Neurosensory development of the four brainstem-projecting sensory systems and their integration in the telencephalon.

Somatosensory, taste, vestibular, and auditory information is first processed in the brainstem. From the brainstem, the respective information is relayed to specific regions within the cortex, where these inputs are further processed and integrated with other sensory systems to provide a comprehensive sensory experience. We provide the organization, genetics, and various neuronal connections of four sensory systems: trigeminal, taste, vestibular, and auditory systems. The development of trigeminal fibers is comparable to many sensory systems, for they project mostly contralaterally from the brainstem or spinal cord to the telencephalon. Taste bud information is primarily projected ipsilaterally through the thalamus to reach the insula. The vestibular fibers develop bilateral connections that eventually reach multiple areas of the cortex to provide a complex map. The auditory fibers project in a tonotopic contour to the auditory cortex. The spatial and tonotopic organization of trigeminal and auditory neuron projections are distinct from the taste and vestibular systems. The individual sensory projections within the cortex provide multi-sensory integration in the telencephalon that depends on context-dependent tertiary connections to integrate other cortical sensory systems across the four modalities.

Collic evoked potentials, myogenic potentials (CEMPs) and postural responses produced by brief 100 Hz vibration of the sternocleidomastoid muscle.

We present an initial report using 5 subjects, of short and long latency collic evoked responses following a half cycle of 100 Hz vibration (5 ms) applied to the sternocleidomastoid (SCM) tendon. These were detected in EEG and extraocular and leg muscles and compared with vestibular-dependent responses from direct mastoid stimulation. The responses from the extraocular recording site are likely to be evoked myogenic potentials, thus "collic evoked myogenic potentials" (CEMPs). An n19/p24 presumed ocular CEMP (oCEMP) was followed by a P22/N28 response over the posterior fossa, referred to as a collic cerebellar evoked potential (CoCEP), with responses in leg muscles starting around 55 ms. In contrast to their vestibular analogues, the oCEMP and CoCEP were predominantly ipsilateral to the side of stimulation, consistent with a double-crossed projection. In addition, their thresholds were just above the threshold of vibrotactile sensation, implying a low threshold, oligo-synaptic projection of SCM afferents to both extraocular and cerebellar targets. Following these short latency responses, SCM tendon stimulation evoked prolonged EMG responses in postural muscles of the legs, consistent with a role in the afferent limb of a short latency, spino-bulbar-spinal postural response to sternal perturbations. These collic evoked responses are likely to be of value in understanding the functions of cervical muscle afferents and have clinical value, for example in monitoring compensation after vestibular loss.

[Cervial vestibular-evoked myogenic potential induced by galvanic vestibular stimulation in normal people].

Objective:To establish a new method for detecting vestibular function by testing cervical vestibular-evoked myogenic potential induced by galvanic vestibular stimulation in normal population. Method:Twenty normal ears were tested for cervical vestibular evoked myogenic potential induced by galvanic vestibular stimulation. SPSS 18.0 software was used to analyze the obtained data. Result:In all healthy subjects mastoid-forehead galvanic vestibular stimulation produced a positive-negative biphasic EMG responses on SCM ipsilateral to the cathodal electrode. The latency of p13 was（11.52±3.05） ms. The latency of n23 was（15.31±3.38） ms. The amplitude of p13-n23 was（40.55±27.93） µV. The interval of p13-n23 was（3.53±1.38） ms. The interaural asymmetry ratio（AR, %） of p13, n23 latency, the amplitude and interval were respectively（6.96±6.79）%, （6.47±5.93）%, （28.08±26.42）% and （16.61±11.11）%. There was no significant difference in all parameters between the right and left ears of all subjects. Conclusion:The value of cervical vestibular-evoked myogenic potential induced by galvanic vestibular stimulation in normal subjects can be established to explore methods for diagnosis, treatment and researching mechanism of auditory neuropathy and vestibular neuropathy.

The influence of vestibular stimulation on metabolism and body composition.

Obesity, diabetes and metabolic disease represent an ongoing and rapidly worsening public health issue in both the developed, and much of the developing world. Although there are many factors that influence fat storage, it has been clearly demonstrated that the homeostatic cornerstone of metabolism lies within the hypothalamus. Moreover, neuronal damage to vital areas of the hypothalamus can drive reregulation or dysregulation of endocrine function, energy expenditure and appetite, thereby promoting a shift in overall metabolic function towards a state of obesity. Therefore, identification of treatments that influence the hypothalamus to improve obesity and associated metabolic diseases has long been a medical goal. Interestingly, evidence from animal studies suggests that activating the vestibular system, specifically the macular gravity receptor, influences the hypothalamus in a way that decreases body fat storage and causes a metabolic shift towards a leaner state. Given that the macular element of the vestibular system has been shown to activate with transdermal electrical stimulation applied to the mastoids, this may be a potential therapeutic approach for obesity, diabetes or related metabolic diseases, whereby repetitive stimulation of the vestibular system influences hypothalamic control of metabolic homeostasis, thereby encouraging decreased fat storage. Here, we present an up-to-date review of the current literature surrounding the vestibular influence of the hypothalamus and associated homeostatic sites in the context of current and novel therapeutic approaches for improved clinical management of obesity and diabetes.

Vestibular Stimulation Modulates Neural Correlates of Own-body Mental Imagery.

There is growing evidence that vestibular information is not only involved in reflexive eye movements and the control of posture but it also plays an important role in higher order cognitive processes. Previous behavioral research has shown that concomitant vestibular stimuli influence performance in tasks that involve imagined self-rotations. These results suggest that imagined and perceived body rotations share common mechanisms. However, the nature and specificity of these effects remain largely unknown. Here, we investigated the neural mechanisms underlying this vestibulocognitive interaction. Participants (n = 20) solved an imagined self-rotation task during caloric vestibular stimulation. We found robust main effects of caloric vestibular stimulation in the core region of the vestibular network, including the rolandic operculum and insula bilaterally, and of the cognitive task in parietal and frontal regions. Interestingly, we found an interaction of stimulation and task in the left inferior parietal lobe, suggesting that this region represents the modulation of imagined body rotations by vestibular input. This result provides evidence that the inferior parietal lobe plays a crucial role in the neural integration of mental and physical body rotation.

Multimodal stimuli modulate rapid visual responses during reaching.

The reticulospinal tract plays an important role in primate upper limb function, but methods for assessing its activity are limited. One promising approach is to measure rapid visual responses (RVRs) in arm muscle activity during a visually cued reaching task; these may arise from a tecto-reticulospinal pathway. We investigated whether changes in reticulospinal excitability can be assessed noninvasively using RVRs, by pairing the visual stimuli of the reaching task with electrical stimulation of the median nerve, galvanic vestibular stimulation, or loud sounds, all of which are known to activate the reticular formation. Surface electromyogram (EMG) recordings were made from the right deltoid of healthy human subjects as they performed fast reaching movements toward visual targets. Stimuli were delivered up to 200 ms before target appearance, and RVR was quantified as the EMG amplitude in a window 75-125 ms after visual target onset. Median nerve, vestibular, and auditory stimuli all consistently facilitated the RVRs, as well as reducing the latency of responses. We propose that this facilitation reflects modulation of tecto-reticulospinal excitability, which is consistent with the idea that the amplitude of RVRs can be used to assess changes in brain stem excitability noninvasively in humans.NEW & NOTEWORTHY Short-latency responses in arm muscles evoked during a visually driven reaching task have previously been proposed to be tecto-reticulospinal in origin. We demonstrate that these responses can be facilitated by pairing the appearance of a visual target with stimuli that activate the reticular formation: median nerve, vestibular, and auditory stimuli. We propose that this reflects noninvasive measurement and modulation of reticulospinal excitability.

Deactivation of somatosensory and visual cortices during vestibular stimulation is associated with older age and poorer balance.

Aging is associated with peripheral and central declines in vestibular processing and postural control. Here we used functional MRI to investigate age differences in neural vestibular representations in response to pneumatic tap stimulation. We also measured the amount of body sway in multiple balance tasks outside of the MRI scanner to assess the relationship between individuals' balance ability and their vestibular neural response. We found a general pattern of activation in canonical vestibular cortex and deactivation in cross modal sensory regions in response to vestibular stimulation. We found that activation amplitude of the vestibular cortex was correlated with age, with younger individuals exhibiting higher activation. Deactivation of visual and somatosensory regions increased with age and was associated with poorer balance. The results demonstrate that brain activations and deactivations in response to vestibular stimuli are correlated with balance, and the pattern of these correlations varies with age. The findings also suggest that older adults exhibit less sensitivity to vestibular stimuli, and may compensate by differentially reweighting visual and somatosensory processes.

Unilateral vestibular deafferentation impairs embodied spatial cognition.

A growing number of studies indicate that cognitive complaints are common in patients with peripheral vestibular disorders. A better understanding of how vestibular disorders influence cognition in these patients requires a clear delineation of the cognitive domains affected by vestibular disorders. Here, we compared the consequences of left and right vestibular neurectomy on third-person perspective taking-a visuo-spatial task requiring mainly own-body mental imagery, and on 3D objects mental rotation imagery-requiring object-based mental imagery, but no perspective taking. Patients tested 1 week after a unilateral vestibular neurectomy and a group of age- and gender-matched healthy participants played a virtual ball-tossing game from their own first-person perspective (1PP) and from the perspective of a distant avatar (third-person perspective, 3PP). Results showed larger response times in the patients with respect to their controls for the 3PP taking task, but not for the 1PP task and the 3D objects mental imagery. In addition, we found that only patients with left vestibular neurectomy presented altered 3PP taking abilities when compared to their controls. This study suggests that unilateral vestibular loss affects mainly own-body mental transformation and that only left vestibular loss seems to impair this cognitive process. Our study also brings further evidence that vestibular signals contribute to the sensorimotor bases of social cognition and strengthens the connections between the so far distinct fields of social neuroscience and human vestibular physiology.

Objective methods to measure vestibular evoked myogenic potential response saccular tuning curves.

Objective: To detect cervical vestibular evoked myogenic potential (cVEMP) responses using objective statistical approaches and to apply this approach to estimate saccular frequency-tuning curves in volunteers and Ménière's disease (MD) patients. Design: Estimates of cVEMP threshold were carried out by 3 expert raters at 500 Hz and compared to objective threshold estimates (using Hotelling's T2 [HT2] and Fsp). Saccular tuning curves were objectively estimated. Study sample: Objective and subjective estimates of cVEMP response thresholds were compared for 13 normal hearing adults. Objective measurement of saccular tuning curves was explored in 20 healthy adults and 15 patients with MD. Results: Significant variability was seen between subjective estimates of cVEMP thresholds. Objective analysis with the HT2 test was more sensitive than 2 of 3 experts in detecting responses. The measurement time of cVEMP was considerably reduced with the HT2 test. Objective saccular tuning curves in volunteers showed strongest responses at 500 Hz. A flatter tuning curve was seen for MD patients. Conclusions: There is significant variability in subjective estimations of cVEMP thresholds. Objective analysis methods are more sensitive than subjective analysis, can detect responses rapidly and have potential to reduce variability in threshold estimates, hence they appear well suited to measure cVEMP tuning curves.

Superior Canal Dehiscence Syndrome: Relating Clinical Findings With Vestibular Neural Responses From a Guinea Pig Model.

HYPOTHESIS: In superior canal dehiscence (SCD), fluid displacement of the endolymph activates type I vestibular hair cells in the crista of the affected canal and thus irregular superior canal (SC) neurons in Scarpa's ganglion, which provides the neurophysiological basis for the clinical presentation of SCD. BACKGROUND: Patients with SCD display sound- and vibration-induced vertigo/nystagmus and increased amplitudes of vestibular evoked myogenic potentials. METHODS: Extracellular recordings from n = 25 primary vestibular neurons of 16 female guinea pigs were analyzed. We recorded from the same vestibular neuron before, during and after creating the dehiscence and after closing the dehiscence. Neurobiotin labeling was employed in n = 11 neurons. RESULTS: After SCD, previously unresponsive irregular SC neurons displayed a stimulus-locked increase in discharge during application of air-conducted sound (ACS) or bone-conducted vibration (BCV) for a broad range of frequencies (ACS: 200-4000 Hz; BCV: 500-1500 Hz). This typical response was only observed for irregular SC neurons (n = 19), but not regular SC neurons, or irregular/regular horizontal canal neurons (n = 2 each), and was abolished after closing the dehiscence. Eleven irregular SC neurons responsive to ACS and/or BCV were traced back to calyx synapses in the central crista of the affected superior canal by neurobiotin labeling. CONCLUSIONS: Stimulus-locked activation of irregular SC neurons by ACS and BCV is the neurophysiological basis for sound- and vibration-induced vertigo/nystagmus and increased VEMP amplitudes in SCD. The results of the present study help to improve vestibular diagnostics in patients with suspected SCD.

Sound-evoked vestibular projections to the splenius capitis in humans: comparison with the sternocleidomastoid muscle.

The short-latency vestibulo-collic reflex in humans is well defined for only the sternocleidomastoid (SCM) neck muscle. However, other neck muscles also receive input from the balance organs and participate in neck stabilization. We therefore investigated the sound-evoked vestibular projection to the splenius capitis (SC) muscles by comparing surface and single motor unit responses in the SC and SCM muscles in 10 normal volunteers. We also recorded surface responses in patients with unilateral vestibular loss but preserved hearing and hearing loss but preserved vestibular function. The single motor unit responses were predominantly inhibitory, and the strongest responses were recorded in the contralateral SC and ipsilateral SCM. In both cases there was a significant decrease or gap in single motor unit activity, in SC at 11.7 ms for 46/66 units and in SCM at 12.7 ms for 51/58 motor units. There were fewer significant responses in the ipsilateral SC and contralateral SCM muscles, and they consisted primarily of weak increases in activity. Surface responses recorded over the contralateral SC were positive-negative during neck rotation, similar to the ipsilateral cervical vestibular evoked myogenic potential in SCM. Responses in SC were present in the patients with hearing loss and absent in the patient with vestibular loss, confirming their vestibular origin. The results describe a pattern of inhibition consistent with the synergistic relationship between these muscles for axial head rotation, with the crossed vestibular projection to the contralateral SC being weaker than the ipsilateral projection to the SCM. NEW & NOTEWORTHY We used acoustic vestibular stimulation to investigate the saccular projections to the splenius capitis (SC) and sternocleidomastoid (SCM) muscles in humans. Single motor unit recordings from within the muscles demonstrated strong inhibitory projections to the contralateral SC and ipsilateral SCM muscles and weak excitatory projections to the opposite muscle pair. This synergistic pattern of activation is consistent with a role for the reflex in axial rotation of the head.

Postural control during galvanic vestibular stimulation in patients with persistent perceptual-postural dizziness.

Over the past years galvanic vestibular stimulation (GVS) has been increasingly applied to stimulate the vestibular system in health and disease, but not in patients with persistent postural-perceptual dizziness (PPPD) yet. We functionally tested motion perception thresholds and postural responses to imperceptible noisy (nGVS) and perceptible bimastoidal GVS intensities in patients with PPPD with normal vestibulo-ocular reflexes. We hypothesized that GVS destabilizes PPPD patients under simple postural conditions stronger compared to healthy controls. They were compared to healthy subjects under several conditions each with the eyes open and closed: baseline with firm platform support, standing on foam and cognitive demand (count backward). Low and high GVS intensities (range 0.8-2.8 mA) were applied according to the individual thresholds and compared with no GVS. PPPD patients showed a reduced perception threshold to GVS compared to healthy control subjects. Median postural sway speed increased with stimulus intensity and on eye closure, but there was no group difference, irrespective of the experimental condition. Romberg's ratio was consistently lower during nGVS than in all other conditions. Group-related dissociable effects were found with the eyes closed in (i) the baseline condition in which high GVS elicited higher postural sway of PPPD patients and (ii) in the foam condition, with better postural stability of PPPD patients during perceptible GVS. Group and condition differences of postural control were neither related to anxiety nor depression scores. GVS may be helpful to identify thresholds of vestibular perception and to modulate vestibulo-spinal reflexes in PPPD, with dissociable effects with respect to perceptible and imperceptible stimuli. The sway increase in the baseline of PPPD may be related to an earlier transition from open- to closed-loop mode of postural control. In contrast, the smaller sway of PPPD in the foam condition under visual deprivation is in line with the known balance improvement under more demanding postural challenges in PPPD. It is associated with a prolonged transition from open- to closed-loop postural feedback control. It could also reflect a shift of intersensory weighting with a smaller dependence on proprioceptive feedback control in PPPD patients under complex tasks. In summary, GVS discloses differences between simple and complex balance tasks in PPPD.

Thalamocortical network: a core structure for integrative multimodal vestibular functions.

PURPOSE OF REVIEW: To apply the concept of nonreflexive sensorimotor and cognitive vestibular functions and disturbances to the current view of separate right and left thalamocortical systems. RECENT FINDINGS: The neuronal modules for sensorimotor and cognitive functions are organized in so-called provincial hubs with intracommunity connections that interact task-dependently via connector hubs. Thalamic subnuclei may serve not only as provincial hubs but also in higher order nuclei as connector hubs. Thus, in addition to its function as a cortical relay station of sensory input, the human thalamus can be seen as an integrative hub for brain networks of higher multisensory vestibular function. Imaging studies on the functional connectivity have revealed a dominance of the right side in right-handers at the upper brainstem and thalamus. A connectivity-based parcellation study has confirmed the asymmetrical organization (i.e., cortical dominance) of the parieto-insular vestibular cortex, an area surrounded by other vestibular cortical areas with symmetrical (nondominant) organization. Notably, imaging techniques have shown that there are no crossings of the vestibular pathways in between the thalamic nuclei complexes. Central vestibular syndromes caused by lesions within the thalamocortical network rarely manifest with rotational vertigo. This can be explained and mathematically simulated by the specific coding of unilateral vestibular dysfunction within different cell systems, the angular velocity cell system (rotational vertigo in lower brainstem lesions) in contrast to the head direction cell system (directional disorientation and swaying vertigo in thalamocortical lesions). SUMMARY: The structural and functional separation of the two thalamic nuclei complexes allowed a lateralization of the right and left hemispheric functions to develop. Furthermore, it made possible the simultaneous performance of sensorimotor and cognitive tasks, which require different spatial reference systems in opposite hemispheres, for example, egocentric manipulation of objects (handedness) and allocentric orientation of the self in the environment by the multisensory vestibular system.

Structural connectome of the human vestibular, pre-motor, and navigation network.

The aim of this study is to characterize modules and hubs within the multimodal vestibular system and, particularly, to test the centrality of posterior peri-sylvian regions. Structural connectivity matrices from 50 unrelated healthy right-handed subjects from the Human Connectome Project (HCP) database were analyzed using multishell diffusion-weighted data, probabilistic tractography (constrained spherical-deconvolution informed filtering of tractograms) in combination with subject-specific grey matter parcellations. Network nodes included parcellated regions within the vestibular, pre-motor and navigation system. Module calculation produced two and three modules in the right and left hemisphere, respectively. On the right, regions were grouped into a vestibular and pre-motor module, and into a visual-navigation module. On the left this last module was split into an inferior and superior component. In the thalamus, a region comprising the mediodorsal and anterior complex, and lateral and inferior pulvinar, was included in the ipsilateral navigation module, while the remaining thalamus was clustered with the ipsilateral vestibular pre-motor module. Hubs were located bilaterally in regions encompassing the inferior parietal cortex and the precuneus. This analysis revealed a dorso-lateral path within the multi-modal vestibular system related to vestibular / motor control, and a ventro-medial path related to spatial orientation / navigation. Posterior peri-sylvian regions may represent the main hubs of the whole modular network.

Effect of noisy galvanic vestibular stimulation in community-dwelling elderly people: a randomised controlled trial.

BACKGROUND: Balance disorders are a risk factor for falls in the elderly. Although noisy galvanic vestibular stimulation (nGVS) has been reported to improve balance in young people, randomised control trials targeting community-dwelling elderly people have not been conducted to date. We aimed to assess the influence of nGVS on COP sway in the open-eye standing posture among community-dwelling elderly people in a randomised controlled trial. METHODS: A randomised controlled trial of 32 community-dwelling elderly people randomly assigned to control (sham stimulation) and an nGVS groups. All participants underwent centre of pressure (COP) sway measurements while standing with open eyes at baseline and during stimulation. The control group underwent sham stimulation and the nGVS group underwent noise stimulation (0.4 mA; 0.1-640 Hz). RESULTS: In the nGVS group, sway path length, mediolateral mean velocity and anteroposterior mean velocity decreased during stimulation compared with baseline (P < 0.01). The effect of nGVS was large in participants with a high COP sway path length at baseline, but there was no significant difference in COP sway in the control group. CONCLUSIONS: We conclude that nGVS decreases the COP sway path length and mean velocity of community-dwelling elderly people when standing with open eyes. This suggests that nGVS could be effective for treating balance dysfunction in the elderly.

Combined exposure to carbon disulfide and low-frequency noise reversibly affects vestibular function.

Chronic occupational exposure to carbon disulfide (CS2) has debilitating motor and sensory effects in humans, which can increase the risk of falls. Although no mention of vestibulotoxic effects is contained in the literature, epidemiological and experimental data suggest that CS2 could cause low-frequency hearing loss when associated with noise exposure. Low-frequency noise might also perturb the peripheral balance receptor through an as-yet unclear mechanism. Here, we studied how exposure to a low-frequency noise combined with 250-ppm CS2 affected balance in rats. Vestibular function was tested based on post-rotary nystagmus recorded by a video-oculography system. These measurements were completed by behavioral tests and analysis of the cerebellum to measure expression levels for gene expression associated with neurotoxicity. Assays were performed prior to and following a 4-week exposure, and again after a 4-week recovery period. Functional measurements were completed by histological analyses of the peripheral organs.Nystagmus was unaltered by exposure to noise alone, while CS2 alone caused a moderate 19% decrease of the saccade number. In contrast, coexposure to 250-ppm CS2 and low-frequency noise decreased both saccade number and duration by 33% and 34%, respectively. After four weeks, recovery was only partial but measures were not significantly different from pre-exposure values. Real-time quantitative polymerase chain reaction (RT-qPCR) analysis of cerebellar tissue revealed a slight but significant modification in expression levels for two genes linked to neurotoxicity in CS2-exposed animals. However, neither histopathological changes to the peripheral receptor nor behavioral differences were observed. Based on all these results, we propose that the effects of CS2 were due to reversible neurochemical disturbance of the efferent pathways managing post-rotatory nystagmus. Because the nervous structures involving the vestibular function appear particularly sensitive to CS2, post-rotary nystagmus could be used as an early, non-invasive measurement to diagnose CS2 intoxication as part of an occupational conservation program.

Exhibition of stochastic resonance in vestibular tilt motion perception.

BACKGROUND: Stochastic Resonance (SR) is a phenomenon broadly described as "noise benefit". The application of subsensory electrical Stochastic Vestibular Stimulation (SVS) via electrodes behind each ear has been used to improve human balance and gait, but its effect on motion perception thresholds has not been examined. OBJECTIVE: This study investigated the capability of subsensory SVS to reduce vestibular motion perception thresholds in a manner consistent with a characteristic bell-shaped SR curve. METHODS: We measured upright, head-centered, roll tilt Direction Recognition (DR) thresholds in the dark in 12 human subjects with the application of wideband 0-30 Hz SVS ranging from ±0-700 µA. To conservatively assess if SR was exhibited, we compared the proportions of both subjective and statistical SR exhibition in our experimental data to proportions of SR exhibition in multiple simulation cases with varying underlying SR behavior. Analysis included individual and group statistics. RESULTS: As there is not an established mathematical definition, three humans subjectively judged that SR was exhibited in 78% of subjects. "Statistically significant SR exhibition", which additionally required that a subject's DR threshold with SVS be significantly lower than baseline (no SVS), was present in 50% of subjects. Both percentages were higher than simulations suggested could occur simply by chance. For SR exhibitors, defined by subjective or statistically significant criteria, the mean DR threshold improved by -30% and -39%, respectively. The largest individual improvement was -47%. CONCLUSION: At least half of the subjects were better able to perceive passive body motion with the application of subsensory SVS. This study presents the first conclusive demonstration of SR in vestibular motion perception.

Multisensory vestibular, vestibular-auditory, and auditory network effects revealed by parametric sound pressure stimulation.

Multisensory convergence and sensorimotor integration are important aspects for the mediation of higher vestibular cognitive functions at the cortical level. In contrast to the integration of vestibulo-visual or vestibulo-tactile perception, much less is known about the neural mechanism that mediates the integration of vestibular-otolith (linear acceleration/translation/gravity detection) and auditory processing. Vestibular-otolith and auditory afferents can be simultaneously activated using loud sound pressure stimulation, which is routinely used for testing cervical and ocular vestibular evoked myogenic potentials (VEMPs) in clinical neurotological testing. Due to the simultaneous activation of afferents there is always an auditory confound problem in fMRI studies of the neural topology of these systems. Here, we demonstrate that the auditory confounding problem can be overcome in a novel way that does not require the assumption of simple subtraction and additionally allows detection of non-linear changes in the response due to vestibular-otolith interference. We used a parametric sound pressure stimulation design that took each subject's vestibular stimulation threshold into account and analyzed for changes in BOLD-response below and above vestibular-otolith threshold. This approach helped to investigate the functional neuroanatomy of sound-induced auditory and vestibular integration using functional magnetic resonance imaging (fMRI). Results revealed that auditory and vestibular convergence are contained in overlapping regions of the caudal part of the superior temporal gyrus (STG) and the posterior insula. In addition, there are regions that were responsive only to suprathreshold stimulations, suggesting vestibular (otolith) signal processing in these areas. Based on these parametric analyses, we suggest that the caudal part of the STG and posterior insula could contain areas of vestibular contribution to auditory processing, i.e., higher vestibular cortices that provide multisensory integration that is important for tasks such as spatial localization of sound.

Psychophysical evidence for auditory motion parallax.

Distance is important: From an ecological perspective, knowledge about the distance to either prey or predator is vital. However, the distance of an unknown sound source is particularly difficult to assess, especially in anechoic environments. In vision, changes in perspective resulting from observer motion produce a reliable, consistent, and unambiguous impression of depth known as motion parallax. Here we demonstrate with formal psychophysics that humans can exploit auditory motion parallax, i.e., the change in the dynamic binaural cues elicited by self-motion, to assess the relative depths of two sound sources. Our data show that sensitivity to relative depth is best when subjects move actively; performance deteriorates when subjects are moved by a motion platform or when the sound sources themselves move. This is true even though the dynamic binaural cues elicited by these three types of motion are identical. Our data demonstrate a perceptual strategy to segregate intermittent sound sources in depth and highlight the tight interaction between self-motion and binaural processing that allows assessment of the spatial layout of complex acoustic scenes.

Stochastic resonance in the human vestibular system - Noise-induced facilitation of vestibulospinal reflexes.

BACKGROUND: There is strong evidence that the presence of noise can enhance information processing in sensory systems via stochastic resonance (SR). OBJECTIVES: To examine the presence of SR in human vestibulospinal reflex function. METHODS: Healthy subjects were stimulated with 1 Hz sinusoidal GVS of varying amplitudes (0-1.9 mA). Coherence between GVS input and stimulation-induced motion responses was determined and psychometric function fits were subsequently used to determine individual vestibulospinal reflex thresholds. This procedure was repeated with additional application of imperceptible white noise GVS (nGVS). RESULTS: nGVS significantly facilitated the detectability of weak subthreshold vestibular inputs (p < 0.001) and thereby effectively lowered the vestibulospinal threshold in 90% of participants (p < 0.001, mean reduction: 17.5 ± 14.6%). CONCLUSION: This finding provides evidence for the presence of SR-dynamics in the human vestibular system and gives a functional explanation for previously observed ameliorating effects of low-intensity vestibular noise stimulation on balance control in healthy subjects and patients with vestibular hypofunction.