Search

Vestibular nuclei and cerebellar function comprise vestibular neural networks that control vestibular-related responses. However, the vestibular-related responses to simultaneous stimulation of these regions are unclear. This study aimed to examine whether the combination of noisy galvanic vestibular stimulation (nGVS) and cerebellar transcranial direct current stimulation (ctDCS) using a complex transcranial electrical stimulation device alters vestibular-dominant standing stability and vestibulo-ocular reflex (VOR) function. The center of foot pressure (COP) sway and VOR of participants (28 healthy, young adults) were assessed under four conditions of transcranial electrical stimulation using nGVS and ctDCS. The COP was calculated with the participant standing on a soft-foam surface with eyes closed using a force plate to evaluate body sway. VOR measurements were collected via passive head movements and fixation on a target projected onto the front wall using a video head impulse test (vHIT). VOR gain was calculated in six directions using a semicircular canal structure based on the ratio of eye movement to head movement. The nGVS + ctDCS and nGVS + sham ctDCS conditions decreased COP sway compared to the sham nGVS + ctDCS and sham nGVS + sham ctDCS conditions. No significant differences were observed in the main effect of stimulation or the interaction of stimulation and direction on the vHIT parameters. The results of this study suggest that postural stability may be independently affected by nGVS. Our findings contribute to the basic neurological foundation for the clinical application of neurorehabilitation using transcranial electrical stimulation of the vestibular system., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Background: Vestibular stimulation causes postural unsteadiness accompanied by a sensation of tilt., Aims/objectives: The mechanism of the sensation of tilt needs to be assessed by accurate calculation of the rotational axis of torsional eye position under various vestibular stimulations., Material and Methods: Twenty-two healthy subjects participated in the study. Thirteen subjects underwent bilateral vestibular stimulation by on-axis yaw rotation under various head positions, and eighteen subjects underwent unilateral vestibular stimulation by caloric irrigation under various head positions. Listing's Plane was plotted for the eye movement data obtained by three-dimensional video-oculography., Results: The offset of Listing's Plane showed sustained deviation of torsional eye position that was more prominent in head positions that stimulated lateral semicircular canals more than vertical semicircular canals. There was a less prominent and directionally reversed offset in head positions that stimulated vertical canals more than lateral semicircular canals., Conclusion and Significance: The sustained torsional eye position was validated by accurate assessment using Listing's Plane. The mechanism behind the deviation may be due to a combination of multiple anatomical components within the vestibular apparatus, with potentially stronger influence from lateral semicircular canals.

Noninvasive electrical stimulation of the vestibular system in humans has become an increasingly popular tool with a broad range of research and clinical applications. However, common assumptions regarding the neural mechanisms that underlie the activation of central vestibular pathways through such stimulation, known as galvanic vestibular stimulation (GVS), have not been directly tested. Here, we show that GVS is encoded by VIIIth nerve vestibular afferents with nonlinear dynamics that differ markedly from those predicted by current models. GVS produced asymmetric activation of both semicircular canal and otolith afferents to the onset versus offset and cathode versus anode of applied current, that in turn produced asymmetric eye movement responses in three awake-behaving male monkeys. Additionally, using computational methods, we demonstrate that the experimentally observed nonlinear neural response dynamics lead to an unexpected directional bias in the net population response when the information from both vestibular nerves is centrally integrated. Together our findings reveal the neural basis by which GVS activates the vestibular system, establish that neural response dynamics differ markedly from current predictions, and advance our mechanistic understanding of how asymmetric activation of the peripheral vestibular system alters vestibular function. We suggest that such nonlinear encoding is a general feature of neural processing that will be common across different noninvasive electrical stimulation approaches. SIGNIFICANCE STATEMENT Here, we show that the application of noninvasive electrical currents to the vestibular system (GVS) induces more complex responses than commonly assumed. We recorded vestibular afferent activity in macaque monkeys exposed to GVS using a setup analogous to human studies. GVS evoked notable asymmetries in irregular afferent responses to cathodal versus anodal currents. We developed a nonlinear model explaining these GVS-evoked afferent responses. Our model predicts that GVS induces directional biases in centrally integrated head motion signals and establishes electrical stimuli that recreate physiologically plausible sensations of motion. Altogether, our findings provide new insights into how GVS activates the vestibular system, which will be vital to advancing new clinical and biomedical applications., (Copyright © 2023 the authors.)

Strong magnetic fields induce dizziness, vertigo, and nystagmus due to Lorentz forces acting on the cupula in the semi-circular canals, an effect called magnetic vestibular stimulation (MVS). In this article, we present an experimental setup in a 7T MRT scanner (MRI scanner) that allows the investigation of the influence of strong magnetic fields on nystagmus as well as perceptual and cognitive responses. The strength of MVS is manipulated by altering the head positions of the participants. The orientation of the participants' semicircular canals with respect to the static magnetic field is assessed by combining a 3D magnetometer and 3D constructive interference in steady-state (3D-CISS) images. This approach allows to account for intra- and inter-individual differences in participants' responses to MVS. In the future, MVS can be useful for clinical research, for example, in the investigation of compensatory processes in vestibular disorders. Furthermore, it could foster insights into the interplay between vestibular information and cognitive processes in terms of spatial cognition and the emergence of self-motion percepts under conflicting sensory information. In fMRI studies, MVS can elicit a possible confounding effect, especially in tasks influenced by vestibular information or in studies comparing vestibular patients with healthy controls.

Objectives: Although vestibular deficits can have severe repercussions on the early motor development in children, vestibular assessment in young children has not yet been routinely integrated in clinical practice and clear diagnostic criteria to detect early vestibular deficits are lacking. In young children, specific adjustments of the test protocol are needed, and normative data are age-dependent as the vestibular pathways mature through childhood. Therefore, this study aims to demonstrate the feasibility of an extensive age-dependent vestibular test battery, to provide pediatric normative data with the concurrent age trends, and to offer a clinical framework for pediatric vestibular testing., Design: This normative study included 133 healthy children below the age of 4 years (mean: 22 mo, standard deviation: 12.3 mo, range: 5-47 mo) without history of hearing loss or vestibular symptoms. Children were divided into four age categories: 38 children younger than 1 year old, 37 one-year olds, 33 two-year olds, and 25 three-year olds. Children younger than 3 years of age were examined with the video Head Impulse Test (vHIT) of the horizontal semicircular canals, cervical vestibular evoked myogenic potentials (cVEMP) with bone conduction stimuli, and the rotatory test at 0.16, 0.04, and 0.01 Hz. In 3-year old children, the vHIT of the vertical semicircular canals and ocular vestibular evoked myogenic potentials (oVEMP) using a minishaker were added to the protocol., Results: The horizontal vHIT appeared to be the most feasible test across age categories, except for children younger than 1-year old in which the success rate was the highest for the cVEMP. Success rates of the rotatory test varied the most across age categories. Age trends were found for the vHIT as the mean vestibulo-ocular reflex (VOR) gain increased significantly with age (r = 0.446, p < 0.001). Concerning the cVEMP, a significant increase with age was found for latency P1 (r = 0.420, p < 0.001), rectified interpeak amplitude P1-N1 (r = 0.574, p < 0.001), and averaged electromyographic (EMG) activity (r = 0.430, p < 0.001), whereas age trends for the latency N1 were less pronounced (r = 0.264, p = 0.004). Overall, the response parameters of the rotatory test did not show significant age effects ( p > 0.01), except for the phase at 0.01 Hz (r = 0.578, p < 0.001). Based on the reported success rates and age-dependent normative vestibular data, straightforward cutoff criteria were proposed (vHIT VOR gain < 0.7, cVEMP rectified interpeak amplitude < 1.3, oVEMP interpeak amplitude < 10 µV) with accompanying clinical recommendations to diagnose early vestibular impairment., Conclusions: In this large cohort of typically developing children below the age of 4 years, the vHIT and cVEMP were the most feasible vestibular tests. Moreover, the age-dependent normative vestibular data could specify age trends in this group of young children. Finally, based on the current results and clinical experience of more than ten years at the Ghent University Hospital (Belgium), a clinical framework to diagnose early vestibular deficits in young patients is proposed., Competing Interests: None of the authors have financial or non-financial conflicts of interest to disclose relevant to this paper., (Copyright © 2022 Wolters Kluwer Health, Inc. All rights reserved.)

Neurophysiological tests probing the vestibulo-ocular, colic and spinal pathways are the gold standard to evaluate the vestibular system in clinics. In contrast, vestibular perception is rarely tested despite its potential usefulness in professional training and for the longitudinal follow-up of professionals dealing with complex man-machine interfaces, such as aircraft pilots. This is explored here using a helicopter flight simulator to probe the vestibular perception of pilots. The vestibular perception of nine professional helicopter pilots was tested using a full flight helicopter simulator. The cabin was tilted six times in roll and six times in pitch (-15°, -10°, -5°, 5°, 10° and 15°) while the pilots had no visual cue. The velocities of the outbound displacement of the cabin were kept below the threshold of the semicircular canal perception. After the completion of each movement, the pilots were asked to put the cabin back in the horizontal plane (still without visual cues). The order of the 12 trials was randomized with two additional control trials where the cabin stayed in the horizontal plane but rotated in yaw (-10° and +10°). Pilots were significantly more precise in roll (average error in roll: 1.15 ± 0.67°) than in pitch (average error in pitch: 2.89 ± 1.06°) (Wilcoxon signed-rank test: p < 0.01). However, we did not find a significant difference either between left and right roll tilts (p = 0.51) or between forward and backward pitch tilts (p = 0.59). Furthermore, we found that the accuracies were significantly biased with respect to the initial tilt. The greater the initial tilt was, the less precise the pilots were, although maintaining the direction of the tilt, meaning that the error can be expressed as a vestibular error gain in the ability to perceive the modification in the orientation. This significant result was found in both roll (Friedman test: p < 0.01) and pitch (p < 0.001). However, the pitch trend error was more prominent (gain = 0.77 vs gain = 0.93) than roll. This study is a first step in the determination of the perceptive-motor profile of pilots, which could be of major use for their training and their longitudinal follow-up. A similar protocol may also be useful in clinics to monitor the aging process of the otolith system with a simplified testing device., (© 2023 The Authors. Physiological Reports published by Wiley Periodicals LLC on behalf of The Physiological Society and the American Physiological Society.)

Background: The video head impulse test (vHIT) is a common assessment of semicircular canal function during high-speed impulses. Reliability of the vHIT for assessing vertical semicircular canals is uncertain. Vertical head impulses require a complex head movement, making it difficult to isolate a single semicircular canal and interpret resulting eye rotations., Objective: The purpose of this study was to provide descriptive head kinematics and vestibular stimuli during vertical plane impulses to ultimately improve impulse delivery and interpretation of vHIT results for vertical semicircular canals., Methods: Six participants received right anterior (RA) and left posterior (LP) semicircular canal impulses. Linear displacements, rotational displacements, and rotational velocities of the head were measured. Peak velocities in semicircular canal planes and peak-to-peak gravitoinertial accelerations at the otolith organs were derived from head kinematics., Results: The largest rotational velocities occurred in the target semicircular canal plane, with non-negligible velocities occurring in non-target planes. Larger vertical displacements and accelerations occurred on the right side of the head compared to the left for RA and LP impulses., Conclusions: These results provide a foundation for designing protocols to optimize stimulation applied to a singular vertical semicircular canal and for interpreting results from the vHIT for vertical semicircular canals.

Background: During a simulated coordinated turn in a gondola centrifuge, experienced pilots show a substantial inter-individual variability in visual measures of perceived roll tilt. Because of the centrifuge's small radius, the pattern of stimuli to the semicircular canals during acceleration of the centrifuge differs in certain respects from that of an aircraft entering a turn., Objective: To explore whether these differences may be of significance for the pilot's roll- plane orientation and whether individual characteristics revealed in the centrifuge correspond to those during real flight., Method: 8 fixed-wing air-force pilots were tested in a centrifuge and a high-performance aircraft. The centrifuge was accelerated to 2 G (gondola inclination 60°) within 10 s. The duration at 2 G was 6 minutes. Similar profiles were created in the aircraft. The subjective visual horizontal (SVH) was measured using an adjustable luminous line in darkness. Each pilot was tested on three occasions: centrifuge (2 runs), aircraft (2 turns), centrifuge (2 runs). For each 2-G exposure, initial and final SVH values were established via curve fitting., Result: Despite a large inter-individual variability (±SD), group means were similar in the aircraft (initial: 43.0±20.6°; final: 22.5±14.8°) and centrifuge (initial: 40.6±17.0°; final: 20.5±16.0°). Further, individual peculiarities in response patterns were similar in the two conditions. For both the initial and final SVH tilt there was a high correlation between centrifuge and aircraft., Conclusion: The correspondence between conditions suggests that the centrifuge is an adequate means for demonstrating the fundamental motion pattern of coordinated flight and also for establishing the individual pilot's ability to perceive an aircraft's roll attitude.Findings are discussed in connection with vestibular learning and the possibility of underlying differences between pilots in the keenness for semicircular canal and somatosensory cues.

The vestibular system detects head motion to coordinate vital reflexes and provide our sense of balance and spatial orientation. A long-standing hypothesis has been that projections from the central vestibular system back to the vestibular sensory organs (i.e., the efferent vestibular system) mediate adaptive sensory coding during voluntary locomotion. However, direct proof for this idea has been lacking. Here we recorded from individual semicircular canal and otolith afferents during walking and running in monkeys. Using a combination of mathematical modeling and nonlinear analysis, we show that afferent encoding is actually identical across passive and active conditions, irrespective of context. Thus, taken together our results are instead consistent with the view that the vestibular periphery relays robust information to the brain during primate locomotion, suggesting that context-dependent modulation instead occurs centrally to ensure that coding is consistent with behavioral goals during locomotion., (© 2022. The Author(s).)

Background: Parkinson's disease (PD) is a common multi-system neurodegenerative disorder with possible vestibular system dysfunction, but prior vestibular function test findings are equivocal., Objective: To report and compare vestibulo-ocular reflex (VOR) gain as measured by the video head impulse test (vHIT) in participants with PD, including tremor dominant and postural instability/gait dysfunction phenotypes, with healthy controls (HC)., Methods: Forty participants with PD and 40 age- and gender-matched HC had their vestibular function assessed. Lateral and vertical semicircular canal VOR gains were measured with vHIT. VOR canal gains between PD participants and HC were compared with independent samples t-tests. Two distinct PD phenotypes were compared to HC using Tukey's ANOVA. The relationship of VOR gain with PD duration, phenotype, severity and age were investigated using logistic regression., Results: There were no significant differences between groups in vHIT VOR gain for lateral or vertical canals. There was no evidence of an effect of PD severity, phenotype or age on VOR gains in the PD group., Conclusion: The impulsive angular VOR pathways are not significantly affected by the pathophysiological changes associated with mild to moderate PD.

Direct and indirect pathways mediating the vestibulo-ocular reflexes, considering contributions made by the vestibular commissure, cerebellum, and vestibular efferent neurons are reviewed. This background leads to a detailed treatment of three-dimensional aspects of the vestibulo-ocular reflex, comparing the planes of the labyrinthine semicircular canals with the pulling directions of the extraocular muscles. By applying matrix algebra and rotational vectors, Robinson provides insights into the comparative anatomy of the vestibular system in different species, how central circuits process raw vestibular signals in three dimensions, and how the directions of eye movement caused by brainstem and cerebellar lesions can be explained., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The biophysical properties of the labyrinthine semicircular canals, and the electrophysiological properties of peripheral vestibular afferent neurons over a range of stimulus frequencies, are reviewed. Resting discharge activity and adaptive properties of vestibular neurons are discussed. Central processing of vestibular signals is then examined, including push-pull organization and the velocity storage mechanism. A detailed treatment of the final common neural integrator for oculomotor signals follows with consideration of its neural substrate and how distributed networks of neurons can overcome several problems posed by conventional control-systems models, such as why neural signals, but not background discharge, are integrated. Next, the behavior of the vestibulo-ocular reflex in darkness is compared with how it satisfies visual demands during natural activities. Finally, the reflex's performance at high frequencies of head rotation is discussed., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Electrical stimulation of the mammalian efferent vestibular system (EVS) predominantly excites primary vestibular afferents along two distinct time scales. Although roles for acetylcholine (ACh) have been demonstrated in other vertebrates, synaptic mechanisms underlying mammalian EVS actions are not well-characterized. To determine if activation of ACh receptors account for efferent-mediated afferent excitation in mammals, we recorded afferent activity from the superior vestibular nerve of anesthetized C57BL/6 mice while stimulating EVS neurons in the brainstem, before and after administration of cholinergic antagonists. Using a normalized coefficient of variation (CV*), we broadly classified vestibular afferents as regularly- (CV* < 0.1) or irregularly-discharging (CV* > 0.1) and characterized their responses to midline or ipsilateral EVS stimulation. Afferent responses to efferent stimulation were predominantly excitatory, grew in amplitude with increasing CV*, and consisted of fast and slow components that could be identified by differences in rise time and post-stimulus duration. Both efferent-mediated excitatory components were larger in irregular afferents with ipsilateral EVS stimulation. Our pharmacological data show, for the first time in mammals, that muscarinic AChR antagonists block efferent-mediated slow excitation whereas the nicotinic AChR antagonist DHβE selectively blocks efferent-mediated fast excitation, while leaving the efferent-mediated slow component intact. These data confirm that mammalian EVS actions are predominantly cholinergic.

Aims: Although the vestibular system has been widely investigated over the past 50 years, there is still an unsolved mystery. Some special vestibular afferent (SVA) neurons responding to both rotation and linear force were found through neurophysiological techniques, however, the sensory overlap mechanism of SVA neurons is still unclear, which may be closely related to vestibular-related diseases., Materials and Methods: To address the above-mentioned problem, a cupula buoyancy theory was established in the present study, where SVA neurons were considered semicircular canal afferent (SCCA) neurons. Then labyrinth anatomy and neural response dynamics of vestibular afferent neurons in chinchilla were investigated through vestibular labyrinth reconstruction and single unit recording technique, respectively., Key Findings: We analyzed the deflections of cupulae under multiple conditions with the help of Amira Software and predicted the neural response law of SCCA neurons to linear force based on the cupula buoyancy theory. Data analysis confirmed that the basic response characteristic of SVA neurons had no significant difference to those of SCCA neurons, but were significantly different from those of otolith afferent neurons. Further, the actual responses of SVA neurons to linear force are completely consistent with our predictions. These results strongly suggest that SVA neurons actually are SCCA neurons, and the cupula buoyancy theory is the key to the sensory overlap mechanism of SCCA neurons., Significance: Our study revealed the real identity of SVA neurons and provided a reasonable mechanism for sensory overlap of rotation and linear force, which improved our understanding about the vestibular system., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests, and all authors confirmed the accuracy of the data., (Copyright © 2020 Elsevier Inc. All rights reserved.)

The normal function of the vestibular system is crucial for the sense of balance. The techniques used to assess the vestibular function plays a vital role in the research of the vestibular system. In this article, we have systematically reviewed some popular methods employing vestibular reflexes and vestibular evoked potentials for assessing the vestibular function in rodent models. These vestibular reflexes and vestibular evoked potentials to effective stimuli have been used as nondestructive and objective functional measures. The main types of vestibular reflexes include the vestibulo-ocular reflex (VOR), vestibulocollic reflex (VCR), and vestibulo-sympathetic reflex (VSR). They are all capable of indicating the functions of the semicircular canals and otoliths. However, the VOR assessment is much more prevalently used because of the relatively stereotypical inputoutput relationship and simple motion pattern of the ocular response. In contrast, the complicated motion pattern and small gain of the VCR response, as well as the undesired component possibly contributed from the acceleration receptors outside the labyrinths in the VSR response, restrict the widespread applications of VCR and VSR in the assessment of the vestibular system. The vestibular evoked myogenic potentials (VEMPs) and vestibular sensory evoked potentials (VsEPs) are the two typical evoked potentials that have been also employed for evaluating the vestibular function. Through exploiting different types of the VEMPs, the saccular and utricular functions can be evaluated separately. The sound-induced VEMPs, moreover, are capable of noninvasively assessing the unilateral vestibular function. The VsEPs, via the morphology of their signal waveforms, enable the access to the location-specific information that indicates the functional statuses of different components within the vestibular neural pathway., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Background: Only a few studies in the literature demonstrate the effect of vestibular rehabilitation (VR) on all vestibular receptor organs. Furthermore, very little evidence of the effect of VR on isolated otolith dysfunction (IOD) is available., Objective: The study aimed to investigate the effect of VR on all vestibular receptor organs in patients with different types of unilateral vestibular hypofunction (UVH)., Methods: We enrolled 80 patients with three different types of UVH; combined and isolated loss of semicircular canal and otolith organ function. All patients performed a 12-week customized program of VR and received a full battery of vestibular function tests, before and after the VR. The DHI and SF-36 were performed before, after 6 weeks, and 12 weeks of the VR., Results: Parameters of the caloric test, video head impulse test, ocular and cervical vestibular evoked myogenic potentials were significantly improved after VR. A total of 59 (74%) patients fully recovered, with no significant difference in recovery regarding the type (p = 0.13) and stage of UVH (p = 0.13). All patients reported significantly lower disability and a better quality of life after the VR based on the DHI and SF-36 score., Conclusions: Vestibular rehabilitation has a positive effect on the recovery of all vestibular receptor organs and it should be used in patients with IOD.

Background: The peripheral vestibular end organ is considered to consist of semi-circular canals (SCC) for detection of angular accelerations and the otoliths for detection of linear accelerations. However, otoliths being phylogenetically the oldest part of the vestibular sensory organs are involved in detection of all motions., Summary: This study elaborates on this property of the otolith organ, as this concept can be of importance for the currently designed vestibular implant devices. Key Message: The analysis of the evolution of the inner ear and examination of clinical examples shows the robustness of the otolith system and inhibition capacity of the SCC. The otolith system must be considered superior to the SCC system as illustrated by evolution, clinical evidence, and physical principles., (© 2020 The Author(s) Published by S. Karger AG, Basel.)

Semicircular canal afferent neurons transmit information about head rotation to the brain. Mathematical models of how they do this have coevolved with concepts of how brains perceive the world. A 19th-century "camera" metaphor, in which sensory neurons project an image of the world captured by sense organs into the brain, gave way to a 20th-century view of sensory nerves as communication channels providing inputs to dynamical control systems. Now, in the 21st century, brains are being modeled as Bayesian observers who infer what is happening in the world given noisy, incomplete, and distorted sense data. The semicircular canals of the vestibular apparatus provide an experimentally accessible, low-dimensional system for developing and testing dynamical Bayesian generative models of sense data. In this review, we summarize advances in mathematical modeling of information transmission by semicircular canal afferent sensory neurons since the first such model was proposed nearly a century ago. Models of information transmission by vestibular afferent neurons may provide a foundation for developing realistic models of how brains perceive the world by inferring the causes of sense data.

Key Points: Considerable debate exists regarding whether electrical vestibular stimuli encoded by vestibular afferents induce a net signal of linear acceleration, rotation or a combination of the two. This debate exists because an isolated signal of head rotation encoded by the vestibular afferents can cause perceptions of both linear and angular motion. We recorded participants' perceptions in different orientations relative to gravity and predicted their responses by modelling the effect of electrical vestibular stimuli on vestibular afferents and a current model of central vestibular processing. We show that, even if electrical vestibular stimuli are encoded as a net signal of head rotation, participants perceive both linear acceleration and rotation motions, provided the electrical stimulation-induced rotational vector has a component orthogonal to gravity. The emergence of a perception of linear acceleration from a single rotational input signal clarifies the origins of the neural mechanisms underlying electrical vestibular stimulation., Abstract: Electrical vestibular stimulation (EVS) is an increasingly popular biomedical tool for generating sensations of virtual motion in humans, for which the mechanism of action is a topic of considerable debate. Contention surrounds whether the evoked vestibular afferent activity encodes a signal of net rotation and/or linear acceleration. Central processing of vestibular self-motion signals occurs through an internal representation of gravity that can lead to inferred linear accelerations in absence of a true inertial acceleration. Applying this model to virtual signals of rotation evoked by EVS, we predict that EVS will induce behaviours attributed to both angular and linear motion, depending on the head orientation relative to gravity. To demonstrate this, 18 subjects indicated their perceived motion during sinusoidal EVS when in one of four head/body positions orienting the gravitational vector parallel or orthogonal to the EVS rotation vector. During stimulation, participants selected one simulated movement from seven that corresponded best to what they perceived. Participants' responses in each orientation were predicted by a model combining the influence of EVS on vestibular afferents with known mechanisms of vestibular processing. When the EVS rotation vector had a component orthogonal to gravity, human perceptual responses were consistent with a non-zero central estimate of interaural or superior-inferior linear acceleration. The emergence of a perception of linear acceleration from a single rotational input signal clarifies the origins of the neural mechanisms underlying EVS, which has important implications for its use in human biomedical or sensory augmentation applications., (© 2019 The Authors. The Journal of Physiology © 2019 The Physiological Society.)

Purkinje neurons in the caudal cerebellar vermis combine semicircular canal and otolith signals to segregate linear and gravitational acceleration, evidence for how the cerebellum creates internal models of body motion. However, it is not known which cerebellar circuit connections are necessary to perform this computation. We first showed that this computation is evolutionarily conserved and represented across multiple lobules of the rodent vermis. Then we tested whether Purkinje neuron GABAergic output is required for accurately differentiating linear and gravitational movements through a conditional genetic silencing approach. By using extracellular recordings from lobules VI through X in awake mice, we show that silencing Purkinje neuron output significantly alters their baseline simple spike variability. Moreover, the cerebellum of genetically manipulated mice continues to distinguish linear from gravitational acceleration, suggesting that the underlying computations remain intact. However, response gain is significantly increased in the mutant mice over littermate controls. Altogether, these data argue that Purkinje neuron feedback regulates gain control within the cerebellar circuit., Competing Interests: Conflict of interest statement: The authors declare no conflict of interest.

Introduction: Orientation in a 3-dimensional environment increases demands on the vestibular system. The anterior, lateral, and posterior semicircular canals sense pitch, roll, and yaw acceleration, respectively. The macular end organs sense linear acceleration and provide gravity reference. Creatures that evolved to fly (birds) share the same sensory systems and probably the same limitations as creatures that did not evolve to fly. However, bird semicircular canals have been noted for their large size relative to body mass since 1907., Methods: A brief literature review was conducted regarding the morphological structure of the vestibular systems in birds, and their evolutionary and functional significance as compared to that in humans., Results: For any given body mass, qualitatively more "agile" species have semicircular canals with larger radius of curvature. Specifically, the anterior and lateral canals were found to be preferentially larger than those of nonavian theropods and humans. Avian canals rarely approach circularity and all three canals typically undergo torsional excursions out of their respective planes., Discussion: Increase in the length of the semicircular canal is associated with greater sensitivity to rotational stimuli. Highly maneuverable birds of prey have the largest overall canal dimensions relative to body mass, most plausibly to match the rapid body rotations that characterize variable speed maneuvering during pursuit. By comparison, the semicircular canals in humans are relatively smaller and lack the adaptive morphological asymmetry in agile flying species. This teleological symmetry may present another inherent but less understood vestibular limitation during aerobatic maneuvers, which could lead to spatial disorientation.Cheung B, Ercoline W. Semicircular canal size and shape influence on disorientation. Aerosp Med Hum Perform. 2018; 89(8):744-748.

Objective: We aimed to quantify peripheral-vestibular deficits that may contribute to imbalanced stance/gait in patients with inflammatory neuropathies., Methods: Twenty-one patients (58 ± 15 y [mean age ± 1SD]; chronic-inflammatory-demyelinating-polyneuropathy = 10, Guillain-Barré Syndrome = 5, Anti-MAG peripheral neuropathy = 2, multifocal-motor-neuropathy = 4) were compared with 26 healthy controls. All subjects received video-head-impulse testing (vHIT), caloric irrigation and cervical/ocular vestibular-evoked myogenic-potentials (VEMPs). The Yardley vertigo-symptom-scale (VSS) was used to rate vertigo/dizziness. Postural stability was assessed using the functional gait-assessment (FGA). Pure-tone audiograms (n = 18), otoacoustic emissions (n = 12) and auditory brainstem responses were obtained (n = 12)., Results: Semicircular-canal hypofunction was noted in 9/21 (43%) patients (vHIT = 6; caloric irrigation = 5), whereas otolith function was impaired in 12/21 (57%) (oVEMPs = 8; cVEMPs = 5), resulting in vestibular impairment of at least one sensor in 13/21 (62%). On average, 2.4 ± 1.1 vestibular end organs (each side: anterior/posterior/horizontal canal, utriculus, sacculus; total = 10) were affected. The VSS-scores were higher in patients (16.8 ± 8.6 vs. 9.5 ± 6.2, p = 0.002) but did not correlate with the number of affected organs. Auditory neuropathy was found in 1/12 (8%) patients., Conclusion: Impairment of one or more vestibular end organs was frequent, but usually mild, possibly contributing to imbalance of stance/gait in inflammatory neuropathies., Significance: While our data does not support routine vestibular testing in inflammatory neuropathies, this may be considered in selected cases., (Copyright © 2017 International Federation of Clinical Neurophysiology. Published by Elsevier B.V. All rights reserved.)

Objectives: To measure the auditory (pure tone audiometry and word recognition scores) and vestibular (video head impulse test and vestibular myogenic potentials) outcomes in patients diagnosed with Meniere's disease (MD) who underwent cochlear implantation., Material and Methods: This prospective study included 23 cochlear implant users with MD and 29 patients diagnosed with far-advanced otosclerosis (the control group)., Results: The preoperative mean pure tone average thresholds were 99 and 122.5 dB for the Meniere's and control groups, respectively. Word recognition scores after cochlear implant yielded a median of 80% and 72% for the Meniere's and control groups, respectively. Semicircular canal gain was not observed to vary post implantation (mean variation for lateral, posterior, and anterior plane was 0, 0.03, and 0, respectively). The mean ocular and cervical myogenic potentials asymmetry varied as 9.65% and 18.39%, respectively., Conclusions: The auditory performance improved in patients with MD similar to the general cochlear implant population. No major dysfunction of otolithic or semicircular canal function was demonstrated after the implantation surgery.

Background: When viewing a far target, the gain of the horizontal vestibulo-ocular reflex (VOR) is around 1.0, but when viewing a near target there is an increased response. It has been shown that while this convergence-mediated modulation is unaffected by canal plugging and clinically practical transmastoid galvanic stimulation, it is eliminated by a partial peripheral gentamicin lesion., Objective: The aim of this study was to determine if convergence increases the gain during peripheral hyposensitivity elicited by cold thermal irrigation., Methods: The high frequency VOR gain was measured using video head impulse testing immediately after the cold caloric stimulus in 9 healthy human subjects with the lateral semicircular canals oriented approximately earth-vertical., Results: Before caloric irrigation, near viewing (15 cm) increased the average VOR gain by 28% (from 1 to 1.28). Cold (24°C) water irrigation of the right ear decreased the gain to 0.66 (far viewing) and 0.82 (near viewing) (22% difference). Although vergence also increased the gain for impulses to the left to the same degree before caloric stimulus, the caloric irrigation itself (applied to the right ear) did not influence the gain for contralateral impulses., Conclusion: In our experiments vergence increased the gain of the human angular VOR during peripheral hyposensitivity elicited by cold thermal irrigation. These results suggest that cold irrigation does not abolish the function of the nonlinear/phasic vestibular afferent pathway.

Objectives: During head rotations, neuronal firing rates increase in ipsilateral and decrease in contralateral vestibular afferents. At low accelerations, this "push-pull mechanism" is linear. At high accelerations, however, the change of firing rates is nonlinear in that the ipsilateral increase of firing rate is larger than the contralateral decrease. This mechanism of stronger ipsilateral excitation than contralateral inhibition during high-acceleration head rotation, known as Ewald's second law, is implemented within the nonlinear pathways. The authors asked whether caloric stimulation could provide an acceleration signal high enough to influence the contribution of the nonlinear pathway to the rotational vestibulo-ocular reflex gain (rVOR gain) during head impulses., Design: Caloric warm (44°C) and cold (24, 27, and 30°C) water irrigations of the left ear were performed in 7 healthy human subjects with the lateral semicircular canals oriented approximately earth-vertical (head inclined 30° from supine) and earth-horizontal (head inclined 30° from upright)., Results: With the lateral semicircular canal oriented earth-vertical, the strongest cold caloric stimulus (24°C) significantly decreased the rVOR gain during ipsilateral head impulses, while all other irrigations, irrespective of head position, had no significant effect on rVOR gains during head impulses to either side., Conclusions: Strong caloric irrigation, which can only be achieved with cold water, reduces the rVOR gain during ipsilateral head impulses and thus demonstrates Ewald's second law in healthy subjects. This unilateral gain reduction suggests that cold-water caloric irritation shifts the set point of the nonlinear relation between head acceleration and the vestibular firing rate toward a less acceleration-sensitive zone.

Objective: To explore ocular vestibular evoked myogenic potentials (oVEMP) to low-frequency vertex vibration (125 Hz) as a diagnostic test for superior canal dehiscence (SCD) syndrome., Methods: The oVEMP using 125 Hz single cycle bone-conducted vertex vibration were tested in 15 patients with unilateral superior canal dehiscence (SCD) syndrome, 15 healthy controls and in 20 patients with unilateral vestibular loss due to vestibular neuritis. Amplitude, amplitude asymmetry ratio, latency and interaural latency difference were parameters of interest., Results: The oVEMP amplitude was significantly larger in SCD patients when affected sides (53 μVolts) were compared to non-affected (17.2 μVolts) or compared to healthy controls (13.6 μVolts). Amplitude larger than 33.8 μVolts separates effectively the SCD ears from the healthy ones with sensitivity of 87% and specificity of 93%. The other three parameters showed an overlap between affected SCD ears and non-affected as well as between SCD ears and those in the two control groups., Conclusions: oVEMP amplitude distinguishes SCD ears from healthy ones using low-frequency vibration stimuli at vertex., Significance: Amplitude analysis of oVEMP evoked by low-frequency vertex bone vibration stimulation is an additional indicator of SCD syndrome and might serve for diagnosing SCD patients with coexistent conductive middle ear problems., (Copyright © 2016 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.)

Key Points: Lying supine in a strong magnetic field, such as in magnetic resonance imaging scanners, can induce a perception of whole-body rotation. The leading hypothesis to explain this invokes a Lorentz force mechanism acting on vestibular endolymph that acts to stimulate semicircular canals. The hypothesis predicts that the perception of whole-body rotation will depend on head orientation in the field. Results showed that the direction and magnitude of apparent whole-body rotation while stationary in a 7 T magnetic field is influenced by head orientation. The data are compatible with the Lorentz force hypothesis of magnetic vestibular stimulation and furthermore demonstrate the operation of a spatial transformation process from head-referenced vestibular signals to Earth-referenced body motion., Abstract: High strength static magnetic fields are known to induce vertigo, believed to be via stimulation of the vestibular system. The leading hypothesis (Lorentz forces) predicts that the induced vertigo should depend on the orientation of the magnetic field relative to the head. In this study we examined the effect of static head pitch (-80 to +40 deg; 12 participants) and roll (-40 to +40 deg; 11 participants) on qualitative and quantitative aspects of vertigo experienced in the dark by healthy humans when exposed to the static uniform magnetic field inside a 7 T MRI scanner. Three participants were additionally examined at 180 deg pitch and roll orientations. The effect of roll orientation on horizontal and vertical nystagmus was also measured and was found to affect only the vertical component. Vertigo was most discomforting when head pitch was around 60 deg extension and was mildest when it was around 20 deg flexion. Quantitative analysis of vertigo focused on the induced perception of horizontal-plane rotation reported online with the aid of hand-held switches. Head orientation had effects on both the magnitude and the direction of this perceived rotation. The data suggest sinusoidal relationships between head orientation and perception with spatial periods of 180 deg for pitch and 360 deg for roll, which we explain is consistent with the Lorentz force hypothesis. The effects of head pitch on vertigo and previously reported nystagmus are consistent with both effects being driven by a common vestibular signal. To explain all the observed effects, this common signal requires contributions from multiple semicircular canals., (© 2015 The Authors. The Journal of Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

This study examined the external morphology and morphometry of the human embryonic inner ear membranous labyrinth and documented its three-dimensional position in the developing embryo using phase-contrast X-ray computed tomography and magnetic resonance imaging. A total of 27 samples between Carnegie stage (CS) 17 and the postembryonic phase during trimester 1 (approximately 6-10 weeks after fertilization) were included. The otic vesicle elongated along the dorso-ventral axis and differentiated into the end lymphatic appendage and cochlear duct (CD) at CS 17. The spiral course of the CD began at CS18, with anterior and posterior semicircular ducts (SDs) forming prominent circles with a common crus. The spiral course of the CD comprised more than two turns at the postembryonic phase, at which time the height of the CD was evident. A linear increase was observed in the length of anterior, posterior, and lateral SDs, in that order, and the length of the CD increased exponentially over the course of development. Bending in the medial direction was observed between the cochlear and vestibular parts from the latero-caudal view, with the angle decreasing during development. The position of the inner ear was stable throughout the period of observation on the lateral to ventral side of the rhombencephalon, caudal to the pontine flexure, and adjacent to the auditory ganglia. The plane of the lateral semicircular canal was approximately 8.0°-14.6° with respect to the cranial caudal (z-)axis, indicating that the orientation of the inner ear changes during growth to adulthood., (© 2015 Wiley Periodicals, Inc.)

The vestibular system operates in a push-pull fashion using signals from both labyrinths and an intricate bilateral organization. Unilateral vestibular lesions cause well-characterized motor deficits that are partially compensated over time and whose neural correlates have been traced in the mean response modulation of vestibular nuclei cells. Here we compare both response gains and neural detection thresholds of vestibular nuclei and semicircular canal afferent neurons in intact vs. unilateral-lesioned macaques using three-dimensional rotation and translation stimuli. We found increased stimulus-driven spike count variability and detection thresholds in semicircular canal afferents, although mean responses were unchanged, after contralateral labyrinth lesion. Analysis of trial-by-trial spike count correlations of a limited number of simultaneously recorded pairs of canal afferents suggests increased noise correlations after lesion. In addition, we also found persistent, chronic deficits in rotation detection thresholds of vestibular nuclei neurons, which were larger in the ipsilesional than the contralesional brain stem. These deficits, which persisted several months after lesion, were due to lower rotational response gains, whereas spike count variability was similar in intact and lesioned animals. In contrast to persistent deficits in rotation threshold, translation detection thresholds were not different from those in intact animals. These findings suggest that, after compensation, a single labyrinth is sufficient to recover motion sensitivity and normal thresholds for the otolith, but not the semicircular canal, system., (Copyright © 2014 the American Physiological Society.)

This paper presents an alternative approach for angular-rate sensing based on the way that the natural vestibular semicircular canals operate, whereby the inertial mass of a fluid is used to deform a sensing structure upon rotation. The presented gyro has been fabricated in a commercially available MEMS process, which allows for microfluidic channels to be implemented in etched glass layers, which sandwich a bulk-micromachined silicon substrate, containing the sensing structures. Measured results obtained from a proof-of-concept device indicate an angular rate sensitivity of less than 1 °/s, which is similar to that of the natural vestibular system. By avoiding the use of a continually-excited vibrating mass, as is practiced in today's state-of-the-art gyroscopes, an ultra-low power consumption of 300 μW is obtained, thus making it suitable for implantation.

The semicircular canals of the inner ear detect head rotations and trigger compensatory movements that stabilize gaze and help maintain visual fixation. Mammals with large eyes and high visual acuity require precise gaze stabilization mechanisms because they experience diminished visual functionality at low thresholds of uncompensated motion. Because semicircular canal radius of curvature is a primary determinant of canal sensitivity, species with large canal radii are expected to be capable of more precise gaze stabilization than species with small canal radii. Here, we examine the relationship between mean semicircular canal radius of curvature, eye size, and visual acuity in a large sample of mammals. Our results demonstrate that eye size and visual acuity both explain a significant proportion of the variance in mean canal radius of curvature after statistically controlling for the effects of body mass and phylogeny. These findings suggest that variation in mean semicircular canal radius of curvature among mammals is partly the result of selection for improved gaze stabilization in species with large eyes and acute vision. Our results also provide a possible functional explanation for the small semicircular canal radii of fossorial mammals and plesiadapiforms., (Copyright © 2014 Wiley Periodicals, Inc.)

Sound-evoked vestibular myogenic potentials recorded from the sternocleidomastoid muscles (the cervical vestibular-evoked myogenic potential or cVEMP) and the extraocular muscles (the ocular VEMP or oVEMP) have proven useful in clinical assessment of vestibular function. VEMPs are commonly interpreted as a test of saccular function, based on neurophysiological evidence showing activation of saccular afferents by intense acoustic click stimuli. However, recent neurophysiological studies suggest that the clicks used in clinical VEMP tests activate vestibular end organs other than the saccule. To provide the neural basis for interpreting clinical VEMP testing results, the present study examined the extent to which air-conducted clicks differentially activate the various vestibular end organs at several intensities and durations in Sprague-Dawley rats. Single unit recordings were made from 562 vestibular afferents that innervated the otoliths [inferior branch otolith (IO) and superior branch otolith (SO)], the anterior canal (AC), the horizontal canal (HC), and the posterior canal (PC). Clicks higher than 60 dB SL (re-auditory brainstem response threshold) activated both semicircular canal and otolith organ afferents. Clicks at or below 60 dB SL, however, activated only otolith organ afferents. Longer duration clicks evoked larger responses in AC, HC, and SO afferents, but not in IO afferents. Intra-axonal recording and labeling confirmed that sound sensitive vestibular afferents innervated the horizontal and anterior canal cristae as well as the saccular and utricular maculae. Interestingly, all sound sensitive afferents are calyx-bearing fibers. These results demonstrate stimulus-dependent acoustic activation of both semicircular canals and otolith organs, and suggest that sound activation of vestibular end organs other than the saccule should not be ruled out when designing and interpreting clinical VEMP tests.

Bilateral loss of vestibular sensation can be disabling. We have shown that a multichannel vestibular prosthesis (MVP) can partly restore vestibular sensation as evidenced by improvements in the 3-dimensional angular vestibulo-ocular reflex (3D VOR). However, a key challenge is to minimize misalignment between the axes of eye and head rotation, which is apparently caused by current spread beyond each electrode's targeted nerve branch. We recently reported that rodents wearing a MVP markedly improve 3D VOR alignment during the first week after MVP activation, probably through the same central nervous system adaptive mechanisms that mediate cross-axis adaptation over time in normal individuals wearing prisms that cause visual scene movement about an axis different than the axis of head rotation. We hypothesized that rhesus monkeys would exhibit similar improvements with continuous prosthetic stimulation over time. We created bilateral vestibular deficiency in four rhesus monkeys via intratympanic injection of gentamicin. A MVP was mounted to the cranium, and eye movements in response to whole-body passive rotation in darkness were measured repeatedly over 1 week of continuous head motion-modulated prosthetic electrical stimulation. 3D VOR responses to whole-body rotations about each semicircular canal axis were measured on days 1, 3, and 7 of chronic stimulation. Horizontal VOR gain during 1 Hz, 50 °/s peak whole-body rotations before the prosthesis was turned on was <0.1, which is profoundly below normal (0.94 ± 0.12). On stimulation day 1, VOR gain was 0.4-0.8, but the axis of observed eye movements aligned poorly with head rotation (misalignment range ∼30-40 °). Substantial improvement of axis misalignment was observed after 7 days of continuous motion-modulated prosthetic stimulation under normal diurnal lighting. Similar improvements were noted for all animals, all three axes of rotation tested, for all sinusoidal frequencies tested (0.05-5 Hz), and for high-acceleration transient rotations. VOR asymmetry changes did not reach statistical significance, although they did trend toward slight improvement over time. Prior studies had already shown that directional plasticity reduces misalignment when a subject with normal labyrinths views abnormal visual scene movement. Our results show that the converse is also true: individuals receiving misoriented vestibular sensation under normal viewing conditions rapidly adapt to restore a well-aligned 3D VOR. Considering the similarity of VOR physiology across primate species, similar effects are likely to occur in humans using a MVP to treat bilateral vestibular deficiency.

Conclusions: The patients who received electric acoustic stimulation (EAS) cochlear implantation had relatively good vestibular function compared with the patients who did not have residual hearing. The vestibular function was well preserved after atraumatic EAS surgery. The round window approach and soft electrode are preferred to decrease the risk of impairing vestibular function., Objectives: The aim of this study was to examine the characteristic features of vestibular functions before and after implantations in patients undergoing EAS., Methods: Vestibular functions in patients who underwent EAS implantation were examined by caloric testing and vestibular evoked myogenic potential (VEMP) in 11 patients before and in 13 patients after implantation., Results: Preoperative evaluation showed that of the 11 patients, most (73%) had good vestibular function. One of 11 patients (9%) had decreased response in postoperative VEMP but all of the patients had unchanged results in postoperative caloric testing.

An implantable prosthesis that stimulates vestibular nerve branches to restore the sensation of head rotation and the three-dimensional (3D) vestibular ocular reflex (VOR) could benefit individuals disabled by bilateral loss of vestibular sensation. Our group has developed a vestibular prosthesis that partly restores normal function in animals by delivering biphasic current pulses via electrodes implanted in semicircular canals. Despite otherwise promising results, this approach has been limited by insufficient velocity of VOR response to head movements that should inhibit the implanted labyrinth and by misalignment between direction of head motion and prosthetically elicited VOR. We report that significantly larger VOR eye velocities in the inhibitory direction can be elicited by adapting a monkey to elevated baseline stimulation rate and current prior to stimulus modulation and then concurrently modulating ("co-modulating") both rate and current below baseline levels to encode inhibitory angular head velocity. Co-modulation of pulse rate and current amplitude above baseline can also elicit larger VOR eye responses in the excitatory direction than do either pulse rate modulation or current modulation alone. Combining these stimulation strategies with a precompensatory 3D coordinate transformation improves alignment and magnitude of evoked VOR eye responses. By demonstrating that a combination of co-modulation and precompensatory transformation strategies achieves a robust VOR response in all directions with significantly improved alignment in an animal model that closely resembles humans with vestibular loss, these findings provide a solid preclinical foundation for application of vestibular stimulation in humans.

The dorsolateral reticular formation of the caudal medulla, or the lateral tegmental field (LTF), has been classified as the brain's "vomiting center", as well as an important region in regulating sympathetic outflow. We examined the responses of LTF neurons in cats to rotations of the body that activate vestibular receptors, as well as to stimulation of baroreceptors (through mechanical stretch of the carotid sinus) and gastrointestinal receptors (through the intragastric administration of the emetic compound copper sulfate). Approximately half of the LTF neurons exhibited graviceptive responses to vestibular stimulation, similar to primary afferents innervating otolith organs. The other half of the neurons had complex responses, including spatiotemporal convergence behavior, suggesting that they received convergent inputs from a variety of vestibular receptors. Neurons that received gastrointestinal and baroreceptor inputs had similar complex responses to vestibular stimulation; such responses are expected for neurons that contribute to the generation of motion sickness. LTF units with convergent baroreceptor and vestibular inputs may participate in producing the cardiovascular system components of motion sickness, such as the changes in skin blood flow that result in pallor. The administration of copper sulfate often modulated the gain of responses of LTF neurons to vestibular stimulation, particularly for units whose spontaneous firing rate was altered by infusion of drug (median of 459%). The present results raise the prospect that emetic signals from the gastrointestinal tract modify the processing of vestibular inputs by LTF neurons, thereby affecting the probability that vomiting will occur as a consequence of motion sickness.

Since inner ear hair cells are mechano-electric transducers the control of hydrostatic pressure in the inner ear is crucial. Most studies analyzing dynamics and regulation of inner ear hydrostatic pressure performed pressure measurements in the cochlea. The present study is the first one reporting about absolute hydrostatic pressure values in the labyrinth. Hydrostatic pressure of the endolymphatic system was recorded in all three semicircular canals. Mean pressure values were 4.06 cmH(2)O ± 0.61 in the posterior, 3.36 cmH(2)O ± 0.94 in the anterior and 3.85 cmH(2)O ± 1.38 in the lateral semicircular canal. Overall hydrostatic pressure in the vestibular organ was 3.76 cmH(2)O ± 0.36. Endolymphatic hydrostatic pressure in all three semicircular canals is the same (p = 0.310). With regard to known endolymphatic pressure values in the cochlea from past studies vestibular pressure values are comparable to cochlear values. Until now it is not known whether the reuniens duct and the Bast's valve which are the narrowest passages in the endolymphatic system are open or closed. Present data show that most likely the endolymphatic system is a functionally open entity.

Directional asymmetries in vestibular reflexes have aided the diagnosis of vestibular lesions; however, potential asymmetries in vestibular perception have not been well defined. This investigation sought to measure potential asymmetries in human vestibular perception. Vestibular perception thresholds were measured in 24 healthy human subjects between the ages of 21 and 68 years. Stimuli consisted of a single cycle of sinusoidal acceleration in a single direction lasting 1 or 2 s (1 or 0.5 Hz), delivered in sway (left-right), surge (forward-backward), heave (up-down), or yaw rotation. Subject identified self-motion directions were analyzed using a forced choice technique, which permitted thresholds to be independently determined for each direction. Non-motion stimuli were presented to measure possible response bias. A significant directional asymmetry in the dynamic response occurred in 27% of conditions tested within subjects, and in at least one type of motion in 92% of subjects. Directional asymmetries were usually consistent when retested in the same subject but did not occur consistently in one direction across the population with the exception of heave at 0.5 Hz. Responses during null stimuli presentation suggested that asymmetries were not due to biased guessing. Multiple models were applied and compared to determine if sensitivities were direction specific. Using Akaike information criterion, it was found that the model with direction specific sensitivities better described the data in 86% of runs when compared with a model that used the same sensitivity for both directions. Mean thresholds for yaw were 1.3±0.9°/s at 0.5 Hz and 0.9±0.7°/s at 1 Hz and were independent of age. Thresholds for surge and sway were 1.7±0.8 cm/s at 0.5 Hz and 0.7±0.3 cm/s at 1.0 Hz for subjects <50 and were significantly higher in subjects >50 years old. Heave thresholds were higher and were independent of age.

This study investigated the impact of visual-vestibular stimulation on performance of an auditory information processing task in young and older adults. Performance on a spatial choice reaction time task was compared to performance on a non-spatial choice reaction time task. The tasks were performed during simultaneous rotational and moving visual stimulation. The non-spatial task was an auditory frequency discrimination task while the spatial task was a right-left lateralization task. Visual and vestibular conditions consisted of a non-movement baseline, sinusoidal earth-vertical axis rotation (EVAR) in darkness, off-vertical axis rotation (OVAR) in darkness at a constant velocity, OVAR in darkness with a sinusoidal profile, EVAR with a lighted visual surround, constant velocity optokinetic stimulation, and sinusoidal optokinetic stimulation. Baseline reaction times were subtracted from reaction times during each stimulus condition to yield "task cost", which was analyzed statistically. Subjects were healthy young (n=20; 24 ± 2.7 yrs; 10F) and older (n=29; 73 ± 6.0 yrs; 18F) adults. Results indicated that task cost was affected by the visual-vestibular condition and a task x condition interaction. There was no main effect of task or age group and no significant interaction with age. Otolithic stimulation and visual stimulation were associated with greater task cost compared to semicircular canal stimulation. Combining semicircular canal with otolithic or visual stimulation had no additional effect beyond otolithic or visual stimulation alone. This pattern of task cost being larger for otolith or visual vs. semicircular canal stimulation was found for both the spatial and non-spatial tasks. The significant interaction between condition and task type revealed that the task cost for the spatial task was larger than the task cost for the non-spatial task during visual conditions but not during non-visual conditions although the visual and non-visual conditions were not entirely comparable. This study suggests that interference of vestibular stimulation with cognitive processing is especially prominent for otolithic and visual stimulation although the strength of the various visual-vestibular stimuli may not have been uniform. Also, spatial tasks are more affected than non-spatial tasks during visual stimulation and interference between vestibular stimulation and cognitive processing is not age dependent for these relatively easy tasks.

In normal individuals, the vestibular labyrinths sense head movement and mediate reflexes that maintain stable gaze and posture. Bilateral loss of vestibular sensation causes chronic disequilibrium, oscillopsia, and postural instability. We describe a new multichannel vestibular prosthesis (MVP) intended to restore modulation of vestibular nerve activity with head rotation. The device comprises motion sensors to measure rotation and gravitoinertial acceleration, a microcontroller to calculate pulse timing, and stimulator units that deliver constant-current pulses to microelectrodes implanted in the labyrinth. This new MVP incorporates many improvements over previous prototypes, including a 50% decrease in implant size, a 50% decrease in power consumption, a new microelectrode array design meant to simplify implantation and reliably achieve selective nerve-electrode coupling, multiple current sources conferring ability to simultaneously stimulate on multiple electrodes, and circuitry for in vivo measurement of electrode impedances. We demonstrate the performance of this device through in vitro bench-top characterization and in vivo physiological experiments with a rhesus macaque monkey.

A fundamental concern of the brain is to establish the spatial relationship between self and the world to allow purposeful action. Response adaptation to unvarying sensory stimuli is a common feature of neural processing, both peripherally and centrally. For the semicircular canals, peripheral adaptation of the canal-cupula system to constant angular-velocity stimuli dominates the picture and masks central adaptation. Here we ask whether galvanic vestibular stimulation circumvents peripheral adaptation and, if so, does it reveal central adaptive processes. Transmastoidal bipolar galvanic stimulation and platform rotation (20 deg s−1) were applied separately and held constant for 2 min while perceived rotation was measured by verbal report. During real rotation, the perception of turn decayed from the onset of constant velocity with a mean time constant of 15.8 s. During galvanic-evoked virtual rotation, the perception of rotation initially rose but then declined towards zero over a period of ∼100 s. For both stimuli, oppositely directed perceptions of similar amplitude were reported when stimulation ceased indicating signal adaptation at some level. From these data the time constants of three independent processes were estimated: (i) the peripheral canal-cupula adaptation with time constant 7.3 s, (ii) the central ‘velocity-storage' process that extends the afferent signal with time constant 7.7 s, and (iii) a long-term adaptation with time constant 75.9 s. The first two agree with previous data based on constant-velocity stimuli. The third component decayed with the profile of a real constant angular acceleration stimulus, showing that the galvanic stimulus signal bypasses the peripheral transformation so that the brainstem sees the galvanic signal as angular acceleration. An adaptive process involving both peripheral and central processes is indicated. Signals evoked by most natural movements will decay peripherally before adaptation can exert an appreciable effect, making a specific vestibular behavioural role unlikely. This adaptation appears to be a general property of the internal coding of self-motion that receives information from multiple sensory sources and filters out the unvarying components regardless of their origin. In this instance of a pure vestibular sensation, it defines the afferent signal that represents the stationary or zero-rotation state.

The sense of orientation during locomotion is derived from our spatial relationship with the external environment, sensed predominantly by sight and sound, and from internal signals of motion, generated by the vestibular sense and the pattern of efferent and afferent signals to the muscles and joints. The sensory channels operate in different reference frames and have different time-dependent adaptive properties and yet the inputs are combined by the central nervous system to create an internal representation of self-motion. In normal circumstances vestibular, visual and proprioceptive cues provide congruent information on locomotor trajectory; however, in cases of sensory discord there must be a recalibration of sensory signals to provide a unitary representation. We develop a means of studying these fusion processes by perturbing each channel in isolation about a consistent behavioural axis. This review focuses on creating the vestibular perturbation of the orientation sense by transmastoidal galvanic stimulation, a technique generally used to evoke balance reflexes. Vector summation across the population of semicircular canal afferents creates a net signal that is interpreted by the brain as a vector of angular acceleration in a craniocentric reference frame. The signal feeds perceptual processes of orientation after transformation that resolves the 3-D signal onto the terrestrial or behavioural plane. Changing head posture changes the interpretation of the galvanic vestibular signal for balance and orientation responses. With appropriate head alignments during locomotion, the galvanic stimulus can be used to either steer trajectory over the terrestrial plane or perturb balance.

We report a precise, simple, and accessible method of mathematically measuring and modeling the three-dimensional (3D) geometry of semicircular canals (SCCs) in living humans. Knowledge of this geometry helps understand the development and physiology of SCC stimulation. We developed a framework of robust techniques that automatically and accurately reconstruct SCC geometry from computed tomography (CT) images and are directly validated using micro-CT as ground truth. This framework measures the 3D centroid paths of the bony SCCs allowing direct comparison and analysis between ears within and between subjects. An average set of SCC morphology is calculated from 34 human ears, within which other geometrical attributes such as nonplanarity, radius of curvature, and inter-SCC angle are examined, with a focus on physiological implications. These measurements have also been used to critically evaluate plane fitting techniques that reconcile many of the discrepancies in current SCC plane studies. Finally, we mathematically model SCC geometry using Fourier series equations. This work has the potential to reinterpret physiology and pathophysiology in terms of real individual 3D morphology.

Background: The aim was to explore vestibular responses to bone vibration-induced nystagmus (BVIN) during bed-side examination in patients with otosclerosis and conductive hearing loss. It is assumed that vibration of the mastoid (at 100 Hz) stimulates all vestibular end organs (semicircular canals and otolith structures). Previous studies described oculomotor responses to vestibular activation by vibratory stimulus. Stimulation of individual semicircular canals produces eye movement in the plane of the stimulated canal. Vibratory stimulation of otolith structures (utricular macula) produces changes in ocular torsional position. Otosclerosis is responsible for conductive hearing loss and symptoms such as dizziness and vertigo. Diagnosis is usually by a combination of family medical history, conductive hearing loss pattern, absence of air-VEMP, and CT scan of the temporal bone., Material/methods: Eye movements of patients diagnosed with otosclerosis and conductive hearing loss were recorded in complete darkness. These oculomotor responses to long-lasting unilateral vibratory stimulation applied to the mastoid surface were evaluated and the patterns of vertical, torsional, and horizontal eye velocity and eye position were measured by three-dimensional computerized infrared video oculography., Results: Mastoid vibration evoked responses in all cases, mostly demonstrating excitation of the affected side. In all cases, three-dimensional infrared video oculography showed mostly horizontal nystagmus directed with the slow phase to the healthy side., Conclusions: Three-dimensional analysis of BVIN in patients with otosclerosis and conductive hearing loss may be appropriate in evaluating the vestibular function and altered immittance related with this clinical entity. This promises to be an interesting new field of research.

Angular head movements in vertebrates are detected by the three semicircular canals of the inner ear and their associated sensory tissues, the cristae. Bone morphogenetic protein 4 (Bmp4), a member of the Transforming growth factor family (TGF-beta), is conservatively expressed in the developing cristae in several species, including zebrafish, frog, chicken, and mouse. Using mouse models in which Bmp4 is conditionally deleted within the inner ear, as well as chicken models in which Bmp signaling is knocked down specifically in the cristae, we show that Bmp4 is essential for the formation of all three cristae and their associated canals. Our results indicate that Bmp4 does not mediate the formation of sensory hair and supporting cells within the cristae by directly regulating genes required for prosensory development in the inner ear such as Serrate1 (Jagged1 in mouse), Fgf10, and Sox2. Instead, Bmp4 most likely mediates crista formation by regulating Lmo4 and Msx1 in the sensory region and Gata3, p75Ngfr, and Lmo4 in the non-sensory region of the crista, the septum cruciatum. In the canals, Bmp2 and Dlx5 are regulated by Bmp4, either directly or indirectly. Mechanisms involved in the formation of sensory organs of the vertebrate inner ear are thought to be analogous to those regulating sensory bristle formation in Drosophila. Our results suggest that, in comparison to sensory bristles, crista formation within the inner ear requires an additional step of sensory and non-sensory fate specification., Competing Interests: The authors have declared that no competing interests exist.

Activation of sympathetic neural traffic via the vestibular system is referred to as the vestibulosympathetic reflex. Investigations of the vestibulosympathetic reflex in humans have been limited to the past decade, and the importance of this reflex in arterial blood pressure regulation is still being determined. This review provides a summary of sympathetic neural responses to various techniques used to engage the vestibulosympathetic reflex. Studies suggest that activation of the semicircular canals using caloric stimulation and yaw rotation do not modulate muscle sympathetic nerve activity (MSNA) or skin sympathetic nerve activity (SSNA). In contrast, activation of the otolith organs appear to alter MSNA, but not SSNA. Specifically, head-down rotation and off-vertical axis rotation increase MSNA, while sinusoidal linear accelerations decrease MSNA. Galvanic stimulation, which results in a nonspecific activation of the vestibule, appears to increase MSNA if the mode of delivery is pulse trained. In conclusion, evidence strongly supports the existence of a vestibulosympathetic reflex in humans. Furthermore, attenuation of the vestibulosympathetic reflex is coupled with a drop in arterial blood pressure in the elderly, suggesting this reflex may be important in human blood pressure regulation.

Objective: This research evaluates effects of vestibular stimulation by Comprehensive Motion Apparatus (CMA) in ADHD., Method: Children ages 6 to 12 (48 boys, 5 girls) with ADHD were randomized to thrice-weekly 30-min treatments for 12 weeks with CMA, stimulating otoliths and semicircular canals, or a single-blind control of equal duration and intensity, each treatment followed by a 20-min typing tutorial., Results: In intent-to-treat analysis (n = 50), primary outcome improved significantly in both groups (p = .0001, d = 1.09 to 1.30), but treatment difference not significant (p = .7). Control children regressed by follow-up (difference p = .034, d = 0.65), but overall difference was not significant (p = .13, d = .47). No measure showed significant treatment differences at treatment end, but one did at follow-up. Children with IQ-achievement discrepancy > or = 1 SD showed significantly more CMA advantage on three measures., Conclusion: This study illustrates the importance of a credible control condition of equal duration and intensity in trials of novel treatments. CMA treatment cannot be recommended for combined-type ADHD without learning disorder.

Central vestibular neurons process head movement-related sensory signals over a wide dynamic range. In the isolated frog whole brain, second-order vestibular neurons were identified by monosynaptic responses after electrical stimulation of individual semicircular canal nerve branches. Neurons were classified as tonic or phasic vestibular neurons based on their different discharge patterns in response to positive current steps. With increasing frequency of sinusoidally modulated current injections, up to 100 Hz, there was a concomitant decrease in the impedance of tonic vestibular neurons. Subthreshold responses as well as spike discharge showed classical low-pass filter-like characteristics with corner frequencies ranging from 5 to 20 Hz. In contrast, the impedance of phasic vestibular neurons was relatively constant over a wider range of frequencies or showed a resonance at approximately 40 Hz. Above spike threshold, single spikes of phasic neurons were synchronized with the sinusoidal stimulation between approximately 20 and 50 Hz, thus showing characteristic bandpass filter-like properties. Both the subthreshold resonance and bandpass filter-like discharge pattern depend on the activation of an I(D) potassium conductance. External current or synaptic stimulation that produced impedance increases (i.e., depolarization in tonic or hyperpolarization in phasic neurons) had opposite and complementary effects on the responses of the two types of neurons. Thus, membrane depolarization by current steps or repetitive synaptic excitation amplified synaptic inputs in tonic vestibular neurons and reduced them in phasic neurons. These differential, opposite membrane response properties render the two neuronal types particularly suitable for either integration (tonic neurons) or signal detection (phasic neurons), respectively, and dampens variations of the resting membrane potential in the latter.

The vestibular semicircular canals are responsible for detection of rotational head motion although the precise mechanisms underlying the transduction and encoding of movement information are still under study. In the present investigation, we utilized neural tracers and immunohistochemistry to quantitatively examine the topology and afferent innervation patterns of the horizontal semicircular canal crista (HCC) in pigeons (Columba livia). Two hundred and eighty-six afferents from five horizontal canal organs were identified of which 92 units were sufficiently labeled and isolated to perform anatomical reconstructions. In addition, a three-dimensional contour map of the crista was constructed. Bouton afferents were located only in the peripheral regions of the receptor epithelium. Bouton afferents had the most complex innervation patterns with significantly longer and more numerous branches as well as a higher branch order than any other fiber type. Bouton fibers also contained significantly more bouton terminals than did dimorph afferents. Calyx afferents were located only in the apex and central planar regions. Calyx fibers had the largest axonal diameters yet the smallest fiber lengths and innervation areas, the fewest number of branches, the lowest branch order, and the fewest total number of terminals of all fiber types. Dimorph afferents were located throughout the central crista with afferent terminations that were larger and more complex than calyx fibers but less so than bouton fibers. Overall, the pigeon HCC morphology and innervation shares many common features with those of other animal classes.

The main objective of this study was to determine whether bone-conducted vibration (BCV) is equally effective in activating both semicircular canal and otolith afferents in the guinea pig or whether there is preferential activation of one of these classes of vestibular afferents. To answer this question a large number (346) of single primary vestibular neurons were recorded extracellularly in anesthetized guinea pigs and were identified by their location in the vestibular nerve and classed as regular or irregular on the basis of the variability of their spontaneous discharge. If a neuron responded to angular acceleration it was classed as a semicircular canal neuron, if it responded to maintained roll or pitch tilts it was classified as an otolith neuron. Each neuron was then tested by BCV stimuli-either clicks, continuous pure tones (200-1,500 Hz) or short tone bursts (500 Hz lasting 7 ms)-delivered by a B-71 clinical bone-conduction oscillator cemented to the guinea pig's skull. All stimulus intensities were referred to that animal's own auditory brainstem response (ABR) threshold to BCV clicks, and the maximum intensity used was within the animal's physiological range and was usually around 70 dB above BCV threshold. In addition two sensitive single axis linear accelerometers cemented to the skull gave absolute values of the stimulus acceleration in the rostro-caudal direction. The criterion for a neuron being classed as activated was an audible, stimulus-locked increase in firing rate (a 10% change was easily detectable) in response to the BCV stimulus. At the stimulus levels used in this study, semicircular canal neurons, both regular and irregular, were insensitive to BCV stimuli and very few responded: only nine of 189 semicircular canal neurons tested (4.7%) showed a detectable increase in firing in response to BCV stimuli up to the maximum 2 V peak-to-peak level we delivered to the B-71 oscillator (which produced a peak-to-peak skull acceleration of around 6-8 g and was usually around 60-70 dB above the animal's own ABR threshold for BCV clicks). Regular otolithic afferents likewise had a poor response; only 14 of 99 tested (14.1%) showed any increase in firing rate up to the maximum BCV stimulus level. However, most irregular otolithic afferents (82.8%) showed a clear increase in firing rate in response to BCV stimuli: of the 58 irregular otolith neurons tested, 48 were activated, with some being activated at very low intensities (only about 10 dB above the animal's ABR threshold to BCV clicks). Most of the activated otolith afferents were in the superior division of the vestibular nerve and were probably utricular afferents. That was confirmed by evidence using juxtacellular injection of neurobiotin near BCV activated neurons to trace their site of origin to the utricular macula. We conclude there is a very clear preference for irregular otolith afferents to be activated selectively by BCV stimuli at low stimulus levels and that BCV stimuli activate some utricular irregular afferent neurons. The BCV generates compressional and shear waves, which travel through the skull and constitute head accelerations, which are sufficient to stimulate the most sensitive otolithic receptor cells.