wrist-Bend, Skin-Pull), nor was the effect of stretch amplitude modulated by the mode of stretch. Locognosic acuity was poorer than baseline accuracy for the large amplitude skin stretches, for both application modes, but did not differ significantly from baseline for either of the small amplitude stretches. We interpret these observations as corroborating the long-held assumption that tactile localisation is primarily dependent upon the RF dimensions, and associated innervation densities, of regional touch units. The finding that performance was reduced to a similar extent under Skin-Pull and Wrist-Bend conditions suggests that non-cutaneous proprioceptors had rather little tonic modulatory effect.

Proprioception in Human Basal Ganglia Disorders

Several lines of evidence indicate that both reflex and voluntary motor responses to proprioceptive input are abnormal in human basal ganglia disorders. Long-latency stretch reflexes are pathologically enhanced in Parkinson’s disease (PD; Tatton & Lee, 1975; Cody, MacDermott, Matthews & Richardson, 1986) and may be reduced or absent in Huntington’s disease (HD; Noth, Podoll & Friedemann, 1985). Additionally, the finding of Moore (1987) that Parkinsonian patients with asymmetrical disease overestimate the trajectory of the more bradykinetic limb when attempting to match slow, active movements of the two arms suggests a disturbance of proprioceptive guidance in basal ganglia dysfunction.