Effects Of Galvanic Vestibular Stimulation Research Articles

Dissociation between subjective vertical and subjective body orientation elicited by galvanic vestibular stimulation

Previous studies demonstrated that sensory stimulation could differentially affect the subjective vertical (SV) and the subjective body orientation (SBO). This suggests that the central nervous system elaborates various references of verticality in function of the task demands and of the available sensory information. In this study, we tested whether the dissociation between SV and SBO appears for a selective stimulation of the vestibular system, by using galvanic vestibular stimulation (GVS). Seated subjects performed vertical settings by controlling the orientation of a visual rod during GVS. Subjects were also instructed to evaluate the orientation of the head and trunk relative to gravity. The results revealed a large variability in the way SV and SBO were affected. In all cases, the effect of GVS on SV was not a mirror image of a distorted SBO. We propose that this dissociation is mainly determined by central processes involved in the estimation of sensory cues reliability. GVS also yielded a tilt of the head when the head was unrestrained. The results suggest that changes in actual head orientation yielded by GVS may be related to the perceived direction of gravity but cannot be explained by a compensation of an illusory orientation of the head.

Vestibulospinal influences on lower limb motoneurons

Galvanic vestibular stimulation (GVS) is a research tool used to activate the vestibular system in human subjects. When a low-intensity stimulus (1-4 mA) is delivered percutaneously to the vestibular nerve, a transient electromyographic response is observed a short time later in lower limb muscles. Typically, galvanically evoked responses are present when the test muscle is actively engaged in controlling standing balance. However, there is evidence to suggest that GVS may be able to modulate the activity of lower limb muscles when subjects are not in a free-standing situation. The purpose of this review is to examine 2 studies from our laboratory that examined the effects of GVS on the lower limb motoneuron pool. For instance, a monopolar monaural galvanic stimulus modified the amplitude of the ipsilateral soleus H-reflex. Furthermore, bipolar binaural GVS significantly altered the onset of activation and the initial firing frequency of gastrocnemius motor units. The following paper examines the effects of GVS on muscles that are not being used to maintain balance. We propose that GVS is modulating motor output by influencing the activity of presynaptic inhibitory mechanisms that act on the motoneuron pool.

Probing the human vestibular system with galvanic stimulation.

Galvanic vestibular stimulation (GVS) is a simple, safe, and specific way to elicit vestibular reflexes. Yet, despite a long history, it has only recently found popularity as a research tool and is rarely used clinically. The obstacle to advancing and exploiting GVS is that we cannot interpret the evoked responses with certainty because we do not understand how the stimulus acts as an input to the system. This paper examines the electrophysiology and anatomy of the vestibular organs and the effects of GVS on human balance control and develops a model that explains the observed balance responses. These responses are large and highly organized over all body segments and adapt to postural and balance requirements. To achieve this, neurons in the vestibular nuclei receive convergent signals from all vestibular receptors and somatosensory and cortical inputs. GVS sway responses are affected by other sources of information about balance but can appear as the sum of otolithic and semicircular canal responses. Electrophysiological studies showing similar activation of primary afferents from the otolith organs and canals and their convergence in the vestibular nuclei support this. On the basis of the morphology of the cristae and the alignment of the semicircular canals in the skull, rotational vectors calculated for every mode of GVS agree with the observed sway. However, vector summation of signals from all utricular afferents does not explain the observed sway. Thus we propose the hypothesis that the otolithic component of the balance response originates from only the pars medialis of the utricular macula.

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.

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.

Vestibular contribution to combined arm and trunk motion.

Recent studies have shown that the hand-pointing movements within arm's reach remain invariant whether the trunk is recruited or not or its motion is unexpectedly prevented. This suggests the presence of compensatory arm-trunk coordination minimizing the deflections of the hand from the intended trajectory. It has been postulated that vestibular signals elicited by the trunk motion and transmitted to the arm motor system play a major role in the compensation. One prediction of this hypothesis is that vestibular stimulation should influence arm posture and movement during reaching. It has been demonstrated that galvanic vestibular stimulation (GVS) can influence the direction of pointing movements when body motion is restrained. In the present study, we analyzed the effects of GVS on trunk-assisted pointing movements. Subjects either moved the hand to a target or maintained a steady-state posture near the target, while moving the trunk forward with the eyes closed. When GVS was applied, the final position of the hand was deviated in the lateral and sagittal direction in both tasks. This was the result of two independent effects: a deviation of the trunk trajectory and a modification of the arm position relative to the trunk. Thus, the vestibular system might be directly involved not only in the control of trunk motion but also in the arm-trunk coordination during trunk-assisted reaching movements.

Galvanic vestibular stimulation evokes sensations of body rotation.

Psychophysical experiments identified effects of galvanic vestibular stimulation (GVS) on the perception of whole-body angular rotation. Subjects lay supine on a platform that could rotate about a vertical axis through the vestibular axis so that linear movements were excluded. Movements were applied sufficiently above perception threshold to enable a reliable report of direction and movement size. In some trials, binaural GVS was applied concurrently at 1-2 mA. When GVS that was incongruent with the movement was applied, subjects reported lesser spin, on average cancelling the movement perception. When the GVS and movement were congruent, subjects reported greater spin. We conclude that GVS produces a vestibular signal of rotation, probably though an effect on semicircular canals.

Somatosensory loss increases vestibulospinal sensitivity.

To determine whether subjects with somatosensory loss show a compensatory increase in sensitivity to vestibular stimulation, we compared the amplitude of postural lean in response to four different intensities of bipolar galvanic stimulation in subjects with diabetic peripheral neuropathy (PNP) and age-matched control subjects. To determine whether healthy and neuropathic subjects show similar increases in sensitivity to galvanic vestibular stimulation when standing on unstable surfaces, both groups were exposed to galvanic stimulation while standing on a compliant foam surface. In these experiments, a 3-s pulse of galvanic current was administered to subjects standing with eyes closed and their heads turned toward one shoulder (anodal current on the forward mastoid). Anterior body tilt, as measured by center of foot pressure (CoP), increased proportionately with increasing galvanic vestibular stimulation intensity for all subjects. Subjects with peripheral neuropathy showed larger forward CoP displacement in response to galvanic stimulation than control subjects. The largest differences between neuropathy and control subjects were at the highest galvanic intensities, indicating an increased sensitivity to vestibular stimulation. Neuropathy subjects showed a larger increase in sensitivity to vestibular stimulation when standing on compliant foam than control subjects. The effect of galvanic stimulation was larger on the movement of the trunk segment in space than on the body's center of mass (CoM) angle, suggesting that the vestibular system acts to control trunk orientation rather than to control whole body posture. This study provides evidence for an increase in the sensitivity of the postural control system to vestibular stimulation when somatosensory information from the surface is disrupted either by peripheral neuropathy or by standing on an unstable surface. Simulations from a simple model of postural orientation incorporating feedback from the vestibular and somatosensory systems suggest that the increase in body lean in response to galvanic current in subjects with neuropathy could be reproduced only if central vestibular gain was increased when peripheral somatosensory gain was decreased. The larger effects of galvanic vestibular stimulation on the trunk than on the body's CoM suggest that the vestibular system may act to control postural orientation via control of the trunk in space.

Effects of galvanic vestibular stimulation during human walking.

1. To identify vestibular influences on human walking, galvanic vestibular stimulation was applied to normal adult subjects as they walked to a previously seen target. A transmastoidal step stimulus commenced as subjects started walking. With the eyes shut, the galvanic stimulus caused large turns towards the side with the anodal current. 2. Ability to perceive the trajectory of gait without visual cues was measured by guiding blindfolded subjects from one arbitrary point to another, either walking or seated in a wheelchair. On reaching a destination position and removing the blindfold, subjects pointed to indicate the starting position. Subjects made considerable errors in estimating the trajectory, but were equally accurate whether in the wheelchair or walking. 3. To determine the effects of vestibular stimulation on the perception of trajectory, the galvanic stimulus was applied to blindfolded subjects as they were guided from one point to another in the wheelchair. The vestibular stimulus produced an illusory shift in the trajectory travelled. This shift was towards the side with the cathode, i.e. in the opposite direction to the turn produced by the stimulus during walking. 4. We conclude that galvanic vestibular stimulation during walking causes subjects to turn from their planned trajectory. In part, this altered course may compensate for an altered perception of trajectory produced by the stimulus. However, altered perception of the vertical or the base of support, or direct vestibulo-fugal influences on the leg muscles could contribute to the changes in gait.

Cortical areas activated by bilateral galvanic vestibular stimulation.

The brain areas activated by bilateral galvanic vestibular stimulation (GVS) were studied using functional magnetic resonance imaging. In six human volunteers, GVS led to activation in the region of the temporoparietal junction, the central sulcus, and the anterior interior intraparietal sulcus, which may correspond to macaque areas PIVC, 3aV, and 2v, respectively. In addition, activation was found in premotor regions of the frontal lobe, presumably analogous to areas 6pa and 8a in the monkey. Since these areas were not detected in previous studies using caloric vestibular stimulation, they could be related to the modulation of otolith afferent activity by GVS. However, the simple paradigm used did not allow separation of the otolithic and semicircular canal effects of GVS. Further studies must be performed to clarify the question of cortical representation of the otolithic information in the human and monkey brain.

The timing of galvanic vestibular stimulation affects responses to platform translation

We compared the effects of galvanic vestibular stimulation applied at 0, 0.5, 1.5 and 2.5 s prior to a backward platform translation on postural responses. The effect of the galvanic stimulation was largest on the final equilibrium position of the center of pressure (CoP). The largest effects occurred for the 0.5 and 0-s pre-period, when the dynamic CoP pressure changes in response to both the galvanic stimulus and the platform translation coincided. The shift in the final equilibrium position was also larger than the sum of the shifts for the galvanic stimulus and the platform translation alone for the 0.5 and 0-s pre-periods. The initial rate of change of the CoP response to the platform translation was not significantly affected in any condition. Changes in the peak CoP position could be accounted for by local interaction of CoP velocity changes induced by the galvanic and translation responses alone, but the changes in final equilibrium position could only be accounted for by a change in global body orientation. These findings suggest that the contribution of vestibulospinal information is greatest during the dynamic phase of the postural response, and that the vestibular system contributes most to the later components of the postural response, particularly to the final equilibrium position. These findings suggest that a nonlinear interaction between the vestibular signal induced by the galvanic current and the sensory stimuli produced by the platform translation occurs when the two stimuli are presented within 1 s, during the dynamic phase of the postural response to the galvanic stimulus. When presented at greater separations in time, the stimuli appear to be treated as independent events, such that no interaction occurs.

Effects of galvanic vestibular stimulation on otolithic and semicircular canal eye movements and perceived vertical

Objective: The aim of this study was to determine the otolithic and semicircular canal effects of galvanic vestibular stimulation with increasing current strengths on eye movements and the perception of verticality. Methods: We measured (1) 3-dimensional eye movements, (2) subjective tilt of the peripheral visual field, and (3) subjective tilt of a central vertical line in 12 healthy subjects during galvanic vestibular stimulation. A rectangular, unipolar binaural electric current was applied to each subject's mastoid. Results: Anodal stimulation of the right mastoid led to an ipsiversive tonic ocular torsion of up to 5.4°, to a contralateral tilt of both the peripheral visual field (1–9°), and a central vertical line (0.5–6.2°) increasing in amplitude with increasing current strengths applied. This reflects otolith stimulation. In most subjects, current strengths of 3 mA or more elicited a slight (horizontal-) torsional nystagmus (amplitude 1–2°) that was superimposed on static torsion. This reflects horizontal and vertical semicircular canal stimulation. A correlation was found in the amount of the 3 measured parameters and the strength of the applied current. Conclusions: Thus, galvanic vestibular stimulation at low current intensities (1– 3 mA) preferably excites otolith responses, which increase with increasing current intensity. With higher current intensity above 3 mA, additional semicircular canal responses are elicited in the form of horizontal-rotatory nystagmus superimposed on static torsional deviations. The lack of a vertical deviation and nystagmus can be explained by the counterdirected vertical components of the anterior and posterior semicircular canal.

Effect of galvanic vestibular stimulation on perception of subjective vertical in standing humans.

The present work reinvestigated the influence of bimastoidal galvanic vestibular stimulation (0.4 mA during 10 sec.) on subjective vertical. We tested the hypothesis that deviations are directed towards the anode side, like postural tilt evoked by galvanic vestibular stimulation. 15 subjects were instructed to orient vertically in darkness a light-rod during 3 experimental conditions of control, anode right, and anode left. The statistical analysis showed that the perception of the vertical was modified according to the experimental conditions and the subjects. Angular deviations occurred toward the anode side. The results are interpreted as a consequence of a modification of the central representation of the vertical or of ocular torsion directed towards the anode side and likely to induce an apparent displacement of the rod towards the cathode side.

Effect of galvanic vestibular stimulation on human postural responses during support surface translations

1. We investigated the role of the vestibular system in postural control by combining galvanic vestibular stimulation (0.2-0.5 mA) with platform translations in standing subjects. Vestibular stimulation delivered 500 ms before and continuously during the platform translation produced little change in the earliest center of pressure (COP) and center of mass (COM) movements in response to platform translations, but resulted in large changes during the execution of the postural movement and in the final equilibrium position. 2. Vestibular stimulation produced anterior or posterior shifts in the position of COP and COM, depending on the polarity of the galvanic current. These shifts were larger during platform translations than during quiet stance. The peak of these shifts in COP and COM occurred at 1.5-2.5 s after the onset of platform translation, and increased in magnitude with increasing platform velocity. The final equilibrium positions of COP and COM were also shifted, but these shifts were smaller and not dependent on platform velocity. 3. These results imply that a tonic step of galvanic current to the vestibular system can change the final equilibrium position for an automatic postural response. Furthermore, these results indicate that the vestibular system may play a larger role in interpreting sensory reafference during postural movements, and especially during fast postural movements, than in controlling quiet stance. Finally, these results indicate that the vestibular system does not play a critical role in triggering the earliest postural responses, but it may be critical in establishing an internal reference for verticality.

Effect of galvanic vestibular stimulation on rotation testing.

Effect of galvanic vestibular stimulation on rotation testing.