Between-subject variability and within-subject reliability of the human eye-movement response to bilateral galvanic (DC) vestibular stimulation

Recent studies have shown that responses to surface galvanic vestibular stimulation (GVS) show substantial interindividual variation. Between-subject variability may be due to individual differences between subjects, or to the poor reliability of the test, or to differences in test details, or to host factors. The aim of the present study was to compare variability between and within subjects in binocular 3-D eye-movement responses to long-duration, maintained, large-amplitude, bilateral, bipolar, surface GVS. Subjects were seated and restrained, and in one condition fixated a small, centrally located visual target; in the other condition, testing was carried out in complete darkness. Surface GVS of 5 mA, with a rectangular waveform was delivered bilaterally for 5 min while eye movements were measured using computerised video-oculography (VTM). In the first experiment, ten subjects participated in both conditions in one session, and in the second experiment, two subjects participated in both conditions for a total of five repeated sessions. The stimulation was well tolerated by all subjects and produced a change in torsional position with the upper pole of both eyes rolling towards the anode and away from the cathode in all subjects in both conditions. Although little vertical nystagmus was evident in either condition, most subjects showed relatively strong horizontal nystagmus (slow phases towards the anode) in darkness. This study confirms previous observations that the torsional response to GVS is highly variable between subjects, whilst also showing for the first time that eye-movement responses to GVS show good within-subject repeatability. This study also demonstrates considerable between-subject variability in the relative ratios of response components (torsional and horizontal nystagmus, torsional position), whereas the relatively small within-subject variability can be characterised more by changes in the overall amplitude of the eye-movement response. Subjects show idiosyncratic oculomotor response patterns to GVS, varying slightly in absolute magnitude between sessions. Thus, GVS may be a more reliable stimulus than may have been anticipated from the literature.     

Short-latency eye movements evoked by near-threshold galvanic vestibular stimulation

To investigate whether the primary planes of eye and body responses to galvanic vestibular stimulation (GVS) are congruent, we have measured the binocular, three-dimensional eye movements (scleral coil technique) to bilateral bipolar GVS in six normal human subjects. Stimulation intensities were kept deliberately low in order to characterize the response to near-threshold intensities of stimulation (0.1–0.9 mA) that had been used previously to characterise body postural responses. Stimuli were applied for 4 s, but only the early responses that occurred within the initial 300 ms of turning the current on or off were measured. At intensities of 0.1–0.7 mA the 'on' response consisted almost exclusively of a torsional slow phase eye movement in which the top of the eyes rotated towards the anode. The latency of the torsional response was ca. 46 ms. A weak polarity-dependent disconjugate response was also observed in which the intorting eye elevated and the extorting eye depressed ('skew eye deviation'). When the current was turned off similar responses occurred in the reverse direction. Removal of the visual fixation light-emitting diode (LED) had no consistent effect on the short-latency ocular responses. The direction of the ocular response was similar to that of the postural response and is compatible with GVS inducing an apparent dynamic roll-tilt of the head towards the cathode. However, weak horizontal eye movements, which became more prominent as the stimulus intensity was increased to 0.9 mA, were also observed. This suggests that an additional weak rotational component about the yaw axis, or a component of lateral translation in the frontal plane, is contained in the GVS-evoked signal. The overall pattern of eye movement suggests that semicircular canal afferents contribute to these low-intensity GVS responses. Electronic Publication     

Comparison of human ocular torsion patterns during natural and galvanic vestibular stimulation

Galvanic vestibular stimulation (GVS) is reported to induce interindividually variable tonic ocular torsion (OT) and superimposed torsional nystagmus. It has been proposed that the tonic component results from the activation of otolith afferents. We tested our hypothesis that both the tonic and the phasic OT are mainly due to semicircular canal (SCC) stimulation by examining whether the OT patterns elicited by GVS can be reproduced by pure SCC stimulations. Using videooculography we measured the OT of six healthy subjects while two different stimuli with a duration of 20 s were applied: 1) transmastoidal GVS steps of 2 mA with the head in a pitched nose-down position and 2) angular head rotations around a combined roll-yaw axis parallel to the gravity vector with the head in the same position. The stimulation profile was individually scaled to match the nystagmus properties from GVS and consisted of a sustained velocity step of 4-12 degrees /s on which a velocity ramp of 0.67-2 degrees /s(2) was superimposed. Since blinks were reported to induce transient torsional eye movements, the subjects were also asked to blink once 10 s after stimulus onset. Analysis of torsional eye movements under both conditions revealed no significant differences. Thus we conclude that both the tonic and the phasic OT responses to GVS can be reproduced by pure rotational stimulations and that the OT-related effects of GVS on SCC afferents are similar to natural stimulations at small amplitudes.     

Direct perturbation of neural integrator by bilateral galvanic vestibular stimulation

Caloric vestibular stimulation (CVS) and galvanic vestibular stimulation (GVS) act primarily on the peripheral vestibular system. Although the electrical current applied during GVS is thought to flow through peripheral vestibular organs, some current may spread into areas within the central nervous system, particularly when the bilateral galvanic vestibular stimulation (bGVS) method is used. According to Alexander’s law, the magnitude of nystagmus increases with eccentric gaze movement, due to the function of the neural integrator (NI); thus, if the information for vestibular stimulation corresponds to Alexander’s law, the peripheral vestibular organ is stimulated. Therefore, it would appear that if CVS results comply with Alexander’s law, and bGVS results do not, the sites stimulated by bGVS are not perfectly located in the peripheral vestibular area. In our experiments on normal human subjects, the magnitude of nystagmus under CVS increased with rising gaze eccentricity in the direction that the magnitude of the nystagmus increases, and this change was found to follow Alexander’s law. However, in the case of nystagmus under bGVS, results did not follow Alexander’s law. In addition, study of the influences of bGVS at different current intensities on nystagmus magnitude showed that bGVS at 5 mA distorted nystagmus magnitude more than at 3 mA, which suggests bGVS acts not only on the peripheral vestibular nerves, but also on some areas of the central nervous system, particularly the NI. According to our experiments, bGVS directly affects neural integrator function.     

Latency and initiation of the human vestibuloocular reflex to pulsed galvanic stimulation

Cathodal galvanic currents activate primary vestibular afferents, whereas anodal currents inhibit them. Pulsed galvanic vestibular stimulation (GVS) was used to determine the latency and initiation of the human vestibuloocular reflex. Three-dimensional galvanic vestibuloocular reflex (g-VOR) was recorded with binocular dual-search coils in response to a bilateral bipolar 100-ms rectangular pulse of current at 0.9 (near-threshold), 2.5, 5.0, 7.5, and 10.0 mA in 11 normal subjects. The g-VOR consisted of three components: conjugate torsional eye rotation away from cathode toward anode; vertical divergence (skew deviation) with hypertropia of the eye on the cathodal and hypotropia of the eye on the anodal sides; and conjugate horizontal eye rotation away from cathode toward anode. The g-VOR was repeatable across all subjects, its magnitude a linear function of the current intensity, its latency about 9.0 ms with GVS of >or=2.5 mA, and was not suppressed by visual fixation. At 10-mA stimulation, the g-VOR [x, y, z] on the cathodal side was [0.77 +/- 0.10, -0.05 +/- 0.05, -0.18 +/- 0.06 degrees ] (mean +/- 95% confidence intervals) and on the anodal side was [0.79 +/- 0.10, 0.16 +/- 0.05, -0.19 +/- 0.06 degrees ], with a vertical divergence of 0.20 degrees . Although the horizontal g-VOR could have arisen from activation of the horizontal semicircular canal afferents, the vertical-torsional g-VOR resembled the vestibuloocular reflex in response to roll-plane head rotation about an Earth-horizontal axis and might be a result of both vertical semicircular canal and otolith afferent activations. Pulsed GVS is a promising technique to investigate latency and initiation of the human vestibuloocular reflex because it does not require a large mechanical apparatus nor does it pose problems of head inertia or slippage.     

Maintained ocular torsion produced by bilateral and unilateral galvanic (DC) vestibular stimulation in humans

 This study was designed to measure ocular movements evoked by galvanic (DC) stimulation using computerised video-oculography. Long duration (>30 s) galvanic vestibular stimulation at currents of up to 5 mA through large-area surface electrodes over the mastoid processes causes maintained changes in the ocular torsional position of both eyes in healthy human subjects. With the subject seated and the head held firmly, torsion was measured by a computer-based image-processing system (VTM). Torsion was recorded in darkness, with or without a single fixation point. With bilateral stimulation, the upper poles of both eyes always torted away from the side of cathode placement and toward the anode. For unilateral stimulation, torsion was directed away from the cathode or toward the anode. The magnitude of ocular torsion was dependent on current strength: with bilateral stimulation the peak torsion was on average 2.88° for 5-mA current intensity compared with 1.58° for 3 mA. A smaller amplitude of torsion was obtained for unilateral stimulation. The average peak torsion was the same for both eyes for all forms of stimulation. Our findings indicate that low-intensity galvanic stimulation evokes ocular torsion in normal subjects, an effect which is consistent with an action on otolith afferents. Received: 27 March 1998 / Accepted: 23 June 1998     

Effects of galvanic vestibular stimulation on human posture and perception while standing

This study examines three hypotheses that have been proposed to explain the effects of galvanic vestibular stimulation (GVS) in standing human subjects. The first assumes realignment to an altered representation of vertical. GVS-evoked body tilt produced under conditions of different stability was compared with perceptions of the vertical which subjects indicated by two means, a visual line and a manipulandum. GVS produced body tilt that increased in unstable conditions but there were no differences in the perceived vertical in any condition. The second hypothesis is that the altered vestibular signal is interpreted as a tilt of the support surface. The postural response evoked by tilting the support surface was compared with the GVS response under conditions of varying stability. These responses were different, particularly for the lower body where movements were oppositely directed. Standing on foam augmented GVS responses whereas standing with feet apart augmented platform-tilt responses. The third hypothesis is that GVS produces an illusion of movement, and this causes a reaction in the opposite direction. Perception of movement during GVS was determined in standing and immobilised subjects. Although immobilised subjects experienced illusions of movement in the direction opposite the sway response, this only happened after long periods of stimulation and never for standing where subjects accurately reported the true direction of sway. Thus, the results do not support any of these proposals. Instead, they and other observations support a simpler interpretation that the GVS signal is consistent with head movement and evokes an automated response to stabilise the head in space.     

Unilateral testing of utricular function

A modified rotatory chair test is reported in which radial acceleration, generated by eccentric displacement of the subject during constant angular velocity, is exploited as a unilateral stimulation to the otolith organs. During constant angular rate rotation, the test subject is displaced laterally on the rotating turntable by 3.5 cm, so that one labyrinth becomes aligned with the rotatory axis while the second – eccentric – labyrinth is solely exposed to the altered gravito-inertial acceleration (GIA). Previously reported results showed that the direction of the response is independent of the direction of turntable rotation, ruling out any canal influence, and indicated that in a normal population the response, measured in one eye, was symmetrical for displacement of the left and right labyrinths. This mode of stimulus thus appears to elicit a unilateral otolith-ocular response (OOR). Examination of this unilateral OOR was extended in the present study; comparative testing with head-tilt to gravity, i.e. involving bilateral stimulation to the otolith organs, was carried out. Movements of both eyes were recorded (by three-dimensional video-oculography), in order to examine response conjugacy. To verify the specificity of the unilateral stimulus, tests were performed with patients who had previously undergone unilateral section of the vestibular nerve as treatment for acoustic neuroma. The eccentric displacement profile (EDP) and head-tilt stimulus each included ten cycles of left-right oscillation in order to permit signal averaging. In the normal subjects (n=12) the torsional component of the OOR proved to be both labyrinth-symmetrical and conjugate, during both bilateral and unilateral otolith stimulation. OOR gain (ocular torsion/GIA tilt) was higher for bilateral than unilateral stimulation. Bilateral OORs, obtained from three of the five unilaterally deafferented patients, proved less symmetrical and conjugate than in the normals. Unilateral OORs in all five patients were characteristically asymmetrical, with little or no response during stimulation of the diseased labyrinth. Received: 28 July 1997 / Accepted: 3 February 1998     

Deficits in vertical and torsional eye movements after uni- and bilateral muscimol inactivation of the interstitial nucleus of Cajal of the alert monkey

The mesencephalic interstitial nucleus of Cajal (iC) is considered the neural integrator for vertical and torsional eye movements and has also been proposed to be involved in saccade generation. The aim of this study was to elucidate the function of iC in neural integration of different types of eye movements and to distinguish eye movement deficits due to iC impairment from that of the immediately adjacent rostral interstitial nucleus of the medial longitudinal fasciculus (riMLF). We addressed the following questions: (1) According to the neural integrator hypothesis, all eye movements including the saccadic system and the vestibulo-ocular reflex (VOR) share a common neural integrator. Do iC lesions impair gaze-holding function for vertical and torsional eye positions and the torsional and vertical VOR gain to a similar degree? (2) What are the dynamic properties of vertical and torsional eye movements deficits after iC lesions, e.g., the specificity of torsional and vertical nystagmus? (3) Is iC involved in saccade generation? We performed 13 uni- and three bilateral iC inactivations by muscimol microinjections in four alert monkeys. Three-dimensional eye movements were studied under head-stationary conditions during vertical and torsional VOR. Under static conditions, unilateral iC injections evoked a shift of Listing’s plane to the contralesional side (up to 20°), which increased (ipsilesional ear down) or decreased (ipsilesional ear up) by additional static vestibular stimulation in the roll plane, i.e., ocular counterroll was preserved. The monkeys showed a spontaneous torsional nystagmus with a profound downbeat component. The fast phases of torsional nystagmus always beat toward the lesion side (ipsilesional). Pronounced gaze-holding deficit for torsional and vertical eye positions (neural integrator failure) was reflected by the reduction of time constants of the exponential decay of the slow phase to 330–370 ms. Whereas the vertical oculomotor range was profoundly decreased (up to 50%) and vertical saccades were reduced in amplitude, saccade velocity remained normal and horizontal eye movements were not affected. Bilateral iC injections reduced the shift of Listing’s plane caused by unilateral injections, i.e., back toward the plane of zero torsion. Torsional nystagmus reversed its direction and ceased, whereas vertical nystagmus persisted. In contrast to unilateral injection, there was additional upbeating nystagmus. Time constants of the position integrator of the gaze-holding system did not differ between unilateral and bilateral injections. The range of stable vertical eye positions and saccade amplitude was smaller when compared with unilateral injections, but the main sequence remained normal. Dynamic vestibular stimulation after unilateral iC injections had virtually no effect on torsional and vertical VOR gain and phase at the same time when time constants already indicated severe integrator failure. Torsional VOR elicited a constant slow-phase velocity offset up to 30° toward the contralesional side, i.e., in the opposite direction to spontaneous torsional nystagmus. Likewise, vertical VOR showed a velocity offset in an upward direction, i.e., opposite to the spontaneous downbeat nystagmus. Contralesional torsional and upward vertical quick phases were missing or severely reduced in amplitude but showed normal velocity. In contrast, bilateral iC injections reduced the gain of the torsional and vertical VOR by 50% and caused a phase lead of 10–20° (eye compared with head velocity). We propose that the slow-phase velocity offset during torsional and vertical VOR reflects a vestibular imbalance. It therefore appears likely that the vertical and torsional nystagmus after iC lesions is not only caused by a neural integrator failure but also by a vestibular imbalance. Unilateral iC injections have clearly differential effects on the VOR and the gaze-holding function. These results are not compatible with a single common neural integrator model, which would predict a much stronger VOR gain reduction and phase advance, as found in our data. Our data support the existence of multiple integrators in iC with parallel processing.     

Orientation of the body response to galvanic stimulation as a function of the inter-vestibular imbalance

We proposed to study and quantify the anteroposterior component, on top of the lateral one, of the body sway induced by different configurations of galvanic vestibular stimulation (GVS) in order to advance the understanding of the orientation of the response. Four stimulation configurations were used in two separate experiments: monaural, binaural, and opposite double monaural in the first experiment (11 subjects); monaural and double monaural in the second (13 subjects). The postural response of the subjects, standing with their eyes closed, to the stimulus (0.6 mA, 4 s) was assessed by measuring the displacement of the center of pressure (CoP) using a force platform. As usual, binaural GVS induced a strictly lateral deviation of the center of pressure. The opposite double monaural condition induced a similar lateral sway to that obtained in the binaural mode, although with a very different stimulation configuration. Monaural GVS induced an oblique, stereotyped deviation in each subject. The anteroposterior component comprised a forward deviation when the anode was on the forehead and a backward deviation when the anode was on the mastoid. The lateral component, directed towards the anode as in the binaural design, was twice as large in the binaural than in the monaural mode. The second experiment showed that double monaural stimulation elicited an anteroposterior deviation (backwards when the anode was on the mastoids and forwards when it was on the forehead) that was equivalent to the addition of two complementary monaural configurations. The present results show that monaural stimulation activates one side of the vestibular apparatus and induces reproducible, stereotyped deviations of the CoP in both the anteroposterior and lateral plane. Secondly, binaural GVS appears to result from the addition of two complementary monaural stimulations. Lateral components of the response to each stimulation, being in the same direction, are summed, whilst anteroposterior components, being in opposite directions, cancel each other out. The opposite happens when both labyrinths are polarized in the same way, as in the double monaural configuration. We suggest that the orientation of the response to GVS is a function of the imbalance between right and left vestibular polarization, rather than a function of the actual position of the electrodes.     

The human horizontal vestibulo-ocular reflex in response to high-acceleration stimulation before and after unilateral vestibular neurectomy

Summary The normal horizontal vestibulo-ocular reflex (HVOR) is largely generated by simultaneous stimulation of the two horizontal semicircular canals (HSCCs). To determine the dynamics of the HVOR when it is generated by only one HSCC, compensatory eye movements in response to a novel vestibular stimulus were measured using magnetic search coils. The vestibular stimulus consisted of low-amplitude, high-acceleration, passive, unpredictable, horizontal rotations of the head with respect to the trunk. While these so called head “impulses” had amplitudes of only 15–20 degrees with peak velocities up to 250 deg/s, they had peak accelerations up to 3000 deg/s/s. Fourteen humans were studied in this way before and after therapeutic unilateral vestibular neurectomy; 10 were studied 1 week or 1 year afterwards; 4 were studied 1 week and 1 year afterwards. The results from these 14 patients were compared with the results from 30 normal control subjects and with the results from one subject with absent vestibular function following bilateral vestibular neurectomy. Compensatory eye rotation in normal subjects closely mirrored head rotation. In contrast there was no compensatory eye rotation in the first 170 ms after the onset of head rotation in the subject without vestibular function. Before unilateral vestibular neurectomy all the patients' eye movement responses were within the normal control range. One week after unilateral vestibular neurectomy however there was a symmetrical bilateral HVOR deficit. The asymmetry was much more profound than has been shown in any previous studies. The HVOR generated in response to head impulses directed away from the intact side largely by ampullofugal disfacilitation from the single intact HSCC (ignoring for the moment the small contribution to the HVOR from stimulation of the vertical SCCs), was severely deficient with an average gain (eye velocity/head velocity) of 0.25 at 122.5 deg/sec head velocity (normal gain=0.94+/−0.08). In contrast the HVOR generated in response to head impulses directed toward the intact side, largely by ampullopetal excitation from the single intact HSCC, was only mildly (but nonetheless significantly) deficient, with an average gain of 0.80 at 122.5 deg/sec head velocity. At these accelerations there was no significant improvement in the average HVOR velocity gain in either direction over the following year. These results indicate that ampullopetal excitation from one HSCC can, even in the absence of ampullofugal disfacilitation from the opposite HSCC, generate a near normal HVOR in response to high-acceleration stimulation. Furthermore, since ampullofugal disfacilitation on its own, can only generate an inadequate HVOR in response to high-acceleration stimulation, it may under some normal circumstances make little contribution to the bilaterally generated HVOR.     

Vestibular contributions across the execution of a voluntary forward step

This work addressed the influence of information arising from the vestibular system on the dynamic control of a forward step. Six subjects performed the stepping task with their eyes closed under three conditions of bipolar, binaural galvanic vestibular stimulation (GVS), including (1) no GVS, (2) GVS with the anode electrode on the side of the swing limb, and (3) GVS with the anode electrode on the side of the stance limb. GVS was delivered 1,500 ms prior to a cue to step. Ground reaction forces were collected from three force platforms and movement was recorded from IRED markers placed bilaterally on the body. The results showed that, following slight deviations caused by GVS onset, the step initiation behaviour was unaffected, but lateral deviations were found during the latter, more dynamic, phases of stepping for centre of mass trajectories, time integrals of the centre of pressure displacement and upper body roll. These findings showed that vestibular information is used differently across the execution of a step without vision. While the initiation phase is run in a feedforward manner without vestibular influence, vestibular information appears to be upregulated during the more dynamic phases. Also, the level of up-regulation may be different across step execution.     

A Short Latency Vestibulomasseteric Reflex Evoked by Electrical Stimulation Over the Mastoid in Healthy Humans

We describe EMG responses recorded in active masseter muscles following unilateral and bilateral electrical vestibular stimulation (EVS, current pulses of 5 mA intensity, 2 ms duration, 3 Hz frequency). Averaged responses in unrectified masseter EMG induced by unilateral EVS were examined in 16 healthy subjects; effects induced by bilateral (transmastoid) stimulation were studied in 10 subjects. Results showed that unilateral as well as bilateral EVS induces bilaterally a clear biphasic response (onset latency ranging from 7.2 to 8.8 ms), that is of equal amplitude and latency contra- and ipsilateral to the stimulation site. In all subjects, unilateral cathodal stimulation induced a positive—negative response termed p11/n15 according to its mean peak latency; the anodal stimulation induced a response of opposite polarity (n11/p15) in 11/16 subjects. Cathodal responses were significantly larger than anodal responses. Bilateral stimulation induced a p11/n15 response significantly larger than that induced by the unilateral cathodal stimulation. Recordings from single motor units showed that responses to cathodal stimulation corresponded to a brief (2–4 ms) silent period in motor unit discharge rate. The magnitude of EVS-induced masseter response was linearly related to current intensity and scaled with the mean level of EMG activity. The size of the p11/n15 response was asymmetrically modulated when subjects were tilted on both sides; in contrast head rotation did not exert any influence. Control experiments excluded a possible role of cutaneous receptors in generating the masseter response. We conclude that transmastoid electrical stimulation evokes vestibulomasseteric reflexes in healthy humans at latencies consistent with a di-trisynaptic pathway.     

Visual-vestibular interactions in postural control during the execution of a dynamic task

The purpose of this experiment was to determine the interaction between visual and vestibular information during the transition from quiet standing to the completion of a forward step. Six subjects were asked to take one step forward at the sound of an audio tone, with their eyes open or closed, and terminate the step in a standing position. During stimulation trials, galvanic vestibular stimulation (GVS) was delivered 1500 ms before the auditory cue. GVS was delivered at an intensity three-fold that of each subject's quiet stance threshold with either stimulus right, left or no stimulation. Force data were collected from three forceplates for the calculation of centre of pressure (CoP), and kinematic data were used to calculate centre of mass (CoM) and body trajectories. In quiet stance all subjects responded to the GVS perturbation by demonstrating upper body segment roll and whole body sway towards the anode electrode. Unexpectedly, in the presence of vision during quiet stance, the upper body roll response was not attenuated, even though the CoP sway patterns were reduced when vision was available. During the initiation phase of the step, despite ongoing GVS stimulation, there were no significant effects seen in CoM, CoP or upper body roll responses. During step execution, however, both CoM displacement and upper body roll demonstrated significant effects and both responses were significantly reduced when subjects' eyes were open. Analysis of the medio-lateral CoP integrals also indicated a strong stimulation effect between conditions late in the execution phase, which were largely attenuated with vision. The results suggest that the importance of visual and vestibular information varies depending on the phase of the task. In addition, the different integration between visual and vestibular input during quiet standing suggests a dual role for vestibular information. We propose that vestibular information in quiet standing has a role in maintaining whole body postural stability, as well as playing an integral role in the alignment of the body segments in preparation for proper movement execution. Vision was demonstrated to differentially attenuate these responses based on the phase of the task. Thus, visual and vestibular information appear to be integrated differently across the different phases of a forward-stepping task.     

Position and velocity responses to galvanic vestibular stimulation in human subjects during standing

Galvanic vestibular stimulation (GVS) in animals modulates the firing of otolith and semicircular canal afferents alike. Here, we look for postural responses evoked by GVS from the otolith organs and semicircular canals. To minimise the modifying effects of somatosensory input on the response, low-intensity (0.3–0.5 mA) GVS was applied for 8 s while subjects stood on foam rubber with the feet together and strapped to the floor. The response had three phases: (i) a rapid movement during the first second, (ii) a slower movement that persisted throughout the stimulus, and (iii) a rapid partial return movement after GVS stopped. The three movement velocities were significantly different. The GVS response therefore appears to be the sum of a step response that returns to the starting point when the stimulus stops, and a constant-velocity ramp response for the duration of the stimulus without a return movement. Subjects' responses differed in size and profile, some with the step or ramp responses almost exclusively but most with a combination of both. The 'step-plus-ramp' model was tested by comparing the three velocities. If the responses add, the initial velocity should not be different from the sum of the velocities during the ramp-only period and the step-only period at offset. ANOVA and pairwise comparisons confirmed this. It is concluded that postural responses to GVS arise through stimulation of both otolith and canal afferents.     

Is the use of vestibular information weighted differently across the initiation of walking?

The purpose of this experiment was to examine vestibular contributions at specific times during the initiation of walking in human subjects. Subjects began walking forward at the sound of an auditory tone, with vision present or occluded. Galvanic vestibular stimulation (GVS) was delivered with the anode electrode on the right or left side at either: (1) onset of the anticipatory postural adjustment (APA), (2) toe-off of the first swing limb (TO) or (3) heel contact of the first swing limb (HC). Ground reaction forces and kinematic data were collected. Upper body (roll angles from head, trunk and pelvis) and lower body (foot placement) data were analysed to determine whether the timing and magnitude of the response to GVS, and therefore the level of vestibular contribution, was modulated at different points during the initiation of gait. With vision present and occluded, the magnitude of the lower body response varied depending on the event in the gait cycle at which the stimulation was delivered. These novel results demonstrate evidence that vestibular weighting during gait initiation is dependent upon the specific gait initiation events. Upper body roll also exhibited magnitude differences between events. However, these changes are proposed to occur due to the transition from a stationary position into a dynamic state, prompting the increased weighting of vestibular information. With vision present no significant changes were seen in the segment roll response. The observations suggest a distinction in vestibular regulation of upper body roll versus foot placement for successful completion of the gait initiation task. Changes in upper body roll are influenced by the dynamic nature of the task, whereas foot placement changes are modulated based on the event during gait initiation at which GVS is delivered.     

The regulation of vestibular afferent information during monocular vision while standing

The purpose of this study was to investigate the contribution of the vestibular system to postural control during monocular vision using binaural-bipolar galvanic vestibular stimulation (GVS). Four visual (both eyes, dominant eye, non-dominant eye, and no vision) conditions were tested during GVS in five healthy subjects while focusing on a target placed in front of them. GVS evoked similar upper body postural sway during both monocular and no vision conditions that were significantly greater to those during binocular vision. Changes in ground reaction forces to the anode side followed that same trend, although data for vision with the dominant eye were not significantly different from that for binocular vision. These data suggest an increase in the weighting of vestibular afferent information during monocular vision for standing postural control.     

Human standing and walking: comparison of the effects of stimulation of the vestibular system

The adoption of bipedalism by hominids including man has complicated the tasks of balance control and the minimisation of body sway. We have investigated the role of the vestibular organs in controlling sway in the roll direction using galvanic vestibular stimulation (GVS). Two stance conditions were studied: during forward lean posterior compartment muscles are activated and during backward lean anterior compartment muscles are activated. GVS-evoked vestibular signals in stance control leg muscles as a group: all the active muscles in the leg on the GVS cathode side are excited together and those in the contralateral leg (anode side) relax. The subject sways towards the anode side. During treadmill walking, vestibular actions are subtly different: the actions are largely restricted to muscles acting at the ankle joint, occur at longer latencies, are not reciprocal in the opposite limb, are modulated throughout the step cycle (largest early in stance) and are reversed in sign in the peroneus longus muscle. The subject deviates towards the anode side. Hand contact with a firm object reduces GVS-evoked responses in leg muscles during treadmill walking. Responses to GVS are observed during over-ground walking but not significantly during bicycling on an ergometer. The observations suggest that these vestibular actions are part of a roll stabilisation mechanism. They may be mediated through different spinal premotor mechanisms during standing and walking and turned off during bicycling, when leg muscles have no balance control function.     

Otolith and canal reflexes in human standing

We used galvanic vestibular stimulation (GVS) to identify human balance reflexes of the semicircular canals and otolith organs. The experiment used a model of vestibular signals arising from GVS modulation of the net signal from vestibular afferents. With the head upright, the model predicts that the GVS-evoked canal signal indicates lateral head rotation while the otolith signal indicates lateral tilt or acceleration. Both signify body sway transverse to the head. With the head bent forward, the model predicts that the canal signal indicates body spin about a vertical axis but the otolith signal still signifies lateral body motion. Thus, we compared electromyograms (EMG) in the leg muscles and body sway evoked by GVS when subjects stood with the head upright or bent forward. With the head upright, GVS evoked a large sway in the direction of the anodal electrode. This response was abolished with the head bent forward leaving only small, oppositely directed, transient responses at the start and end of the stimulus. With the head upright, GVS evoked short-latency (60–70 ms), followed by medium-latency (120 ms) EMG responses, of opposite polarity. Bending the head forward abolished the medium-latency but preserved the short-latency response. This is compatible with GVS evoking separate otolithic and canal reflexes, indicating that balance is controlled by independent canal and otolith reflexes, probably through different pathways. We propose that the short-latency reflex and small transient sway are driven by the otolith organs and the medium-latency response and the large sway are driven by the semicircular canals.     

Modeling postural instability with Galvanic vestibular stimulation

In this study the effect of a pseudorandom binaural bipolar Galvanic stimulus generated by a sum of nonharmonically related sine waves on postural control was functionally assessed using computerized dynamic posturography (CDP), and the results compared to vestibulopathic patient populations and astronauts exposed to microgravity. The standardized CDP test battery comprised six sensory organization tests (SOTs) that combined three visual conditions (eyes open, eyes closed, and sway-referenced vision) with two proprioceptive conditions (fixed and sway-referenced support surfaces). Subjects (12) performed 18 randomized trials (three trials of each of the six SOTs) as a baseline, repeated the 18 trials with Galvanic vestibular stimulation (GVS), and then performed a post-GVS baseline. A 10 min rest period was inserted between each test battery. Anterioposterior postural sway increased significantly and was in the abnormal range (fifth percentile) during GVS for SOTs where visual input was compromised (sway-referenced surround) or absent. Postural stability returned to baseline when GVS was removed. An analysis of sensory input scores (somatosensory, visual, and vestibular) demonstrated the specificity of GVS in distorting vestibular input to postural control. The SOT scores observed in astronauts on landing day did not differ significantly to that generated by GVS in our normal subjects. GVS also induced a similar pattern of instability on CDP as profound bilateral vestibular loss, although not as severe. The results suggest that unpredictably varying GVS quantitatively and qualitatively models postural instability of vestibular origin.