Reproduction of ON-center and OFF-center self-rotations

In self-rotation reproduction tasks, subjects appear to estimate the displacement angle and then reproduce this angle without necessarily replicating the entire temporal velocity profile. In contrast, subjects appear to reproduce the entire temporal velocity profile during linear motion stimulating the otoliths. To investigate what happens during combined rotation and translation, we investigated in darkness the central processing of vestibular cues during eccentric rotation. Controlling a centrifuge with a joystick, nine healthy subjects were asked to reproduce the angle of the previously imposed rotation. Rotations were either ON-center, or 50 cm OFF-center with inter-aural centripetal acceleration. Rotation duration was either variable (proportional to the traveled angle), or constant. We examined whether the stimulation of the otoliths during OFF-center rotation changes self-rotation reproduction, and whether rotation duration is processed differently by the nervous system with and without otolith stimulation. As postulated, the subjects indeed reproduced more closely the stimulus velocity profile when OFF-center. But the primary result is that the additional supra-threshold linear acceleration cues, measured by the otoliths, did not improve performance. More specifically, to our surprise, the ability to reproduce rotation angle degraded slightly in the presence of additional information from the otolith organs, with the linear acceleration cues appearing to interfere with the reproduction of movement duration.     

Space-time relativity in self-motion reproduction

Experiments on reproducing imposed self-motion showed that not only final distance or angle of motion, but also the temporal profile are reproduced. Reproduction errors have been attributed to sensory inputs, inaccurate memorization of the motion variable, or motor errors. However, another possible source of error has so far been neglected. The internal time base for path integration or movement memorization may be distorted and thus not reflect physical time. Because additional cognitive load was previously shown to affect subjective estimation of duration, we used a dual-task paradigm during either the stimulation or reproduction phase of three different movement reproduction tasks. We asked subjects 1) on a rotating chair to reproduce imposed passive whole body rotations by controlling the chair with a joystick, 2) on a treadmill to actively reproduce locomotion with respect to the treadmill, and 3) while blindfolded to reproduce a previously walked straight trajectory. The cognitive load changed the distance of reproduced self-motion by about 25% depending on whether the mental task was performed while experiencing or reproducing the motion. Although imposed velocity was reproduced accurately in all conditions, reproduced movement duration was affected in the same way as distance. This result implies that for the perception of distance traveled, perceptual space and time are closely interrelated. The findings are consistent with shared processing of temporal and spatial information. A computational model of motion reproduction including a discrete path integrator is proposed that is able to explain the experimental results within one coherent framework.     

The use of optical velocities for distance discrimination and reproduction during visually simulated self motion

Successful navigation through an environment requires precise monitoring of direction and distance traveled (”path integration” or ”dead reckoning”). Previous studies in blindfolded human subjects showed that velocity information arising from vestibular and somatosensory signals can be used to reproduce passive linear displacements. In these studies, visual information was excluded as sensory cue. Yet, in our everyday life, visual information is very important and usually dominates vestibular and somatosensory cues. In the present study, we investigated whether visual signals can be used to discriminate and reproduce simulated linear displacements. In a first set of experiments, subjects viewed two sequences of linear motion and were asked in a 2AFC task to judge whether the travel distance in the second sequence was larger or shorter than in the first. Displacements in either movement sequence could be forward (f) or backward (b). Subjects were very accurate in discriminating travel distances. Average error was less than 3% and did not depend on displacements being into the same (ff, bb) or opposite direction (fb, bf). In a second set of experiments, subjects had to reproduce a previously seen forward motion (passive condition), either in light or in darkness, i.e., with or without visual feedback. Passive displacements had different velocity profiles (constant, sinusoidal, complex) and speeds and were performed across a textured ground plane, a 2-D plane of dots or through a 3-D cloud of dots. With visual feedback, subjects reproduced distances accurately. Accuracy did not depend on the kind of velocity profile in the passive condition. Subjects tended to reproduce distance by replicating the velocity profile of the passive displacement. Finally, in the condition without visual feedback, subjects reproduced the shape of the velocity profile, but used much higher speeds, resulting in a substantial overshoot of travel distance. Our results show that visual, vestibular, and somatosensory signals are used for path integration, following a common strategy: the use of the velocity profile during self-motion. Received: 3 June 1998 / Accepted: 15 February 1999     

Self-motion perception during a sequence of whole-body rotations in darkness

The main aim of this study was to examine how postrotatory effects, induced by passive whole-body rotations in darkness, could alter the perception of motion and eye movements during a subsequent rotation. Perception of angle magnitude was assessed in a reproduction task: blindfolded subjects were first submitted to a passive rotation about the earth-vertical axis on a mobile robot. They were then asked to reproduce this angle by controlling the robot with a joystick. Stimulus rotations ranged from 80 degrees to 340 degrees. Subjects were given one of two delay instructions: after the stimulus, they either had to await the end of postrotatory sensations before starting reproduction (condition free delay, FD), or they had to start immediately after the end of the stimulus rotation (no delay, ND). The delay in FD was used as an incidental measure of the subjective duration of these sensations. Eye movements were recorded with an infrared measuring system (IRIS). Results showed that in both conditions subjects accurately reproduced rotation angles, though they did not reproduce the stimulus dynamics. Peak velocities reached in ND were higher than in FD. This difference suggests that postrotatory effects induced a bias in the perception of angular velocity in the ND condition.     

Multifactorial interactions involved in linear self-transport distance estimate: a place for time.

As the vestibular system is the only sensory organ whose primary function is self-motion detection, we examined the conditions under which the otoliths, which detect the linear acceleration of the head, could be used to estimate traveled distance. In order to isolate the contribution of the otoliths (with the somatosensory system) from contributions of the visual and motor systems subjects were transported in darkness. We initially hypothesized that self-transport with continuously varying linear velocity should facilitate distance computation by continuously stimulating the otoliths, and that active control of self-motion should also help subjects estimate the distance traveled. However, it was found that the distance covered during self-motion is actually better estimated when transport velocity is quasi-constant. Nevertheless, such estimates strongly depend upon velocity magnitude; subjects show an idiosyncratic preferred self-motion velocity for which distance measurements are most accurate. Furthermore, the active control of self-transport improves estimates of self-motion mainly because the subjects can then adopt a constant velocity, and more precisely their preferred one. It was finally found that subjects mentally count in order to assess their displacement length, and that time perception is indeed disturbed by varying self-motion velocity.     

Gaze stabilization during dynamic posturography in normal and vestibulopathic humans

Dynamic posturography by measurement of center of pressure (COP) is a widely employed technique for evaluating the vestibular system. However, the relationship of COP motion to vestibulo-ocular reflex (VOR) function and image stability on the retina has not been determined previously. To assess these relationships, we report gaze, head, and trunk stability during dynamic posturography in 11 normal volunteers, 7 subjects with unilateral vestibular lesions, and 3 subjects with bilateral vestibular lesions. Posturographic tasks consisted of standing still and standing on a platform that was sliding (0.2 Hz), tilting (0.1 Hz), or covered with a foam cushion 6 cm thick while tilting (0.1 Hz). Each perturbation was imposed in the anterior-posterior and repeated in the medial-lateral direction, in both light and darkness. Subjects viewed (or in darkness remembered) a target located 50, 100, or 500 cm distant. COP, angular eye position, and angular and linear orbit and trunk positions were measured using magnetic search coils and flux gate magnetometer sensors. With the target visible, the velocity of image motion on the retina was on average always less than 1°/s, well within the range consistent with high visual acuity. In darkness, gaze velocity increased for normal and vestibulopathic subjects. During tilt, vestibulopathic subjects had a significantly greater gaze velocity than controls. Gain of the angular VOR (eye velocity/head velocity) was significantly lower in darkness than in light and in vestibulopathic as compared to control subjects. Gain of the VOR was significantly correlated with gaze instability, but variation in VOR gain accounted for only 20–40% of the variance. In darkness, the velocity of the COP was significantly greater in vestibulopathic than control subjects for every condition tested. In light, this difference was small and often not significant. Although spectral analysis of the COP indicated frequencies above 1 Hz that were not observed in motion of the trunk and orbit, root mean square (RMS) velocities of the trunk and orbit in the horizontal plane were higher in darkness and in vestibulopathic subjects, mirroring COP findings. Only in vestibulopathic subjects tested in darkness was there a correlation between COP velocity and gaze velocity; COP velocity was otherwise uncorrelated with gaze. Gaze velocity was greater with near than with distant targets. Vertical VOR gain was higher with near targets. No other significant effects of target distance were found. Head movement strategy, VOR gain, and COP were all unaffected by target proximity. These data show that gaze velocity measurements during dynamic posturography in darkness are sensitive to vestibular loss. With a visible target, both COP and gaze stability of vestibulopathic subjects are difficult to distinguish from normal. During visual feedback, it is likely that image stabilization over the range of frequencies tested is achieved through better head stability and through visual tracking, allowing vestibulopathic subjects to maintain adequate visual acuity. Received: 25 November 1997 / Accepted: 24 April 1998     

Heading judgments during active and passive self-motion

Previous studies have generally considered heading perception to be a visual task. However, since judgments of heading direction are required only during self-motion, there are several other relevant senses which could provide supplementary and, in some cases, necessary information to make accurate and precise judgments of the direction of self-motion. We assessed the contributions of several of these senses using tasks chosen to reflect the reference system used by each sensory modality. Head-pointing and rod-pointing tasks were performed in which subjects aligned either the head or an unseen pointer with the direction of motion during whole body linear motion. Passive visual and vestibular stimulation was generated by accelerating subjects at sub- or supravestibular thresholds down a linear track. The motor-kinesthetic system was stimulated by having subjects actively walk along the track. A helmet-mounted optical system, fixed either on the cart used to provide passive visual or vestibular information or on the walker used in the active walking conditions, provided a stereoscopic display of an optical flow field. Subjects could be positioned at any orientation relative to the heading, and heading judgments were obtained using unimodal visual, vestibular, or walking cues, or combined visual-vestibular and visual-walking cues. Vision alone resulted in reasonably precise and accurate head-pointing judgments (0.3° constant errors, 2.9° variable errors), but not rod-pointing judgments (3.5° constant errors, 5.9° variable errors). Concordant visual-walking stimulation slightly decreased the variable errors and reduced constant pointing errors to close to zero, while head-pointing errors were unaffected. Concordant visual-vestibular stimulation did not facilitate either response. Stimulation of the vestibular system in the absence of vision produced imprecise rod-pointing responses, while variable and constant pointing errors in the active walking condition were comparable to those obtained in the visual condition. During active self-motion, subjects made large headpointing undershoots when visual information was not available. These results suggest that while vision provides sufficient information to identify the heading direction, it cannot, in isolation, be used to guide the motor response required to point toward or move in the direction of self-motion.     

Velocity storage contribution to vestibular self-motion perception in healthy human subjects

Self-motion perception after a sudden stop from a sustained rotation in darkness lasts approximately as long as reflexive eye movements. We hypothesized that, after an angular velocity step, self-motion perception and reflexive eye movements are driven by the same vestibular pathways. In 16 healthy subjects (25-71 years of age), perceived rotational velocity (PRV) and the vestibulo-ocular reflex (rVOR) after sudden decelerations (90°/s(2)) from constant-velocity (90°/s) earth-vertical axis rotations were simultaneously measured (PRV reported by hand-lever turning; rVOR recorded by search coils). Subjects were upright (yaw) or 90° left-ear-down (pitch). After both yaw and pitch decelerations, PRV rose rapidly and showed a plateau before decaying. In contrast, slow-phase eye velocity (SPV) decayed immediately after the initial increase. SPV and PRV were fitted with the sum of two exponentials: one time constant accounting for the semicircular canal (SCC) dynamics and one time constant accounting for a central process, known as velocity storage mechanism (VSM). Parameters were constrained by requiring equal SCC time constant and VSM time constant for SPV and PRV. The gains weighting the two exponential functions were free to change. SPV were accurately fitted (variance-accounted-for: 0.85 ± 0.10) and PRV (variance-accounted-for: 0.86 ± 0.07), showing that SPV and PRV curve differences can be explained by a greater relative weight of VSM in PRV compared with SPV (twofold for yaw, threefold for pitch). These results support our hypothesis that self-motion perception after angular velocity steps is be driven by the same central vestibular processes as reflexive eye movements and that no additional mechanisms are required to explain the perceptual dynamics.     

Storing upright turns: how visual and vestibular cues interact during the encoding and recalling process

Many previous studies have focused on how humans combine inputs provided by different modalities for the same physical property. However, it is not yet very clear how different senses providing information about our own movements combine in order to provide a motion percept. We designed an experiment to investigate how upright turns are stored, and particularly how vestibular and visual cues interact at the different stages of the memorization process (encoding/recalling). Subjects experienced passive yaw turns stimulated in the vestibular modality (whole-body rotations) and/or in the visual modality (limited lifetime star-field rotations), with the visual scene turning 1.5 times faster when combined (unnoticed conflict). Then they were asked to actively reproduce the rotation displacement in the opposite direction, with body cues only, visual cues only, or both cues with either the same or a different gain factor. First, we found that in none of the conditions did the reproduced motion dynamics follow that of the presentation phase (Gaussian angular velocity profiles). Second, the unimodal recalling of turns was largely uninfluenced by the other sensory cue that it could be combined with during the encoding. Therefore, turns in each modality, visual, and vestibular are stored independently. Third, when the intersensory gain was preserved, the bimodal reproduction was more precise (reduced variance) and lay between the two unimodal reproductions. This suggests that with both visual and vestibular cues available, these combine in order to improve the reproduction. Fourth, when the intersensory gain was modified, the bimodal reproduction resulted in a substantially larger change for the body than for the visual scene rotations, which indicates that vision prevails for this rotation displacement task when a matching problem is introduced.     

Retention of habituation of vestibulo-ocular reflex and sensation of rotation in humans

In humans, habituation of vestibulo-ocular reflex (VOR) by repeated caloric or rotational stimulation has been well documented. However, less attention has been directed to the effect of habituation on the sensation of self-rotation and little is known about the retention duration of vestibular habituation. To investigate these characteristics, subjects were exposed to ten sessions of angular velocity steps in yaw, with a chair rotating either alternatively in both CW and CCW directions (bidirectional protocol) or always in the same direction (unidirectional protocol), i.e., CW or CCW. The retention of habituation of VOR and sensation of rotation induced by both protocols was studied for a period up to 8 months following the end of the habituation protocols. There was a progressive decline in the VOR peak slow phase velocity and time constant throughout the sessions during both protocols. These parameters then followed an exponential recovery with a time constant of about 1 month. The duration of the sensation of rotation also habituated during repeated angular velocity steps, but it was shorter for both directions of stimulation, including after the unidirectional protocol. Sinusoidal VOR gain was not affected by vestibular habituation to velocity steps, but sinusoidal VOR phase showed an increase in phase lead at 0.02 and 0.04 Hz, which also returned to baseline values within about 1 month. We conclude that vestibular habituation is a long-lasting phenomenon. These results may be helpful for designing and scheduling the protocols for drug studies using crossover design, rehabilitation of balance disorder patients, and for the application of intermittent artificial gravity during space missions.     

Long-term deficits in otolith, canal and optokinetic ocular reflexes of pigmented rats after unilateral vestibular nerve section

 Static and dynamic otolith, horizontal vestibular and optokinetic ocular reflexes were investigated in pigmented rats 1–6 and more months after unilateral vestibular nerve (UVN) section. Evoked responses were compared with published data from control rats studied under identical conditions. Static lateral tilt of UVN rats in the light evoked a vertical deviation in static eye position that was as large as in controls. In darkness, the evoked responses in UVN rats 6 months after the lesion were consistently smaller than in controls. Linear horizontal acceleration in darkness evoked vertical and torsional response components in UVN rats that were parallel-shifted towards lower gains and larger phase lags. Off-vertical axis rotation on a platform provoked responses that differed markedly from those recorded in intact rats with respect to the bias velocity component. These results suggest a permanent deficiency in the static and dynamic otolith-ocular reflex performance of UVN rats. Ocular responses to horizontal table velocity steps in darkness exhibited a direction-specific asymmetry in UVN rats. Step responses evoked by acceleration towards the intact side were larger in gain and longer in duration than responses evoked by acceleration towards the operated side. When compared with control data, responses to either side were reduced in UVN rats and the velocity store mechanism was barely activated by velocity steps towards the operated side. Responses evoked by horizontal optokinetic stimulation with constant pattern velocities were below control values in either direction. Slow-phase eye velocity saturated at much lower values than in intact rats, particularly during pattern motion towards the intact side. The duration of the optokinetic afternystagmus was asymmetrically reduced with respect to control data. Practically identical reductions in duration were found for vestibulo-ocular responses in the opposite directions. Behaving animals exhibited no obvious impairment in their spontaneous locomotory or exploratory activities. However, each UVN rat was impaired, even 2 years after the lesion, in its postural reaction to being lifted by the tail in the air. This observation suggests the presence of a permanent deficit in static and dynamic otolith-spinal reflexes that may be substituted on the ground by proprioceptive inputs. Received: 26 February 1997 / Accepted: 2 July 1997     

Integration of visual and inertial cues in the perception of angular self-motion

The brain is able to determine angular self-motion from visual, vestibular, and kinesthetic information. There is compelling evidence that both humans and non-human primates integrate visual and inertial (i.e., vestibular and kinesthetic) information in a statistically optimal fashion when discriminating heading direction. In the present study, we investigated whether the brain also integrates information about angular self-motion in a similar manner. Eight participants performed a 2IFC task in which they discriminated yaw-rotations (2-s sinusoidal acceleration) on peak velocity. Just-noticeable differences (JNDs) were determined as a measure of precision in unimodal inertial-only and visual-only trials, as well as in bimodal visual–inertial trials. The visual stimulus was a moving stripe pattern, synchronized with the inertial motion. Peak velocity of comparison stimuli was varied relative to the standard stimulus. Individual analyses showed that data of three participants showed an increase in bimodal precision, consistent with the optimal integration model; while data from the other participants did not conform to maximum-likelihood integration schemes. We suggest that either the sensory cues were not perceived as congruent, that integration might be achieved with fixed weights, or that estimates of visual precision obtained from non-moving observers do not accurately reflect visual precision during self-motion.     

Self-motion perception training: thresholds improve in the light but not in the dark

We investigated perceptual learning in self-motion perception. Blindfolded participants were displaced leftward or rightward by means of a motion platform and asked to indicate the direction of motion. A total of eleven participants underwent 3,360 practice trials, distributed over twelve (Experiment 1) or 6 days (Experiment 2). We found no improvement in motion discrimination in both experiments. These results are surprising since perceptual learning has been demonstrated for visual, auditory, and somatosensory discrimination. Improvements in the same task were found when visual input was provided (Experiment 3). The multisensory nature of vestibular information is discussed as a possible explanation of the absence of perceptual learning in darkness.     

Evaluation of the vestibulo-ocular reflex using sinusoidal off-vertical axis rotation in patients with acoustic neurinoma

The vestibulo-ocular reflex (VOR) was studied to examine the utility of off-vertical axis rotation (OVAR) in the diagnosis of acoustic neurinoma. Subjects were sinusoidally rotated with eyes open in complete darkness at frequencies of 0.4 and 0.8 Hz with a maximum angular velocity of 60°/s at either earth-vertical axis rotation (EVAR) or OVAR. Thirteen patients with acoustic neurinomas were investigated. Results showed that VOR gain during OVAR at 0.8 Hz and in a 30° nose-up position in patients with internal auditory canal tumors was significantly less than the gain measured during EVAR. The VOR gain measured from all patients (including those with tumors extending to the cerebellopontine angle) was not significantly different when the patients were subjected to EVAR and OVAR. These observations were possibly due to superior vestibular nerve dysfunction. We concluded that certain stimulating parameters—patient's nose tilted up 30°; sinusoidal OVAR at 0.8 Hz and 60°/s maximum angular head velocity—were useful for evaluating vestibular function in patients suffering from an acoustic neurinoma located within the internal auditory canal.     

Vestibular perception of passive whole-body rotation about horizontal and vertical axes in humans: goal-directed vestibulo-ocular reflex and vestibular memory-contingent saccades

This study was aimed at complementing the existing knowledge about vestibular perception of self-motion in humans. Both goal-directed vestibulo-ocular reflex and vestibular memory-contingent saccade (VM-CS) tasks were used, respectively as concurrent and retrospective magnitude estimators for passive whole-body rotation. Rotations were applied about the earth-vertical and earth-horizontal axes to study the effect of the otolith signal in self-rotation evaluation, and both in yaw and pitch to examine the horizontal and vertical semi-circular canals. Two different magnitudes of constant angular acceleration (50°/s² and 100°/s²) were used. The main findings were (1) strong correlation between both oculomotor responses of both tasks, (2) greater accuracy with rotations about the earth-vertical than the earth: -horizontal axis, (3) greater accuracy for yaw than for pitch rotations, (4) greater accuracy for high acceleration than for low, and (5) no effect of the delay (2s or 12s) in the VMCS task. Adequacy of both tasks as subjective magnitude estimators of vestibular perception of self-motion is discussed.On leave from the Laboratoíre de Physiologie Neurosensorielle, CNRS, Paris, FrancePresent address: Laboratoire de Physiologie de la Perception et de l'Action, CNRS, Collége de France, 15, rue de l'Ecole de Médecine, F-75270 Paris Cedex 06, France     

Latency of adaptive vergence eye movements induced by vergence-vestibular interaction training in monkeys

Clear vision of objects that move in depth toward or away from an observer requires vergence eye movements. The vergence system must interact with the vestibular system to maintain the object images on the foveae of both eyes during head movement. Previous studies have shown that training with sinusoidal vergence-vestibular interaction improves the frequency response of vergence eye movements during pitch rotation: vergence eye velocity gains increase and phase-lags decrease. To further understand the changes in eye movement responses in this adaptation, we examined latencies of vergence eye movements before and after vergence-vestibular training. Two head-stabilized Japanese monkeys were rewarded for tracking a target spot moving in depth that required vergence eye movements of 10°/s. This target motion was synchronized with pitch rotation at 20°/s. Both target and chair moved in a trapezoidal waveform interspersed with random inter-trial intervals. Before training, pitch rotation in complete darkness without a target did not induce vergence eye movements. Mean latencies of convergence and divergence eye movements induced by vergence target motion alone were 182 and 169 ms, respectively. After training, mean latencies of convergence and divergence eye movements to a target synchronized with pitch rotation shortened to 65 and 53 ms, and vergence eye velocity gains (relative to vergence target velocity) at the normal latencies were 0.68 and 1.53, respectively. Pitch rotation alone without a target induced vergence eye movements with similar latencies after training. These results indicate that vestibular information can be used effectively to initiate vergence eye movements following vergence-vestibular training.     

Electrophysiological evidence for visual-vestibular interaction in man

The aim of the experiments reported here was to confirm electrophysiologically the results of psychophysical experiments, which demonstrated that thresholds for object-motion detection are significantly raised during both concurrent active or passive sinusoidal head oscillations and during visually induced self-motion perception (circularvection, CV). This intersensory inhibition could now be demonstrated electrophysiologically by recording visual motion evoked potentials both during concurrent sinusoidal head oscillations and during visually induced apparent self-motion of the objectively stationary subject. Recordings of visual contrast reversal evoked potentials failed to reveal such an interaction. Perceptual phenomena with multisensory stimulation are well described in the literature. Berthoz et al. demonstrated the dominant influence of the visual channel on vestibular thresholds such that the detection of a suprathreshold vestibular stimulation was clearly impaired by a simultaneously moving visual pattern inducing linearvection and vice versa. Comparable results are reported for circularvection. Evidence for inhibitory interaction between object-motion and simultaneous self-motion perception also exists. Electrophysiological data on intersensory interaction in humans have only been reported between electrical stimulation of a limb and its concurrent movement by means of scalp-recorded somatosensory-evoked potentials (SSEPs) (e.g. refs. 3, 5). Electrophysiological evidence for the interaction of visual object-motion and vestibular self-motion perception in humans has never been reported in the literature thus far, though Hood and Kayan demonstrated that retinal image motion makes a contribution to the vestibularly evoked bioelectric response.     

Stabilization of gaze during circular locomotion in darkness. II. Contribution of velocity storage to compensatory eye and head nystagmus in the running monkey.

1. Yaw eye in head (Eh) and head on body velocities (Hb) were measured in two monkeys that ran around the perimeter of a circular platform in darkness. The platform was stationary or could be counterrotated to reduce body velocity in space (Bs) while increasing gait velocity on the platform (Bp). The animals were also rotated while seated in a primate chair at eccentric locations to provide linear and angular accelerations similar to those experienced while running. 2. Both animals had head and eye nystagmus while running in darkness during which slow phase gaze velocity on the body (Gb) partially compensated for body velocity in space (Bs). The eyes, driven by the vestibuloocular reflex (VOR), supplied high-frequency characteristics, bringing Gb up to compensatory levels at the beginning and end of the slow phases. The head provided substantial gaze compensation during the slow phases, probably through the vestibulocollic reflex (VCR). Synchronous eye and head quick phases moved gaze in the direction of running. Head movements occurred consistently only when animals were running. This indicates that active body and limb motion may be essential for inducing the head-eye gaze synergy. 3. Gaze compensation was good when running in both directions in one animal and in one direction in the other animal. The animals had long VOR time constants in these directions. The VOR time constant was short to one side in one animal, and it had poor gaze compensation in this direction. Postlocomotory nystagmus was weaker after running in directions with a long VOR time constant than when the animals were passively rotated in darkness. We infer that velocity storage in the vestibular system had been activated to produce continuous Eh and Hb during running and to counteract postrotatory afterresponses. 4. Continuous compensatory gaze nystagmus was not produced by passive eccentric rotation with the head stabilized or free. This indicates that an aspect of active locomotion, most likely somatosensory feedback, was responsible for activating velocity storage. 5. Nystagmus was compared when an animal ran in darkness and in light. the beat frequency of eye and head nystagmus was lower, and the quick phases were larger in darkness. The duration of head and eye quick phases covaried. Eye quick phases were larger when animals ran in darkness than when they were passively rotated. The maximum velocity and duration of eye quick phases were the same in both conditions. 6. The platform was counterrotated under one monkey in darkness while it ran in the direction of its long vestibular time constant.(ABSTRACT TRUNCATED AT 400 WORDS)     

Posterior parietal rTMS disrupts human Path Integration during a vestibular navigation task

In contrast to vision, the neuro-anatomical substrates of vestibular perception are obscure. The vestibular apparati provide a head angular velocity signal allowing perception of self-motion velocity. Perceived change of angular position-in-space can also be obtained from the vestibular head velocity signal via a process called Path Integration (so-called since displacement is obtained by a mathematical temporal integration of the vestibular velocity signal). It is unknown however, if distinct cortical loci sub-serve vestibular perceptions of velocity versus displacement (i.e. Path Integration). Previous studies of human brain activity have not used head motion stimuli hence precluding localisation of vestibular cortical areas specialised for Path Integration distinct from velocity perception. We inferred vestibular cortical function by measuring the disrupting effect of repetitive transcranial magnetic stimulation on the performance of a displacement-dependent vestibular navigation task. Our data suggest that posterior parietal cortex is involved in encoding contralaterally directed vestibular-derived signals of perceived angular displacement and a similar effect was found for both hemispheres. We separately tested whether right posterior parietal cortex was involved in vestibular-sensed velocity perception but found no association. Overall, our data demonstrate that posterior parietal cortex is involved in human Path Integration but not velocity perception. We suggest that there are separate brain areas that process vestibular signals of head velocity versus those involved in Path Integration.     

Neural correlates of oddball detection in self-motion heading: A high-density event-related potential study of vestibular integration

The perception of self-motion is a product of the integration of information from both visual and non-visual cues, to which the vestibular system is a central contributor. It is well documented that vestibular dysfunction leads to impaired movement and balance, dizziness and falls, and yet our knowledge of the neuronal processing of vestibular signals remains relatively sparse. In this study, high-density electroencephalographic recordings were deployed to investigate the neural processes associated with vestibular detection of changes in heading. To this end, a self-motion oddball paradigm was designed. Participants were translated linearly 7.8 cm on a motion platform using a one second motion profile, at a 45° angle leftward or rightward of straight ahead. These headings were presented with a stimulus probability of 80–20 %. Participants responded when they detected the infrequent direction change via button-press. Event-related potentials (ERPs) were calculated in response to the standard (80 %) and target (20 %) movement directions. Statistical parametric mapping showed that ERPs to standard and target movements differed significantly from 490 to 950 ms post-stimulus. Topographic analysis showed that this difference had a typical P3 topography. Individual participant bootstrap analysis revealed that 93.3 % of participants exhibited a clear P3 component. These results indicate that a perceived change in vestibular heading can readily elicit a P3 response, wholly similar to that evoked by oddball stimuli presented in other sensory modalities. This vestibular-evoked P3 response may provide a readily and robustly detectable objective measure for the evaluation of vestibular integrity in various disease models.