Natural Vestibular Stimulation Research Articles

Directional effects of whole-body spinning and visual flow in virtual reality on vagal neuromodulation.

Neural circuits allow whole-body yaw rotation to modulate vagal parasympathetic activity, which alters beat-to-beat variation in heart rate. The overall output of spinning direction, as well as vestibular-visual interactions on vagal activity still needs to be investigated. This study investigated direction-dependent effects of visual and natural vestibular stimulation on two autonomic responses: heart rate variability (HRV) and pupil diameter. Healthy human male subjects (n = 27) underwent constant whole-body yaw rotation with eyes open and closed in the clockwise (CW) and anticlockwise (ACW) directions, at 90°/s for two minutes. Subjects also viewed the same spinning environments on video in a VR headset. CW spinning significantly decreased parasympathetic vagal activity in all conditions (CW open p = 0.0048, CW closed p = 0.0151, CW VR p = 0.0019,), but not ACW spinning (ACW open p = 0.2068, ACW closed p = 0.7755, ACW VR p = 0.1775,) as indicated by an HRV metric, the root mean square of successive RR interval differences (RMSSD). There were no direction-dependent effects of constant spinning on sympathetic activity inferred through the HRV metrics, stress index (SI), sympathetic nervous system index (SNS index) and pupil diameter. Neuroplasticity in the CW eyes closed and CW VR conditions post stimulation was observed. Only one direction of yaw spinning, and visual flow caused vagal nerve neuromodulation and neuroplasticity, resulting in an inhibition of parasympathetic activity on the heart, to the same extent in either vestibular or visual stimulation. These results indicate that visual flow in VR can be used as a non-electrical method for vagus nerve inhibition without the need for body motion in the treatment of disorders with vagal overactivity. The findings are also important for VR and spinning chair based autonomic nervous system modulation protocols, and the effects of motion integrated VR.

Vestibular-Evoked Cerebral Potentials.

The human vestibular cortex has mostly been approached using functional magnetic resonance imaging and positron emission tomography combined with artificial stimulation of the vestibular receptors or nerve. Few studies have used electroencephalography and benefited from its high temporal resolution to describe the spatiotemporal dynamics of vestibular information processing from the first milliseconds following vestibular stimulation. Evoked potentials (EPs) are largely used to describe neural processing of other sensory signals, but they remain poorly developed and standardized in vestibular neuroscience and neuro-otology. Yet, vestibular EPs of brainstem, cerebellar, and cortical origin have been reported as early as the 1960s. This review article summarizes and compares results from studies that have used a large range of vestibular stimulation, including natural vestibular stimulation on rotating chairs and motion platforms, as well as artificial vestibular stimulation (e.g., sounds, impulsive acceleration stimulation, galvanic stimulation). These studies identified vestibular EPs with short latency (<20 ms), middle latency (from 20 to 50 ms), and late latency (>50 ms). Analysis of the generators (source analysis) of these responses offers new insights into the neuroimaging of the vestibular system. Generators were consistently found in the parieto-insular and temporo-parietal junction—the core of the vestibular cortex—as well as in the prefrontal and frontal areas, superior parietal, and temporal areas. We discuss the relevance of vestibular EPs for basic research and clinical neuroscience and highlight their limitations.

Pain modulation by illusory body rotation: A new way to disclose the interaction between the vestibular system and pain processing.

Clinical and experimental evidence advocates a structural and functional link between the vestibular and other sensory systems. For instance, visuo-vestibular and vestibular-somatosensory interactions have been widely reported. However, whether visual inputs carrying vestibular information can modulate pain is not yet clear. Recent evidence using natural vestibular stimulation or moving visual stimuli, points at an unspecific effect of distraction. By using immersive virtual reality (VR), we created a new way to prompt the vestibular system through the vision of static visual cues, studying the possible interaction with pain. Twenty-four healthy participants were visually immersed in a virtual room which could appear with five different degrees of rotation in the sagittal axis, either towards the right, left or with no rotation. Participants' heat pain thresholds and subjective reports of perceived body rotation, sense of presence and attention were measured. 'Being' in a tilted room induced the sensation of body rotation in our participants, even though they were always in an upright position. We also found that rotating the visual scenario can modulate the participants' pain thresholds, determining a significant increase when a left tilt is displayed. In addition,a positive correlation between the perceived body midline rotation and pain threshold wasfoundwhen the virtual room was titled 15 degrees toward the left. Importantly, all VR conditions were found to beequally distractive. Vestibular information present in static visual cues can modulate experimentally-induced acute pain according to a side-dependent manner and bypassing supramodal attentional mechanisms. These findings may help refining pain management approaches based on multimodal stimulation. This study explored how the visualization ofstatic environments in immersive virtual reality can lead to pain threshold modulation through the activation of the vestibular system. Immersion into rotated virtualenvironments led to the illusorysensation of body rotation, and this sensation was found to be related witha modulationof pain perception. Possible analgesic effectsdue to distractioncould beruled out.These results expandour current knowledge about how the visual, vestibular and somatosensory (pain) systems interact. These findings may influence future pain treatmentstrategies based on multisensory stimulation.

PET Visualized Stimulation of the Vestibular Organ in Menière's Disease

Introduction: The cortical metabolic activity in patients with Menière's disease has not been investigated. The aim of this study was to investigate the 18F-FDG cerebral uptake in Menière's patients compared to healthy controls.Method: Eight patients with right-sided Menière's disease and fourteen healthy controls underwent a video head impulse test (vHIT), test of utricular function with ocular vestibular evoked myogenic potentials (oVEMP) and three 18F-FDG-based PET examinations of the brain. Participants were seated in a self-propelled chair, injected with 18F-FDG and then exposed to 35 min of chair motion stimulation, followed by a PET scan. Two types of natural vestibular stimuli were applied, predominantly toward the right horizontal semicircular canal (angular acceleration) and right utriculus (linear acceleration). For baseline scans, participants were injected with 18F-FDG while seated without movement.Results: Analyses of baseline scans revealed decreased 18F-FDG-uptake in the medial part of Heschl's gyrus in the left hemisphere in patients with Menière's disease compared to healthy controls. During angular vestibular stimulation there was also a significantly decreased 18F-FDG uptake in the intersection between the medial part of Heschl's gyrus and the parietal operculum in the left hemisphere and bilaterally in the posterior part of insula. During linear stimulation, Menière's patients showed decreased 18F-FDG uptake in the medial part of Heschl's gyrus in the right hemisphere and also bilaterally in the posterior insula. In addition, decreased 18F-FDG uptake was seen in the thalamus during vestibular stimulation.Conclusion: Heschl's gyrus, the posterior part of insula, and thalamus have previously been shown to be core areas for processing vestibular inputs. Patients with Menière's disease solely differed from the healthy controls with lower cortical activity in these areas at baseline and during natural vestibular stimulation.

Positron emission tomography visualized stimulation of the vestibular organ is localized in Heschl's gyrus.

The existence of a human primary vestibular cortex is still debated. Current knowledge mainly derives from functional magnetic resonance imaging (fMRI) and positron emission tomography (PET) acquisitions during artificial vestibular stimulation. This may be problematic as artificial vestibular stimulation entails coactivation of other sensory receptors. The use of fMRI is challenging as the strong magnetic field and loud noise during MRI may both stimulate the vestibular organ. This study aimed to characterize the cortical activity during natural stimulation of the human vestibular organ. Two fluorodeoxyglucose (FDG)‐PET scans were obtained after natural vestibular stimulation in a self‐propelled chair. Two types of stimuli were applied: (a) rotation (horizontal semicircular canal) and (b) linear sideways movement (utriculus). A comparable baseline FDG‐PET scan was obtained after sitting motion‐less in the chair. In both stimulation paradigms, significantly increased FDG uptake was measured bilaterally in the medial part of Heschl's gyrus, with some overlap into the posterior insula. This is the first neuroimaging study to visualize cortical processing of natural vestibular stimuli. FDG uptake was demonstrated in the medial‐most part of Heschl's gyrus, normally associated with the primary auditory cortex. This anatomical localization seems plausible, considering that the labyrinth contains both the vestibular organ and the cochlea.

Canal-otolith interactions alter the perception of self-motion direction.

Few studies have investigated the perception of vestibular stimuli when they occur in sequences. Here, three experiments (ntotal = 33) are presented that focus on intravestibular motion sequences and the underlying perceptual decision-making process. Natural vestibular stimulation (yaw rotation or translation) was used to investigate the discrimination process of the direction of a subsequent spatially congruent or incongruent translation or rotation. The few existing studies focusing on unimodal motion sequences have uncovered self-motion aftereffects, similar to the visual motion aftereffect, possibly due to altered processing of sensory stimuli. An alternative hypothesis predicts a shift of spatial attention due to the cue motion influencing perception of the subsequent motion stimulus. The results show that participants systematically misjudged the direction of motion stimuli well above the detection threshold if the direction of the preceding cue motion stimulus was congruent with the direction of the target (a motion aftereffect). Hierarchical drift diffusion models were used to analyze the data. The results suggest that altered perceptual decision-making and the resulting misperceptions are likely to originate in altered processing of sensory vestibular information.

Effect of Natural Vestibular Stimulation on Self-Esteem Levels in Relation to Stress

Effect of Natural Vestibular Stimulation on Self-Esteem Levels in Relation to Stress

Oscillatory neural responses evoked by natural vestibular stimuli in humans.

While there have been numerous studies of the vestibular system in mammals, less is known about the brain mechanisms of vestibular processing in humans. In particular, of the studies that have been carried out in humans over the last 30 years, none has investigated how vestibular stimulation (VS) affects cortical oscillations. Here we recorded high-density electroencephalography (EEG) in healthy human subjects and a group of bilateral vestibular loss patients (BVPs) undergoing transient and constant-velocity passive whole body yaw rotations, focusing our analyses on the modulation of cortical oscillations in response to natural VS. The present approach overcame significant technical challenges associated with combining natural VS with human electrophysiology and reveals that both transient and constant-velocity VS are associated with a prominent suppression of alpha power (8-13 Hz). Alpha band suppression was localized over bilateral temporo-parietal scalp regions, and these alpha modulations were significantly smaller in BVPs. We propose that suppression of oscillations in the alpha band over temporo-parietal scalp regions reflects cortical vestibular processing, potentially comparable with alpha and mu oscillations in the visual and sensorimotor systems, respectively, opening the door to the investigation of human cortical processing under various experimental conditions during natural VS.

Changing perspective: The role of vestibular signals

Social interactions depend on mechanisms such as the ability to take another person's viewpoint, i.e. visuo-spatial perspective taking. However, little is known about the sensorimotor mechanisms underpinning perspective taking. Because vestibular signals play roles in mental rotation and spatial cognition tasks and because damage to the vestibular cortex can disturb egocentric perspective, vestibular signals stand as important candidates for the sensorimotor foundations of perspective taking. Yet, no study merged natural full-body vestibular stimulations and explicit visuo-spatial perspective taking tasks in virtual environments. In Experiment 1, we combined natural vestibular stimulation on a rotatory chair with virtual reality to test how vestibular signals are processed to simulate the viewpoint of a distant avatar. While they were rotated, participants tossed a ball to a virtual character from the viewpoint of a distant avatar. Our results showed that vestibular signals influence perspective taking in a direction-specific way: participants were faster when their physical body rotated in the same direction as the mental rotation needed to take the avatar's viewpoint. In Experiment 2, participants realized 3D object mental rotations, which did not involve perspective taking, during the same whole-body vestibular stimulation. Our results demonstrated that vestibular stimulation did not affect 3D object mental rotations. Altogether, these data indicate that vestibular signals have a direction-specific influence on visuo-spatial perspective taking (self-centered mental imagery), but not a general effect on mental imagery. Findings from this study suggest that vestibular signals contribute to one of the most crucial mechanisms of social cognition: understanding others’ actions.

Balancing awareness: Vestibular signals modulate visual consciousness in the absence of awareness.

The processing of visual and vestibular information is crucial for perceiving self-motion. Visual cues, such as optic flow, have been shown to induce and alter vestibular percepts, yet the role of vestibular information in shaping visual awareness remains unclear. Here we investigated if vestibular signals influence the access to awareness of invisible visual signals. Using natural vestibular stimulation (passive yaw rotations) on a vestibular self-motion platform, and optic flow masked through continuous flash suppression (CFS) we tested if congruent visual-vestibular information would break interocular suppression more rapidly than incongruent information. We found that when the unseen optic flow was congruent with the vestibular signals perceptual suppression as quantified with the CFS paradigm was broken more rapidly than when it was incongruent. We argue that vestibular signals impact the formation of visual awareness through enhanced access to awareness for congruent multisensory stimulation.

Restricted Neural Plasticity in Vestibulospinal Pathways after Unilateral Labyrinthectomy as the Origin for Scoliotic Deformations

Adolescent idiopathic scoliosis in humans is often associated with vestibulomotor deficits. Compatible with a vestibular origin, scoliotic deformations were provoked in adult Xenopus frogs by unilateral labyrinthectomy (UL) at larval stages. The aquatic ecophysiology and absence of body-weight-supporting limb proprioceptive signals in amphibian tadpoles as a potential sensory substitute after UL might be the cause for a persistent asymmetric descending vestibulospinal activity. Therefore, peripheral vestibular lesions in larval Xenopus were used to reveal the morphophysiological alterations at the cellular and network levels. As a result, spinal motor nerves that were modulated by the previously intact side before UL remained permanently silent during natural vestibular stimulation after the lesion. In addition, retrograde tracing of descending pathways revealed a loss of vestibular neurons on the ipsilesional side with crossed vestibulospinal projections. This loss facilitated a general mass imbalance in descending premotor activity and a permanent asymmetric motor drive to the axial musculature. Therefore, we propose that the persistent asymmetric contraction of trunk muscles exerts a constant, uncompensated differential mechanical pull on bilateral skeletal elements that enforces a distortion of the soft cartilaginous skeletal elements and bone shapes. This ultimately provokes severe scoliotic deformations during ontogenetic development similar to the human syndrome.

Alexander's law during high-acceleration head rotations in humans

Alexander's law states that the amplitude of the spontaneous nystagmus grows with increasing gaze in the direction of the fast phase. Using the search-coil method we employed head impulses at various eye-in-orbit azimuth angles to test (i) whether the normal vestibulo-ocular reflex (VOR) in the behaviorally relevant high-frequency range has intrinsic properties that could account for Alexander's law and (ii) whether such properties can also be shown in patients with unilateral vestibulopathy. We showed that the gain of the VOR remained unaffected by eye-in-orbit position in contols and in patients, both on ipsilesional and contralesional stimuli. These findings suggest that eye-in-orbit position does not directly modulate the activity in VOR pathways, neither during unbalanced but reciprocal (in controls), nor during unbalanced and nonreciprocal natural vestibular stimulation (in patients).

Responses of primary vestibular neurons to galvanic vestibular stimulation (GVS) in the anaesthetised guinea pig

Previous studies in humans and animals which have shown that DC galvanic vestibular stimulation (GVS) induces horizontal and torsional eye movements have been interpreted as being due to a preferential activation of primary vestibular afferents innervating the horizontal semicircular canals and otoliths by GVS. The present study sought to determine in guinea pigs whether GVS does indeed selectively activate primary horizontal canal and otolith afferents. Constant-current GVS was passed between electrodes implanted in the tensor-tympani muscle of each middle ear or between electrodes on the skin over the mastoid. During this stimulation, responses from single primary vestibular neurons were recorded extracellularly by glass microelectrodes in Scarpa's ganglion. Afferents from all vestibular sensory regions were activated by both surface and tensor-tympani galvanic stimulation. Tensor tympani GVS was approximately 10 times more effective than surface GVS. At larger current intensities irregularly discharging afferents showed an asymmetrical response: cathodal stimulation resulted in a larger change in firing (increase) than anodal stimulation (decrease), whereas regularly discharging afferents responded symmetrically to the two polarities of GVS. Across all afferents tuned for different types of natural vestibular stimulation, neuronal sensitivity for GVS was found to increase with discharge variability (as indexed by CV*). Anterior canal afferents showed a slightly higher sensitivity than afferents from other vestibular sensory regions. Hence, the present study concluded that GVS activates primary vestibular afferents innervating all sensory regions in a uniform fashion. Therefore, the specific pattern of GVS-induced eye movements reported in previous studies are not due to differential sensitivity between different vestibular sensory regions, but are likely to reflect an involvement of central processing.

Ultrastructural Changes in the Mixed Synapses of Mauthner Neurons Related to Long-Term Potentiation and Natural Modification of the Motor Function

We examined changes in the ultrastructure of afferent mixed synapses on the membrane of Mauthner neurons (M cells) of the goldfish, which were related to two functional states, long-term potentiation (LTP) of the electrotonic response (a model form of the memory trace) and adaptation (resistivity to fatigue resulting from long-lasting motor training and considered a natural form of the memory trace manifested on the neuronal level). LTP was induced in medullary slices using high-frequency electrical stimulation of the afferent input. Adaptation was produced using natural vestibular stimulation (everyday motor training, which modified motor behavior of the fish and function of the M cell). It was supposed that if the LTP phenomenon is involved in the formation of natural memory, both the adaptation and the LTP states should be accompanied by similar specific structural modifications. Indeed, it was found that in both cases the number of fibrillar bridges in the gaps of desmosome-like contacts (DLC) in the mixed synapses on the M cell surface demonstrated an about twofold increase. These bridges are known to include actin filaments, which function as conductors of cationic signals; thus, the LTP-related increase in the density of bridges corresponds to increased efficacy of electrotonic coupling via mixed synapses. Such a structural correlate of LTP, which probably has the same functional significance in mixed synapses of the “adapted” M cells, allows us to suppose that LTP is a natural property of the nervous system. The LTP-type intensification of the relay function of mixed synapses, which corresponds to adaptation, is probably a compensatory rearrangement allowing M cells to maintain some balance of the synaptic influences and, at the same time, to remain in a stable and plastic state; this is necessary for stable functioning under changing environmental conditions.

Vestibularly evoked climbing-fiber responses modulate simple spikes in rabbit cerebellar Purkinje neurons.

The nodulus receives a primary vestibular afferent input from the ipsilateral labyrinth and a vestibularly related climbing-fiber input originating from the contralateral labyrinth. Previously we demonstrated that increased discharge of vestibularly evoked climbing-fiber responses (CFRs) in nodular Purkinje cells was correlated with decreased discharge of simple spikes (SSs). This left unresolved the question of whether vestibularly evoked antiphasic behavior of CFRs and SSs reflects a common neural mechanism or the activation of two separate parallel pathways. We answered this question using natural vestibular stimulation to modulate the discharge of uvula-nodular Purkinje cells recorded extracellularly in unilaterally labyrinthectomized, chloralose urethane-anesthetized rabbits. In such animals, vestibular primary afferents projecting to the uvula-nodulus as mossy fibers remained intact on the side contralateral to the unilateral labyrinthectomy. The discharge of CFRs recorded in ipsilateral nodular Purkinje cells was increased by ipsilateral roll-tilt while the discharge of SSs was increased by contralateral roll-tilt. These polarities were reversed for Purkinje cells recorded in the contralateral uvula-nodulus. The polarity of SS discharge recorded from Purkinje cells on both sides of the nodulus was opposite to that of the vestibular primary mossy-fiber afferents. SSs continued to respond to contralateral roll-tilt even when the primary vestibular afferent mossy-fiber pathway was destroyed by the unilateral labyrinthectomy. Although the discharge of SSs recorded in the contralateral uvula-nodulus was increased by contralateral roll-tilt, this modulation was reduced relative to that observed in Purkinje cells recorded in the ipsilateral uvula-nodulus. We conclude that vestibularly evoked CFRs caused the modulation of SS discharge.

Short-term (FOS) and long-term (FRA) protein expression in rat locus coeruleus neurons during the neurolab mission: contribution of altered gravitational fields, stress, and other factors

Changes in immediate-early gene (IEG) expression during and after space flight were studied in the rat locus coeruleus (LC) during the NASA Neurolab mission. The LC sends widespread projections throughout the brain and releases the neuromodulator norepinephrine. LC neurons respond to natural vestibular stimulation; their firing rate also increases during waking and decreases or ceases during sleep. LC neurons express IEGs such as c-fos after activation. Adult male albino Fisher 344 rats were killed at four mission time points, and the number of Fos- and Fos-related antigen (FRA)-positive LC cells were counted in flight and ground-based control rats. Half of the subjects at each time point were exposed to light for 60 min prior to killing to standardize their sleep–waking state. FRA-expressing cells were more numerous than Fos-expressing cells in both flight- and ground-based subjects. The difference between FRA- and Fos-expressing cells within individuals was significantly larger 24 h after landing in subjects exposed to both space flight and light pulse than in all other subjects at any mission time point. Fos and FRA responses scaled in proportion to the maximum response observed in any single individual showed similar patterns of variation. Analysis of the scaled and combined responses showed that LC IEG levels responded to both gravity changes and light pulses. Subjects exposed to either single stimulus had equivalent responses, significantly greater than those of control subjects maintained in dim light. The combination of gravity change and light pulse gave significantly higher LC responses than either stimulus alone 24 h after takeoff, and to a lesser extent after 12 days in space; the highest responses were obtained 24 h after landing. By 14 days after landing, animals exposed to space flight and light pulse responded no differently than ground-based subjects. No difference in LC IEG expression was clearly attributable to changes in the sleep–waking state of subjects. Activity of noradrenergic LC neurons has been previously shown to modulate IEG expression in target structures. The increased IEG LC activity (seen most especially 24 h after landing) may reflect large-scale activation of noradrenergic neurons that may serve as a trigger for molecular changes in target structures, and be critical for adaptation to gravity changes.

Cerebellar nodulectomy impairs spatial memory of vestibular and optokinetic stimulation in rabbits.

Natural vestibular and optokinetic stimulation were used to investigate the possible role of the cerebellar nodulus in the regulation and modification of reflexive eye movements in rabbits. The nodulus and folium 9d of the uvula were destroyed by surgical aspiration. Before and after nodulectomy the vertical and horizontal vestibuloocular reflexes (VVOR, HVOR) were measured during sinusoidal vestibular stimulation about the longitudinal (roll) and vertical (yaw) axes. Although the gain of the HVOR (G(HVOR) = peak eye movement velocity/peak head velocity) was not affected by the nodulectomy, the gain of the VVOR (G(VVOR)) was reduced. The gains of the vertical and horizontal optokinetic reflexes (G(VOKR), G(HOKR)) were measured during monocular, sinusoidal optokinetic stimulation (OKS) about the longitudinal and vertical axes. Following nodulectomy, there was no reduction in G(VOKR) or G(HOKR). Long-term binocular OKS was used to generate optokinetic afternystagmus, OKAN II, that lasts for hours. After OKAN II was induced, rabbits were subjected to static pitch and roll, to determine how the plane and velocity of OKAN II is influenced by a changing vestibular environment. During static pitch, OKAN II slow phase remained aligned with earth-horizontal. This was true for normal and nodulectomized rabbits. During static roll, OKAN II remained aligned with earth-horizontal in normal rabbits. During static roll in nodulectomized rabbits, OKAN II slow phase developed a centripetal vertical drift. We examined the suppression and recovery of G(VVOR) following exposure to conflicting vertical OKS for 10-30 min. This vestibular-optokinetic conflict reduced G(VVOR) in both normal and nodulectomized rabbits. The time course of recovery of G(VVOR) after conflicting OKS was the same before and after nodulectomy. In normal rabbits, the head pitch angle, at which peak OKAN II velocity occurred, corresponded to the head pitch angle maintained during long-term OKS. If the head was maintained in a "pitched-up" or "pitched-down" orientation during long-term OKS, the subsequently measured OKAN II peak velocity occurred at the same orientation. This was not true for nodulectomized rabbits, who had OKAN II peak velocities at head pitch angles independent of those maintained during long-term OKS. We conclude that the nodulus participates in the regulation of compensatory reflexive movements. The nodulus also influences "remembered" head position in space derived from previous optokinetic and vestibular stimulation.

Local vestibular blood flow and systemic vascular responses to natural vestibular stimulation in the Mongolian gerbil.

Natural stimulation of the vestibular end organs will produce alterations in the local vestibular microvascular blood flow. The vestibular and cardiovascular systems require a coordinated interaction to maintain organ perfusion during rapid positional and postural changes. However, the detailed relationship of these systems is not well understood. There have been no previous descriptions of local vestibular blood flow (VBF) during natural stimulation (NS) conditions. In vivo VBF and systemic blood pressure (BP) in the Mongolian gerbil during natural stimulation. Using laser Doppler flowmetry, the authors obtained continuous measures of local VBF in both anesthetized and alert gerbils during sinusoidal rotational stimuli. Simultaneous recordings of systemic BP were collected from the contralateral common carotid artery. The anesthetized gerbils showed stable VBF and BP during all vestibular stimuli. By contrast, alert subjects demonstrated a significant response to natural stimulation. The VBF increased 28% over baseline, and systemic BP increased 8% during a 45-second, 0.133-Hz sinusoid. Decreases in BP of 8% and 5%, respectively, were seen with a 0.10 and 0.20 Hz, 360-second stimulus. A corresponding determination of VBF during the extended stimulus conditions was not technically possible. To the authors' knowledge, these are the first in vivo descriptions of vestibular blood flow during natural stimulation. In the alert animals, VBF increased in response to NS. This increase in flow does not appear to be directly dependent on systemic blood pressure changes and indicates that the vestibular microvasculature is closely regulated.

Vestibular Signals in the Parasolitary Nucleus

Vestibular primary afferents project to secondary vestibular neurons located in the vestibular complex. Vestibular primary afferents also project to the uvula-nodulus of the cerebellum where they terminate on granule cells. In this report we describe the physiological properties of neurons in a "new" vestibular nucleus, the parasolitary nucleus (Psol). This nucleus consists of 2,300 GABAergic neurons that project onto the ipsilateral inferior olive (beta-nucleus and dorsomedial cell column) as well as the nucleus reticularis gigantocellularis. These olivary neurons are the exclusive source of vestibularly modulated climbing fiber inputs to the cerebellum. We recorded the activity of Psol neurons during natural vestibular stimulation in anesthetized rabbits. The rabbits were placed in a three-axis rate table at the center of a large sphere, permitting vestibular and optokinetic stimulation. We recorded from 74 neurons in the Psol and from 23 neurons in the regions bordering Psol. The activity of 72/74 Psol neurons and 4/23 non-Psol neurons was modulated by vestibular stimulation in either the pitch or roll planes but not the horizontal plane. Psol neurons responded in phase with ipsilateral side-down head position or velocity during sinusoidal stimulation. Approximately 80% of the recorded Psol neurons responded to static roll-tilt. The optimal response planes of evoked vestibular responses were inferred from measurement of null planes. Optimal response planes usually were aligned with the anatomical orientation of one of the two ipsilateral vertical semicircular canals. The frequency dependence of null plane measurements indicated a convergence of vestibular information from otoliths and semicircular canals. None of the recorded neurons evinced optokinetic sensitivity. These results are consistent with the view that Psol neurons provide the vestibular signals to the inferior olive that eventually reached the cerebellum in the form of modulated climbing fiber discharges. These signals provide information about spatial orientation about the longitudinal axis.

Vestibular compensation and substitution.

This is a very brief update on the major papers since August 1998. Unilateral vestibular loss causes oculomotor, postural and sensory symptoms, all of which would be appropriate responses in a healthy person to a strong maintained angular and linear acceleration stimulus directed towards the healthy side. Within hours or days these static symptoms (so called because they are present without any externally imposed vestibular stimulation) reduce, and their progressive disappearance is called 'vestibular compensation'. However, careful testing with natural vestibular stimuli shows that the dynamic vestibular response after unilateral vestibular loss to passively imposed vestibular stimuli does not recover; it is usually asymmetric and functionally ineffective. Major recent developments are: (1) the permanent asymmetrical and functionally ineffective dynamic rotational vestibulo-ocular reflex responses to passive natural vestibular stimulation after unilateral vestibular loss and canal blocks in human patients; (2) evidence for the substitution of other sensory input and responses during vestibular compensation; (3) perceptual testing using visual perception of a horizontal line to confirm permanent otolith dysfunction; (4) the clear and substantial differences in post-unilateral vestibular loss vestibulo-ocular reflex responses between passive and active head turning; and (5) new results in brainstem physiology explaining the disappearance of static symptoms.