1. We have studied the effects of changes in posture on the motor response to galvanic vestibular stimulation (GVS). The purpose of the experiments was to investigate whether the function of the GVS-evoked response is to stabilize the body or the head in space. Subjects faced forwards with eyes closed standing with various stance widths and sitting. In all cases the GVS-evoked response consisted of a sway of the body towards the anodal ear. 2. In the first set of experiments the response was measured from changes in (i) electromyographic activity of hip and ankle muscles, (ii) the lateral ground reaction force, and (iii) lateral motion of the body at the level of the neck (C7). For all measurements the response became smaller as the feet were placed further apart. 3. In the second set of experiments we measured the GVS-evoked tilts of the head, torso and pelvis. The basic response consisted of a tilt in space (anodal ear down) of all three segments. The head tilted more than the trunk and the trunk tilted more than the pelvis producing a leaning and bending of the body towards the anodal ear. This change in posture was sustained for the duration of the stimulus. 4. The tilt of all three segments was reduced by increasing the stance width. This was due to a reduction in evoked tilt of the pelvis, the bending of the upper body remaining relatively unchanged. Changing from a standing to a sitting posture produced additional reductions in tilt by reducing the degree of upper body bending. 5. The results indicate that the response is organized to stabilize the body rather than the head in space. We suggest that GVS produces a vestibular input akin to that experienced on an inclined support surface and that the function of the response is to counter any threat to balance by keeping the centre of mass of the body within safe limits.Passing a small current across the mastoid processes stimulates the vestibular system and in standing subjects evokes a sway of the body. This 'galvanic vestibular stimulation' (GVS) acts by modulating the spontaneous firing of vestibular afferents, increasing their firing frequency on

We assessed the effect of directed attention on early neurophysiological indices of face processing, measuring the N170 event-related potential (ERP). Twelve subjects were tested on two tasks each in which they attended either to eyes only or to faces with eyes closed, presented within series of facial and control stimuli. Consistent with the ERP literature, N170 was recorded to facial stimuli at posterior temporal electrodes and a concomitant positive peak at the vertex, with latencies around 150 ms for faces and 174 ms for eyes. However, unlike fMRI studies, neither the latency nor the amplitude of the peaks were sensitive to the target/non-target status of either the eyes or the face stimuli. This suggests that early stages of face processing indexed by N 170 are automatic and unmodified by selective attention.

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.

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.

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Visuo-vestibular integration is crucial for locomotion, yet the cortical mechanisms involved remain poorly understood. We combined binaural monopolar galvanic vestibular stimulation (GVS) and functional magnetic resonance imaging (fMRI) to characterize the cortical networks activated during antero-posterior and lateral stimulations in humans. We focused on functional areas that selectively respond to egomotion-consistent optic flow patterns: the human middle temporal complex (hMT+), V6, the ventral intraparietal (VIP) area, the cingulate sulcus visual (CSv) area and the posterior insular cortex (PIC). Areas hMT+, CSv, and PIC were equivalently responsive during lateral and anteroposterior GVS while areas VIP and V6 were highly activated during antero-posterior GVS, but remained silent during lateral GVS. Using psychophysiological interaction (PPI) analyses, we confirmed that a cortical network including areas V6 and VIP is engaged during antero-posterior GVS. Our results suggest that V6 and VIP play a specific role in processing multisensory signals specific to locomotion during navigation.

We have investigated whether vestibular information plays a role in the control of voluntary movement of the upper body. Movement consisted of a lateral tilt of the upper body in the frontal plane through an angle of about 8 deg. The influence of vestibular input was assessed from the effect of long duration (3–6 s), low‐intensity (0.7 mA) galvanic vestibular stimulation (GVS) applied at different times relative to the movement. GVS always produced a tilt of the body in the frontal plane but the response was larger and more prolonged when the onset of stimulation coincided with the cue to start moving compared with when it was applied some seconds after movement onset (i.e. while the subject was stationary in a tilted posture). When the stimulus began 2 s before the voluntary movement the response consisted of two distinct components separated in time, one that was linked to the onset of GVS and another that was linked to onset of the voluntary movement. The large response observed when GVS onset coincided with the movement cue resembled the sum (after realignment in time) of these two separate components. We suggest that these two components of the response to GVS relate to two different uses of vestibular information for whole‐body control: first, to help maintain balance of the body, and second, to help guide and improve the accuracy of voluntary movements involving motion of the head in space.

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