Artificial vestibular feedback in conditions of modified body scheme

Subjects standing in darkness on the rigid support, kept a vertical posture which was destabilized by vibration of the Achilles tendons. To create a feedback on the vestibular input, transmastoidal bipolar galvanic stimulation was used. Changes of current in the feedback contour looked as linear function considering amplitude and velocity of the subject's head displacements in reference to the vertical. To change the body scheme we used some posture configurations: turning of the head in relation to the trunk; turning of the trunk with the head fixed; joint turning of the head and trunk. As a result of these configurations, the head could be turned approximately at right angle in relation to the feet. In addition turning of one foot at right angle in relation to the other foot was used. Artificial feedback reduces body fluctuations caused by vibration only in the vertical plane which passes through interaural axis of the head. The authors assume that directional changes of vestibulo-motor responses and results of application of artificial feedback during changes of orientation of the head in relation to the feet can be connected to change of ensembles of vestibular hair cells, which signals dominate in responses of vestibulo-spinal neurones.     

Vestibular effects on posture instability evoked by moving visual environment and footing

Subjects held the vertical posture standing up on hard footing, having small degree of the freedom in the frontal plane. The stability of the vertical posture has been assessed by the standard deviations (sigma) from average amplitudes of the fluctuations of the subject's head (in frontal and sagittal planes) from conditional zero. Sinusoidal rotations of optokinetic cylinder, sinusoidal rotations of the footing, and combinations of these rotations, under phase shifts between the optokinetic cylinder and the footing, caused increase of sigma. The amplitude and velocity signal of the head deviations was transformed into low galvanic current applied to the mastoids and used as the artifical vestibular biofeedback. It was possible to reduce the value of the sigma for lateral tilts (raised in comparison with their values during stance in the dark as a result of destabilizing influence), varying coefficients of the biofeedback. At the same time, appropriate fluctuations in sagittal plane were not systematic.     

Velocity of body lean evoked by leg muscle vibration potentiate the effects of vestibular stimulation on posture

To investigate the vestibular and somatosensory interaction in human postural control, a galvanic vestibular stimulation of cosine bell shape resulting in a small forward or backward body lean was paired with three vibrations of both soleus muscles. The induced body lean was registered by the position of the center of foot pressure (CoP). During a quiet stance with eyes closed the vibration of both soleus muscles with frequency (of) 40 Hz, 60 Hz and 80 Hz resulted in the body lean backward with velocities related to the vibration frequencies. The vestibular galvanic stimulation with the head turned to the right caused forward or backward modification of CoP backward response to the soleus muscles vibration and peaked at 1.5-2 s following the onset of the vibration. The effect of the paired stimulation was larger than the summation of the vestibular stimulation during the quiet stance and a leg muscle vibration alone. The enhancement of the galvanic stimulation was related to the velocity of body lean induced by the leg muscle vibration. The galvanic vestibular stimulation during a faster body movement had larger effects than during a slow body lean or the quiet stance. The results suggest that velocity of a body postural movement or incoming proprioceptive signal from postural muscles potentiate the effects of simultaneous vestibular stimulations on posture.     

Cross-Modal Calibration of Vestibular Afference for Human Balance

To determine how the vestibular sense controls balance, we used instantaneous head angular velocity to drive a galvanic vestibular stimulus so that afference would signal that head movement was faster or slower than actual. In effect, this changed vestibular afferent gain. This increased sway 4-fold when subjects (N = 8) stood without vision. However, after a 240 s conditioning period with stable balance achieved through reliable visual or somatosensory cues, sway returned to normal. An equivalent galvanic stimulus unrelated to sway (not driven by head motion) was equally destabilising but in this situation the conditioning period of stable balance did not reduce sway. Reflex muscle responses evoked by an independent, higher bandwidth vestibular stimulus were initially reduced in amplitude by the galvanic stimulus but returned to normal levels after the conditioning period, contrary to predictions that they would decrease after adaptation to increased sensory gain and increase after adaptation to decreased sensory gain. We conclude that an erroneous vestibular signal of head motion during standing has profound effects on balance control. If it is unrelated to current head motion, the CNS has no immediate mechanism of ignoring the vestibular signal to reduce its influence on destabilising balance. This result is inconsistent with sensory reweighting based on disturbances. The increase in sway with increased sensory gain is also inconsistent with a simple feedback model of vestibular reflex action. Thus, we propose that recalibration of a forward sensory model best explains the reinterpretation of an altered reafferent signal of head motion during stable balance.     

Adaptational Modifications of the Vestibular Postural Response in Humans under Conditions of Horizontal Visual Reversal

In healthy volunteers, we recorded stabilograms and studied postural responses evoked by galvanic stimulation of the labyrinth (binaurally applied 1-mA current, 4 sec) with the subjects' eyes open and closed and under conditions of reversed visual perception. Horizontal reversal of the visual space was provided by using spectacles with the Dove's prisms. In series consisting of 10 sequential tests with eyes open, we observed a gradual drop in the response amplitude, while there were practically no changes in the maximum velocity of the displacement. Postural responses with eyes closed were considerably greater than those with eyes open, but their amplitude and velocity demonstrated no changes with sequential tests. Under conditions of reversal of the visual perception, both the amplitude and maximum velocity of the postural responses decreased with successive testing. Under the above conditions, at the beginning of a test series responses to vestibular stimulation were greater than those with eyes closed, but in repeated tests they decreased and attained the same magnitude as in the tests with eyes closed. Therefore, the effect of short-term adaptation to visual reversal on the system controlling vertical posture resulted in simple rejection of the information coming via the visual input. In another experimental mode, we studied the adaptation effects at longer (3 h long) visual reversal. Postural responses to galvanic stimulation of the labyrinth (monaurally applied, 2-mA current, 4 sec) were tested with 1-h-long intervals; tests with visual reversal and with eyes closed were made in a random order with each other. A 3-h-long interval with the prismatic spectacles on did not modify the amplitude and velocity of the vestibular postural responses when the tests were made with the eyes closed. When the tests were performed with the eyes open, but in the inverting spectacles, postural responses significantly decreased (by about 50-60%) to the 2nd and 3rd h of the experiment. Such selective suppression of the vestibular input under conditions of visual reversal can be interpreted as a result of adaptational transformation of the visual-vestibular relation directed toward minimization of the visual-vestibular conflict.     

Effects of Near-Threshold Stimulation of the Vestibular Apparatus on Postural Responses Evoked by Displacements of the Visualized Surrounding

In healthy subjects in the relaxed upward stance and perceiving a virtual visual environment (VVE), we recorded postural reactions to isolated visual and vestibular stimulations or their combinations. Lateral displacements of the visualized virtual scene were used as visual stimuli. The vestibular apparatus was stimulated by application of near-threshold galvanic current pulses to the proc. mastoidei of the temporal bones. Isolated VVE shifts evoked mild, nonetheless clear, body tilts readily distinguished in separate trials; at the same time, postural effects of isolated vestibular stimulation could be detected only after averaging of several trials synchronized with respect to the beginning of stimulation. Under conditions of simultaneous combined presentation of visual and vestibular stimuli, the direction of the resulting postural responses always corresponded to the direction of responses induced by VVE shifts. The contribution of an afferent volley from the vestibular organ depended on the coincidence/mismatch of the direction of motor response evoked by such a volley with the direction of response to visual stimulation. When both types of stimulations evoked unidirectional body tilts, postural responses were facilitated, and the resulting effect was greater than that of simple summation of the reactions to isolated actions of the above stimuli. In the case where isolated galvanic stimulation evoked a response opposite with respect to that induced by visual stimulation, the combined action of these stimuli of different modalities evoked postural responses identical in their magnitude, direction, and shape to those evoked by isolated visual stimulation. The above findings allow us to conclude that the effects of visual afferent input on the vertical posture under conditions of our experiments clearly dominate. In general, these results confirm the statement that neuronal structures involved in integrative processing of different afferent volleys preferably select certain type of afferentation carrying more significant or more detailed information on displacements (including oscillations) of the body in space.     

Somatosensory influence on postural response to galvanic vestibular stimulation

We investigated how postural responses to galvanic vestibular stimulation were affected by standing on a translating support surface and by somatosensory loss due to diabetic neuropathy. We tested the hypothesis that an unstable surface and somatosensory loss can result in an increase of vestibulospinal sensitivity. Bipolar galvanic vestibular stimulation was applied to subjects who were standing on a force platform, either on a hard, stationary surface or during a backward platform translation (9 cm, 4.2 cm/s). The intensity of the galvanic stimulus was varied from 0.25 to 1 mA. The amplitude of the peak body CoP displacement in response to the galvanic stimulus was plotted as a function of stimulus intensity for each individual. A larger increase in CoP displacement to a given increase in galvanic current was interpreted as an increase of vestibulospinal sensitivity. Subjects with somatosensory loss in the feet due to diabetes showed higher vestibulospinal sensitivity than healthy subjects when tested on a stationary support surface. Control subjects and patients with somatosensory loss standing on translating surface also showed increased galvanic response gains compared to stance on a stationary surface. The severity of the somatosensory loss in the feet correlated with the increased postural sensitivity to galvanic vestibular stimulation. These results showed that postural responses to galvanic vestibular stimulus were modified by somatosensory information from the surface. Somatosensory loss due to diabetic neuropathy and alteration of somatosensory input during stance on translating support surface resulted in increased vestibulospinal sensitivity.     

Enhancing human balance control with galvanic vestibular stimulation

 With galvanic vestibular stimulation (GVS), electrical current is delivered transcutaneously to the vestibular afferents through electrodes placed over the mastoid bones. This serves to modulate the continuous firing levels of the vestibular afferents, and causes a standing subject to lean in different directions depending on the polarity of the current. Our objective in this study was to test the hypothesis that the sway response elicited by GVS can be used to reduce the postural sway resulting from a mechanical perturbation. Nine subjects were tested for their postural responses to both galvanic stimuli and support-surface translations. Transfer-function models were fit to these responses and used to calculate a galvanic stimulus that would act to counteract sway induced by a support-surface translation. The subjects' responses to support-surface translations, without and with the stabilizing galvanic stimulus, were then measured. With the stabilizing galvanic stimulus, all subjects showed significant reductions in both sway amplitude and sway latency. Thus, with GVS, subjects maintained a more erect stance and followed the support-surface displacement more closely. These findings suggest that GVS could possibly form the basis for a vestibular prosthesis by providing a means through which an individual's posture can be systematically controlled. Received: 11 May 2000 / Accepted in revised form: 20 November 2000     

Velocity of head movements and sensory-motor adaptation during and after short spaceflight

To investigate to time course of sensory-motor adaptation to microgravity, we tested spatially-directed voluntary head movements before, during and after short spaceflight. We also tested the re-adaptation of postural responses to sensory stimulation after space flight. The cosmonaut performed in microgravity six cycles of voluntary head rotation in pitch, roll and yaw directions. During the first days of weightlessness the angular velocity of head movements increased. Over the next days of microgravity the velocity of head movements gradually decreased. On landing day a significant decrease of head rotation velocity was observed compared to the head movement velocity before spaceflight. Re-adaptation to Earth condition measured by body sway on soft support showed similar time course, but re-adaptation measured by postural responses to vestibular galvanic stimulation was prolonged. These results showed that the angular velocity of aimed head movements of cosmonauts is a good indicator of sensory-motor adaptation in altered gravity conditions.     

Vestibular reafference shapes voluntary movement

The vestibular organs in the inner ear are commonly thought of as sensors that serve balance, gaze control, and higher spatial functions such as navigation. Here, we investigate their role in the online control of voluntary movements. The central nervous system uses sensory feedback information during movement to detect and correct errors as they develop. Vestibular organs signal three-dimensional head rotations and translations and so could provide error information for body movements that transport the head in space. To test this, we electrically stimulated human vestibular nerves during a goal-directed voluntary tilt of the trunk. The stimulating current waveform was made identical to the angular velocity profile of the head in the roll plane. With this, we could proportionally increase or decrease the rate of vestibular nerve firing, as if the head were rotating faster or slower than it actually was. In comparison to movements performed without stimulation, subjects tilted their trunk faster and further or slower and less far, depending upon the polarity of the stimulus. The response was negligible when identical stimulus waveforms were replayed to stationary subjects. We conclude that the brain uses vestibular information for online error correction of planned body-movement trajectories.     

Sensory processing in the vestibular nuclei during active head movements

Many secondary vestibular neurons are sensitive to head on trunk rotation during reflex-induced and voluntary head movements. During passive whole body rotation the interaction of head on trunk signals related to the vestibulo-collic reflex with vestibular signals increases the rotational gain of many secondary vestibular neurons, including many that project to the spinal cord. In some units, the sensitivity to head on trunk and vestibular input is matched and the resulting interaction produces an output that is related to the trunk velocity in space. In other units the head on trunk inputs are stronger and the resulting interaction produces an output that is larger during the reflex. During voluntary head movements, inputs related to head on trunk movement combine destructively with vestibular signals, and often cancel the sensory reafferent consequences of self-generated movements. Cancellation of sensory vestibular signals was observed in all of the antidromically identified secondary vestibulospinal units, even though many of these units were not significantly affected by reflexive head on trunk movements. The results imply that the inputs to vestibular neurons related to head on trunk rotation during reflexive and voluntary movements arise from different sources. We suggest that the relative strength of reflexive head on trunk input to different vestibular neurons might reflect the different functional roles they have in controlling the posture of the neck and body.     

A model of the nystagmus induced by off vertical axis rotation

A three-dimensional model is proposed that accounts for a number of phenomena attributed to the otoliths. It is constructed by extending and modifying a model of vestibular velocity storage. It is proposed that the otolith information about the orientation of the head to gravity changes the time constant of vestibular responses by modulating the gain of the velocity storage feedback loop. It is further proposed that the otolith signals, such as those that generate L-nystagmus (linear acceleration induced nystagmus), are partially coupled to the vestibular system via the velocity storage integrator. The combination of these two hypotheses suggests that a vestibular neural mechanism exists that performs correlation in the mathematical sense which is multiplication followed by integration. The multiplication is performed by the otolith modulation of the velocity storage feedback loop gain and the integration is performed by the velocity storage mechanism itself. Correlation allows calculation of the degree to which two signals are related and in this context provides a simple method of determining head angular velocity from the components of linear acceleration induced by off-vertical axis rotation. Correlation accounts for the otolith supplementation of the VOR and the sustained nystagmus generated by off-vertical axis rotation. The model also predicts the cross-coupling of horizontal and vertical optokinetic afternystagmus that occurs in head-lateral positions and the reported effects of tilt on vestibular responses.     

The Vestibular-Evoked Postural Response of Adolescents with Idiopathic Scoliosis Is Altered

Adolescent idiopathic scoliosis is a multifactorial disorder including neurological factors. A dysfunction of the sensorimotor networks processing vestibular information could be related to spine deformation. This study investigates whether feed-forward vestibulomotor control or sensory reweighting mechanisms are impaired in adolescent scoliosis patients. Vestibular evoked postural responses were obtained using galvanic vestibular stimulation while participants stood with their eyes closed and head facing forward. Lateral forces under each foot and lateral displacement of the upper body of adolescents with mild (n = 20) or severe (n = 16) spine deformation were compared to those of healthy control adolescents (n = 16). Adolescent idiopathic scoliosis patients demonstrated greater lateral displacement and net lateral forces than controls both during and immediately after vestibular stimulation. Altered sensory reweighting of vestibular and proprioceptive information changed balance control of AIS patients during and after vestibular stimulation. Therefore, scoliosis onset could be related to abnormal sensory reweighting, leading to altered sensorimotor processes.     

The Subjective Visual Vertical and the Subjective Haptic Vertical Access Different Gravity Estimates

The subjective visual vertical (SVV) and the subjective haptic vertical (SHV) both claim to probe the underlying perception of gravity. However, when the body is roll tilted these two measures evoke different patterns of errors with SVV generally becoming biased towards the body (A-effect, named for its discoverer, Hermann Rudolph Aubert) and SHV remaining accurate or becoming biased away from the body (E-effect, short for Entgegengesetzt-effect, meaning “opposite”, i.e., opposite to the A-effect). We compared the two methods in a series of five experiments and provide evidence that the two measures access two different but related estimates of gravitational vertical. Experiment 1 compared SVV and SHV across three levels of whole-body tilt and found that SVV showed an A-effect at larger tilts while SHV was accurate. Experiment 2 found that tilting either the head or the trunk independently produced an A-effect in SVV while SHV remained accurate when the head was tilted on an upright body but showed an A-effect when the body was tilted below an upright head. Experiment 3 repeated these head/body configurations in the presence of vestibular noise induced by using disruptive galvanic vestibular stimulation (dGVS). dGVS abolished both SVV and SHV A-effects while evoking a massive E-effect in the SHV head tilt condition. Experiments 4 and 5 show that SVV and SHV do not combine in an optimally statistical fashion, but when vibration is applied to the dorsal neck muscles, integration becomes optimal. Overall our results suggest that SVV and SHV access distinct underlying gravity percepts based primarily on head and body position information respectively, consistent with a model proposed by Clemens and colleagues.     

Dynamic Reweighting of Three Modalities for Sensor Fusion

We simultaneously perturbed visual, vestibular and proprioceptive modalities to understand how sensory feedback is re-weighted so that overall feedback remains suited to stabilizing upright stance. Ten healthy young subjects received an 80 Hz vibratory stimulus to their bilateral Achilles tendons (stimulus turns on-off at 0.28 Hz), a ±1 mA binaural monopolar galvanic vestibular stimulus at 0.36 Hz, and a visual stimulus at 0.2 Hz during standing. The visual stimulus was presented at different amplitudes (0.2, 0.8 deg rotation about ankle axis) to measure: the change in gain (weighting) to vision, an intramodal effect; and a change in gain to vibration and galvanic vestibular stimulation, both intermodal effects. The results showed a clear intramodal visual effect, indicating a de-emphasis on vision when the amplitude of visual stimulus increased. At the same time, an intermodal visual-proprioceptive reweighting effect was observed with the addition of vibration, which is thought to change proprioceptive inputs at the ankles, forcing the nervous system to rely more on vision and vestibular modalities. Similar intermodal effects for visual-vestibular reweighting were observed, suggesting that vestibular information is not a “fixed” reference, but is dynamically adjusted in the sensor fusion process. This is the first time, to our knowledge, that the interplay between the three primary modalities for postural control has been clearly delineated, illustrating a central process that fuses these modalities for accurate estimates of self-motion.     

Integrative responses of neurons in nucleus tractus solitarius to visceral afferent stimulation and vestibular stimulation in vertical planes

Anatomical studies have demonstrated that the vestibular nuclei project to nucleus tractus solitarius (NTS), but little is known about the effects of vestibular inputs on NTS neuronal activity. Furthermore, lesions of NTS abolish vomiting elicited by a variety of different triggering mechanisms, including vestibular stimulation, suggesting that emetic inputs may converge on the same NTS neurons. As such, an emetic stimulus that activates gastrointestinal (GI) receptors could alter the responses of NTS neurons to vestibular inputs. In the present study, we examined in decerebrate cats the responses of NTS neurons to rotations of the body in vertical planes before and after the intragastric administration of the emetic compound copper sulfate. The activity of more than one-third of NTS neurons was modulated by vertical vestibular stimulation, with most of the responsive cells having their firing rate altered by rotations in the head-up or head-down directions. These responses were aligned with head position in space, as opposed to the velocity of head movements. The activity of NTS neurons with baroreceptor, pulmonary, and GI inputs could be modulated by vertical plane rotations. However, injection of copper sulfate into the stomach did not alter the responses to vestibular stimulation of NTS neurons that received GI inputs, suggesting that the stimuli did not have additive effects. These findings show that the detection and processing of visceral inputs by NTS neurons can be altered in accordance with the direction of ongoing movements.     

Stabilometry and definition of the threshold of galvanic vestibular stimulation in normal subjects: an experimental study

The effect of the galvanic stimulation on the vestibular apparatus has been evaluated by registration on the postural deviations, using a stabilometry platform. We have studied the galvanic body-sway responses in a group of normal subjects, using a binauricolar bipolar stimulation, with the electrodes attached by means of surgical tape to the mastoid area. The records of body-sway responses have demonstrated in 80% of the considered cases a significant variation of all positional parameters after a current intensity of 2 mA, according the body sways toward the positive stimulus. At the same current intensity only five of the studied subjects have shown multidirectional swinging, in three cases joined with a subjective slight sway toward the ear stimulated with positive polarity. Therefore the galvanic test, joined with the posturography, proves to be a useful auxiliary method in vestibular investigation, allowing us to lower the threshold of galvanic stimulation and to make the electric stimulus better supported for the patient.     

Interaction between Visual and Proprioceptive Analyzers in Vertical Posture Control

The CNS can precisely assess the spatial position of the human body only by simultaneously processing and integrating the visual, proprioceptive, and vestibular inputs. Postural stability data make it possible to estimate changes taking place in the function of analyzers involved in the maintenance of the upright posture. The vertical posture stability was assessed in healthy children and children with spastic diplegic cerebral palsy from their postural responses to the presentation of single optokinetic or somatosensory stimuli. The visual analyzer was found to play a significant role in maintaining the upright position under natural gravity conditions in healthy children. A single exposure of the proprioceptive system to variable forces directed with the gravity of the earth (vertical) decreased the contribution of the visual analyzer. Healthy children maintained the upright position relying on the direction of movement of the optokinetic stimuli, which, however, produced no effect on the maintenance of posture in the patients. A hypothesis is proposed that prenatal or early postnatal CNS lesions decrease the contribution of phylogenetically newer brain structures to the regulation of upright posture.     

前庭电刺激联合前庭康复治疗周围性眩晕的疗效观察

目的：观察前庭电刺激联合前庭康复治疗周围性眩晕的疗效。方法：在常规药物治疗基础上将2008年5月．2012年5月我科眩晕门诊收治的226例诊断明确的单侧前庭周围性眩晕患者随机分成两组：前庭康复组和前庭康复＋前庭电刺激组。前庭康复组行常规前庭康复治疗,前庭康复＋前庭电刺激组在药物治疗及前庭康复基础上加用前庭电刺激,即在双侧乳突采取双极直流电刺激,每次15-20分钟,每天2次,共6周。治疗前及治疗后第2、4、6周行BBS评分及计时平衡试验时间测定以评判和比较两组的疗效。结果：两组患者治疗后第2、4、6周BBS评分及计时平衡试验时间较治疗前均明显增加（P〈0．05）,且B组各时点BBS评分及计时平衡试验时间均明显高于A组（P〈0．05）。结论：前庭电刺激联合前庭康复是较单纯前庭康复治疗前庭周围性眩晕更加有效的方法,其简单、无创、值得推广。     

Extracting phase-dependent human vestibular reflexes during locomotion using both time and frequency correlation approaches

Daily activities, such as walking, may require dynamic modulation of vestibular input onto motoneurons. This dynamic modulation is difficult to identify in humans due to limitations in the delivery and analysis of current vestibular probes, such as galvanic vestibular stimulation. Stochastic vestibular stimulation, however, provides an alternative method to extract human vestibular reflexes. Here, we used time-dependent coherence and time-dependent cross-correlation, coupled with stochastic vestibular stimulation, to investigate the phase dependency of human vestibular reflexes during locomotion. We found that phase-dependent activity from the medial gastrocnemius muscles is correlated with the vestibular signals over the 2- to 20-Hz bandwidth during the stance phase of locomotion. Vestibular-gastrocnemius coherence and time-dependent cross-correlations reached maximums at 21 ± 4 and 23 ± 8% of the step cycle following heel contact and before the period of maximal electromyographic activity (38 ± 5%). These results demonstrate 1) the effectiveness of these techniques in extracting the phase-dependent modulation of vestibulomuscular coupling during a cyclic task; 2) that vestibulomuscular coupling is phasically modulated during locomotion; and 3) that the period of strongest vestibulomuscular coupling does not correspond to the period of maximal electromyographic activity in the gastrocnemius. Therefore, we have shown that stochastic vestibular stimulation, coupled with time-frequency decomposition, provides an effective tool to assess the contribution of vestibular ex-afference to the muscular control during locomotion.