Selective changes of ocular vestibular myogenic potentials in Parkinson's disease.

BACKGROUND: Vestibular evoked myogenic potentials represent electrophysiological tools to measure vestibular reflex actions at different levels of the brainstem in Parkinson's disease. OBJECTIVE: To investigate cervical and ocular vestibular myogenic potentials in Parkinsonian patients with mild disability. METHODS: In 13 Parkinsonian patients and 13 age-matched healthy controls, cervical and ocular vestibular myogenic potentials were recorded after unilateral air-conducted tone bursts and bone-conducted stimuli delivered at the forehead or mastoids. RESULTS: In contrast to relatively preserved cervical vestibular evoked myogenic potentials, ocular vestibular evoked myogenic potentials were significantly delayed and of reduced amplitude, particularly after impulsive stimulation in Parkinsonian patients. Levodopa had no significant effect on either type of response. CONCLUSION: In mild to moderate Parkinson's disease, altered ocular vestibular myogenic potentials may indicate early functional involvement of the upper brainstem, in contrast to preserved lower brainstem function as reflected by normal cervical vestibular myogenic potentials.

Properties of rectified averaging of an evoked-type signal: theory and application to the vestibular-evoked myogenic potential.

The properties of rectified averages were investigated using the VEMP (vestibular-evoked myogenic potential) as an example of an evoked-type response. Recordings were made of surface EMG from the sternocleidomastoid (SCM) muscles of six volunteers, unstimulated, at different levels of tonic activation and then in response to clicks of different intensities. The stochastic properties of the surface EMG recorded were shown to be well modelled using a zero mean normal distribution with a standard deviation equivalent to the mean RMS (root mean squared) value (mean residual error variance 0.87%). Assuming a normal distribution, equations were derived for the expected value of both the rectified and RMS average with the addition of constant waveforms of different sizes. A simulation using recorded EMG and added sine waves of different amplitudes demonstrated that the equations predicted the rectified averages accurately. It also confirmed the importance of the relative amplitude of the added signal in determining whether it was detected using rectified averages. The same equations were then applied to actual data consisting of VEMPs of different relative amplitudes recorded from the volunteers. Whilst the signal-to-noise ratio (measured by corrected amplitude) was a major determinant of the nature of the rectified average, consistent deviations were detected between the predicted and actual rectified averages. Deviations from predicted values indicated that the VEMP did not behave simply like a constant signal added to tonic background EMG. A more complicated model, which included temporal jitter as well as inhibition of background EMG during the VEMP, was required to fit the physiological recordings. Rectified averages are sensitive to physiological properties, which are not apparent when using unrectified averages alone. Awareness of the properties of rectified averages should improve their interpretation.

Ocular vestibular evoked myogenic potentials in superior canal dehiscence.

OBJECTIVE: Patients with superior canal dehiscence (SCD) have large sound-evoked vestibular reflexes with pathologically low threshold. We wished to determine whether a recently discovered measure of the vestibulo-ocular reflex-the ocular vestibular evoked myogenic potential (OVEMP)-produced similar high-amplitude, low-threshold responses in SCD, and could differentiate patients with SCD from normal control patients. METHODS: Nine patients with CT-confirmed SCD and 10 normal controls were stimulated with 500 Hz, 2 ms tone bursts and 0.1 ms clicks at intensities up to 142 dB peak SPL. Conventional VEMPs were recorded from the ipsilateral sternocleidomastoid muscle to determine threshold, and OVEMPs were recorded from electrode pairs placed superior and inferior to the eyes. Three-dimensional eye movements were measured with scleral dual-search coils. RESULTS: In patients with SCD, OVEMP amplitudes were significantly larger than normal (p<0.001) and thresholds were pathologically low. The n10 OVEMP in the contralateral inferior electrode became particularly large with increasing stimulus intensity (up to 25 microV) and with up-gaze (up to 40 microV). Sound-evoked (slow-phase) eye movements were present in all patients with SCD (vertical: upward; torsional: upper pole away from the affected side; and horizontal: towards or away from the affected side), but began only as the OVEMP response became maximal, which is consistent with the surface potentials being produced by activation of the extraocular muscles that generated the eye movements. CONCLUSIONS: OVEMP amplitude and threshold (particularly the contralateral inferior n10 response) differentiated patients with SCD from normal controls. Our findings suggest that both the OVEMPs and induced eye movements in SCD are a result of intense saccular activation in addition to superior canal stimulation.

Vestibular hypersensitivity to sound in superior canal dehiscence: large evoked responses in the legs produce little postural sway.

OBJECTIVE: Patients with superior canal dehiscence (SCD) typically have enhanced sound-evoked vestibular reflexes, such as vestibulo-collic and vestibulo-ocular reflexes. We wished to investigate whether sound-evoked lower limb EMG responses and postural sway are also enhanced in this condition. METHODS: Eight patients with CT confirmed SCD (11 affected ears) and 8 age-matched normal controls participated. Three sound-evoked responses were measured; vestibulo-collic reflexes (i.e. vestibular-evoked myogenic potentials, VEMPs), lower limb vestibulo-spinal reflexes and body sway (centre of pressure in mm). Sound stimuli were 500 Hz air-conducted tone bursts of varying lengths (VEMPs: 2 ms; vestibulo-spinal: 20 ms; sway: 1s and 200 ms) set at fixed levels above each subject's VEMP threshold. RESULTS: SCD patients had very large VEMP and vestibulo-spinal responses following high intensity stimulation, but at the matched intensity of 15 dB above threshold amplitudes were similar in both SCD patients and controls. The amplitude of both responses increased linearly with increasing stimulus intensity in both groups. Large ( approximately 20mm), stereotyped sway responses were present in only one (atypical) patient with high intensity stimulation. Small ( approximately 2mm) sway responses were present in the remaining patients, and began immediately following the vestibulo-spinal responses. CONCLUSIONS: Despite the presence of large vestibular reflexes, there is usually very little body sway in response to loud sounds in SCD patients. SIGNIFICANCE: Large short-latency vestibulo-spinal reflexes in SCD do not necessarily evoke large sway responses.

Vestibular-evoked myogenic potentials.

The vestibular-evoked myogenic potential (VEMP) is a short-latency potential evoked through activation of vestibular receptors using sound or vibration. It is generated by modulated electromyographic signals either from the sternocleidomastoid muscle for the cervical VEMP (cVEMP) or the inferior oblique muscle for the ocular VEMP (oVEMP). These reflexes appear to originate from the otolith organs and thus complement existing methods of vestibular assessment, which are mainly based upon canal function. This review considers the basis, methodology, and current applications of the cVEMP and oVEMP in the assessment and diagnosis of vestibular disorders, both peripheral and central.

Vestibular-dependent inter-stimulus interval effects on sound evoked potentials of central origin.

Todd et al. (2014ab) have recently demonstrated the presence of vestibular-dependent contributions to auditory evoked potentials (AEPs) when passing through the vestibular threshold as determined by vestibular evoked myogenic potentials (VEMPs), including a particular deflection labeled as an N42/P52 prior to the long-latency AEPs N1 and P2. In this paper we report the results of an experiment to determine the effect of inter-stimulus interval (ISI) and regularity on potentials recorded above and below VEMP threshold. Five healthy, right-handed subjects were recruited and evoked potentials were recorded to binaurally presented sound stimulation, above and below vestibular threshold, at seven stimulus rates with ISIs of 212, 300, 424, 600, 848, 1200 and 1696 ms. The inner five intervals, i.e. 300, 424, 600, 848, 1200 ms, were presented twice in both regular and irregular conditions. ANOVA on the global field power (GFP) were conducted for each of four waves, N42, P52, N1 and P2 with factors of intensity, ISI and regularity. Both N42 and P52 waves showed significant ANOVA effects of intensity but no other main effects or interactions. In contrast both N1 and P2 showed additional effects of ISI, as well as intensity, and evidence of non-linear interactions between ISI and intensity. A source analysis was carried out consistent with prior work suggesting that when above vestibular threshold, in addition to bilateral superior temporal cortex, ocular, cerebellar and cingulate sources are recruited. Further statistical analysis of the source currents indicated that the origin of the interactions with intensity may be the ISI sensitivity of the vestibular-dependent sources. This in turn may reflect a specific vestibular preference for stimulus rates associated with locomotion, i.e. rates close to 2 Hz, or ISIs close to 500 ms, where saccular afferents show increased gain and the corresponding reflexes are most sensitive.

Vestibular responses to sound.

Research into vestibular responses to sound has evolved in four stages. The first, largely the work of Tullio in the 1920s, involved inspection of the eye, head, and postural responses to sound of alert animals with surgical fenestrae into various parts of the bony labyrinth. The second, begun in 1964 by Bickford and his group and continued by our group and then by others in the last 10 years, involves the measurement of evoked myogenic potentials to air-conducted and bone-conducted clicks and tones in normal humans. The third, begun by Mikaelian at about the same time as Bickford and continued by McCue, our group, and others, involves electrophysiological recordings of primary vestibular afferent neuron responses to sound in anesthetized animals. The fourth involves measurements of vestibulo-ocular responses to sound in humans with the Tullio phenomenon. It was begun by Minor and his group in 1998 with the observation that sound-induced nystagmus in humans, the Tullio phenomenon, aligned with the rotation axis of the superior semicircular canal. They then showed a defect in the temporal bone between the apex of the superior semicircular canal and the middle cranial fossa, which was the cause of most, if not all, cases of sound-induced nystagmus. Here some of the key observations made in each of these four stages are reviewed.

Vestibular-evoked extraocular potentials produced by stimulation with bone-conducted sound.

OBJECTIVE: To investigate the origin, whether ocular or extraocular, of the short latency frontal potential (N15) reported by following vestibular stimulation. METHODS: Fourteen subjects with low VEMP thresholds (V(T)) and 9 patients with vestibular or ocular disorders were stimulated at the mastoid with bone-conducted tone bursts (500 Hz, 8 ms) above vestibular threshold, using a B71 bone vibrator. Surface potentials were recorded from Fpz and around the eyes and referred to linked earlobes. RESULTS: The N15 was present at Fpz, but was largest around the eyes (mean amplitude 2.6 microV, peak latency 13.4 ms, with stimulation at +18 dB above threshold) and was generally in phase above and below the eyes. The response was vestibular-dependent and modulated by alteration of gaze direction. The potentials were delayed in a patient with Miller Fisher syndrome and were larger in patients with superior canal dehiscence than in controls. CONCLUSIONS: We report a new vestibular-evoked extraocular potential. Its properties are not consistent with an eye movement. It is likely to be produced, mainly or exclusively, by synchronous activity in extraocular muscles (i.e. a myogenic potential). SIGNIFICANCE: Vestibular-evoked extraocular potentials extend the range of vestibular pathways that can be assessed electrophysiologically, and may be a useful additional test of vestibular function.

Decreases in regional cerebral blood flow with normal aging.

Positron emission tomographic (PET) images of regional cerebral blood flow (rCBF) from 30 normal, resting volunteers aged 30 to 85 years were analysed to identify areas where rCBF fell with age. Images were anatomically normalised, and a pixel-by-pixel linear regression was performed to remove differences in global CBF between subjects. Pixels at which rCBF then showed a significant (p less than 0.01) negative correlation with age were identified. They were displayed as a statistical parametric map (SPM) of correlations. We demonstrate an age-related decrease in adjusted rCBF in the cingulate, parahippocampal, superior temporal, medial frontal, and posterior parietal cortices bilaterally, and in the left insular and left posterior prefrontal cortices (omnibus significance, chi 2 = 2,291, p less than 0.0001, df = 1). Decreases in rCBF suggest a regionally specific loss of cerebral function with age. The affected areas were all limbic, or association, cortices. Therefore, these decreases may constitute the cerebral substrate of the cognitive changes that occur during normal aging.

Tapping the head activates the vestibular system: a new use for the clinical reflex hammer.

We investigated the use of skull taps with a modified clinical reflex hammer as a method of vestibular activation. Using recently described EMG techniques to measure vestibulocollic reflexes in response to clicks, we were able to show analogous short-latency potentials to taps. The earliest responses were invariably absent on the side of a previous vestibular nerve section but were preserved in profound sensorineural or conductive hearing loss. We propose that the taps activated the vestibular apparatus directly by a bone-conducted vibration wave.

Vestibular hypersensitivity to sound (Tullio phenomenon): structural and functional assessment.

OBJECTIVES: To establish the role of high-resolution CT imaging and tests of vestibulocollic reflexes in diagnosing and understanding the pathogenesis of the Tullio phenomenon. BACKGROUND: The Tullio phenomenon is a syndrome in which acoustic stimulation produces symptoms and signs of vestibular activation. It has previously been associated with an abnormally low threshold for click-evoked vestibulocollic responses and also with dehiscence of the roof of the anterior (superior) semicircular canal on high-resolution CT scans of the temporal bones. METHODS: High-resolution CT scans of the temporal bones and vestibulocollic responses in sternocleidomastoid to both clicks and transmastoid galvanic stimulation (3 mA/2 msec) were studied in four patients with the Tullio phenomenon (one bilateral). RESULTS: Click-evoked thresholds were low for all affected ears (four at 65 dB nHL, one at 55 dB nHL) and normal (>70 dB nHL) for the three unaffected ears. In contrast, galvanic-evoked vestibulocollic responses were symmetric and of normal size in all patients. The bony roof of the anterior (superior) semicircular canal was thin, possibly absent, on CT of all affected ears and also in two out of three unaffected ears. CONCLUSIONS: The normal galvanic vestibulocollic responses indicate that sound sensitivity in patients with the Tullio phenomenon is likely to occur distal to the vestibular nerve, probably at the level of the receptors. Both click hypersensitivity and dehiscence of the anterior (superior) semicircular canal are associated with the Tullio phenomenon but as the CT scan abnormality can occur in clinically unaffected ears, click testing is important for specific diagnosis. Abnormal sound sensitivity, as demonstrated by click responses, confirms that the radiologic abnormality is function significant.

Measurement of capillary pressure in humans using a venous occlusion method.

The venous occlusion technique was used to measure capillary pressure in the forearm and foot of man over a wide range of venous pressures. In six recumbent subjects venous pressure (Pv) in the forearm (mean +/- SE) was 9.3 +/- 1.4 mmHg and the venous occlusion estimate of capillary pressure (Pc) was 17.0 +/- 1.6 mmHg, whereas in another six subjects Pv in the foot was 17.1 +/- 1.2 mmHg and Pc was 23.4 +/- 2.5 mmHg. Venous pressure in the limbs was increased either by changes in posture or by venous congestion with a sphygmomanometer cuff. On standing Pv in the foot increased to 95.2 +/- 1.5 mmHg and Pc rose to 112.8 +/- 3.1 mmHg. The relationship established between venous pressure and capillary pressure in the forearm is Pc = 1.16 Pv + 8.1, whereas in the foot the relationship is Pc = 1.2 Pv + 1.6. The magnitude and duration of the changes in capillary pressure were also recorded during reactive hyperemia. The venous occlusion method of measuring capillary pressure is simple and easily applied to studies in humans.

A modified method of estimating phase resetting of rhythmical movement.

A computer-based method of estimating resetting index is presented which was successfully applied to a study of resetting of human voluntary rhythmical movement. Peaks were detected before and after the stimulus (magnetic cortical stimulation) from position records and the phase of stimulation and the phase change induced by the stimulus estimated. The phase change versus phase of stimulation relationship was rotated until the best linear fit was obtained: the gradient of this relationship is the resetting index. The calculation of resetting index was thus independent of the initial assignment of reference phase. A constant, the null phase, was derived from the best linear fit and indicates the phase to which the original waveform is being reset by the stimulus. Statistical tests of significance and linearity of the final relationship are also described.

Effects of externally imposed elastic loads on the ability to estimate position and force.

Normal adult subjects attempted, with vision excluded, to match the position of the left index finger by pointing to it with the right one, using only flexion-extension movements at the right elbow. An elastic load applied at the right wrist was altered from trial to trial. When instructed to align the fingers, subjects were found to select the position of the right forearm by taking into account both a position signal and some measure of the force exerted by the elbow flexors. When instructed to match a target force, instead of position, the subjects were able to give greater weighting to signals of force than in the position matching task.

Motor unit excitability changes mediating vestibulocollic reflexes in the sternocleidomastoid muscle.

OBJECTIVE: To investigate the patterns of motor unit firing in single motor units from the sternocleidomastoid (SCM) muscles in response to stimuli previously shown to be capable of evoking vestibulocollic reflexes (loud clicks and electrical stimulation) and to relate these to the previously described surface potentials (VEMPs, vestibular evoked myogenic potentials) evoked by the same stimuli. METHODS: Eleven male subjects (30-43-years-old) were studied. Local anaesthetic was used to block the SCM and confirm that the surface potentials (p13n23) arose from it. At another time, fine wire or needle electrodes were used to record single motor unit activity and peristimulus time histograms were constructed. RESULTS: Local anaesthetic block reduced or abolished the p13n23 response in 5 of 6 subjects. A total of 94 histograms of motor unit discharges were recorded. The excitability changes seen were always small. Loud click stimuli given ipsilaterally evoked short latency (mean 14.2 ms, uncorrected for triggering delays), and short duration (mean 3.6 ms) inhibition. Contralaterally, the effect was excitatory (latency 11.9 ms, uncorrected, mean duration 2.3 ms). For electrical stimulation, short latency inhibition occurred ipsilateral to the cathode (average latency 14.0 ms, uncorrected, mean duration 2.9 ms) and excitation occurred contralaterally. CONCLUSIONS: We confirmed that the SCM is the sole or dominant source of the VEMPs recorded from electrodes over it. Short latency, short duration motor unit firing changes are evoked in SCM by loud clicks and electrical stimulation, stimuli known to be capable of evoking vestibulocollic reflexes. VEMPs beginning with a positivity correspond to inhibition of the underlying motor unit firing and those starting with a negativity correspond to an underlying excitation, findings consistent with intracellular recordings of otolith effects. Qualitative explanations of how the surface potentials are generated by these excitability changes and relating to other properties of the surface responses are proposed. SIGNIFICANCE: This study has shown consistent patterns of single motor unit firing which underlie VEMPs evoked by both clicks and short duration galvanic stimulation.

Galvanic evoked vestibulospinal and vestibulocollic reflexes in stroke.

OBJECTIVE: Following stroke, the startle reflex, mediated via the reticulospinal tract, is often facilitated. Vestibulospinal reflexes are another bulbospinal reflex, abnormalities of which may contribute to impaired body posture and stance following stroke. We recorded galvanic evoked vestibulospinal and vestibulocollic reflexes to assess whether these showed similar changes to those for startle following stroke affecting the pons and above. METHODS: Twenty-four stroke subjects (aged 40-82) were studied in the vestibulospinal part of the study, 21 stroke subjects (aged 40-81 years) were studied in the vestibulocollic part, including 18 studied in both. Transmastoid galvanic (DC) current was used to stimulate the vestibular nerve. Vestibulocollic responses were recorded from the sternocleidomastoid muscles and vestibulospinal responses from over soleus in standing subjects. RESULTS: Vestibulocollic reflex amplitudes and latencies showed no significant differences between the two sides. Similarly short latency (SL) and medium latency (ML) vestibulospinal reflexes did not differ significantly in frequency, latency or amplitude between the affected and unaffected legs. CONCLUSIONS: Vestibular reflexes are not facilitated by stroke at or above the pontine level. The exaggeration of startle by stroke may be specific to this reflex.

Vestibulocollic reflexes: normal values and the effect of age.

OBJECTIVES: To define normal values and examine the influence of ageing on vestibulocollic reflexes (VCR). METHODS: Vestibulocollic responses to 100 dB (normal hearing level; NHL) clicks, forehead taps and galvanic stimulation were measured in 70 healthy adults aged 25-85 years. RESULTS: Click- and galvanic-evoked responses were present bilaterally in all subjects below 60. Average click-evoked response amplitudes decreased with age, with a pronounced decline of 25-30% per decade from the 6th decade. The average click thresholds increased from 85 dB in the third decade to 96.5 dB in the 8th and 9th decades. Average galvanic-evoked VCR amplitudes decreased sharply from the seventh decade. Tap-evoked reflex amplitudes showed a milder decrease. When side to side differences in amplitude were expressed as asymmetry ratios (AR) in subjects below the age of 60, values of up to 35 and 46% were obtained for click amplitudes corrected and uncorrected for background electromyogram (EMG), up to 61% for both corrected and uncorrected tap response amplitudes, and up to 41 and 55% for corrected and uncorrected galvanic-evoked responses. CONCLUSIONS: A normative range of values can be specified for click- and galvanic-evoked VCRs for subjects up to the age of 60. Click- and galvanic-evoked VCR amplitudes decrease rapidly thereafter while tap-evoked responses are less affected. These changes are probably due to morphological changes in the vestibular system occurring with ageing and are more marked than in several previous reports of age-related changes in caloric responses and vestibulo-ocular reflexes.

Selective effects of ageing on vestibular-dependent lower limb responses following galvanic stimulation.

OBJECTIVES: To examine the influence of ageing on electromyographic (EMG) responses to galvanic vestibular stimulation (GVS) in the lower limbs and to define normal values. METHODS: EMG responses to 4 mA/20 ms transmastoid binaural GVS were recorded from the soleus muscles of 70 healthy adults aged 24-85 years. Short (SL) and medium latency (ML) vestibular-dependent reflex latencies and amplitudes were measured from averaged rectified EMG. Side to side differences were examined in 30 subjects, using monaural stimulation, and expressed as an asymmetry ratio (AR). RESULTS: SL reflexes to transmastoid binaural stimulation were absent in 10 of the 25 subjects over the age of 60. ML responses were present in all subjects. SL reflex amplitudes decreased from 24.4+/-12.4% (mean+/-SD) in the 3rd decade to 7.6+/-6.2% in the 8th and 9th decades. The ML reflex amplitudes showed a tendency to increase with age. The mean AR for SL reflex amplitudes increased from 12.1% in the 3rd decade to 80.7% in the 8th and 9th decades but that for ML amplitudes did not change significantly. The average SL and ML response onset latencies were 56.5+/-7.5 and 97.2+/-9.4 ms. SL onset latencies were significantly delayed with age. Both SL and ML responses were highly reproducible between experiments. CONCLUSIONS: SL and ML responses behave differently in response to age. The decrease in SL reflex amplitudes correlates well with previously described age-related morphological changes. Preservation of the ML reflex may reflect central adaptation to reduced afferent input. SL and ML responses are potentially useful measures of vestibular-spinal function, which may have a role in the assessment of older subjects with dysequilibrium. These reflexes need to be interpreted in the context of age.

Differential effect of current rise time on short and medium latency vestibulospinal reflexes.

OBJECTIVES: To investigate the effect of varying current rise time on galvanic-evoked short (SL) and medium (ML) latency vestibulospinal reflexes. METHODS: We recorded the soleus EMG of standing subjects in response to 3 mA direct current transmastoid stimulation with a series of current ramps with rise times of 0-300 ms. RESULTS: Longer current rise times significantly delayed the onset of both SL (P<<0.001) and ML (P<<0.001) vestibulospinal responses, by approximately 20 and 39 ms, respectively. The SL response amplitude was reduced with increasing rise time (P<<0.001), whereas the ML response amplitude was relatively unaffected by stimulus rise time. With very slow rise times a prolonged ML response alone was evoked. CONCLUSIONS: Both SL and ML reflexes can be evoked by changes in vestibular activity produced by transmastoid galvanic stimulation with a ramp onset. We found a differential effect of current rise time on SL and ML vestibulospinal reflexes, suggesting different potential functional roles for the two reflexes. SL reflexes can participate in the response to abrupt disturbances only. ML reflexes are evoked by both fast and slow changes in vestibular discharge and may be particularly effective for slowly-changing disturbances.

EMG responses evoked by the termination of galvanic (DC) vestibular stimulation: 'off-responses'.

OBJECTIVE: Vestibular responses in soleus electromyography (EMG) evoked by the sudden onset of galvanic (DC) stimulation ('on-responses') have been described in detail previously. The aim of the present study was to describe responses in soleus triggered by the termination of galvanic stimulation ('off-responses'). METHODS: In 10 healthy human subjects, we studied responses to transmastoid (bilateral) stimuli of 200 ms and 2 s average duration and 3 or 4 mA intensity. We obtained both on- and off-responses using the same raw data. EMG activity was recorded onto tape while current pulses of systematically varying duration were delivered. Averaged on-responses were obtained by triggering from the beginning of the current pulses. Averaged off-responses were obtained by triggering from the termination of the current pulses. RESULTS: Short-latency (SL) and medium latency (ML) off-responses were both obtained in all but one study. The SL and the ML components of the off-responses were present and had similar latencies and amplitudes, but opposite excitability, to the on-responses obtained with the same stimuli. CONCLUSIONS: Off-responses to galvanic vestibular stimulation can be recorded from soleus EMG. Our findings imply that vestibular SL and ML reflex responses in the legs are dependent on the change in the rate of vestibular nerve discharge, not its absolute level. Both on- and off-responses have properties appropriate to a role in maintaining body stability.