Watching the Effects of Gravity. Vestibular Cortex and the Neural Representation of “Visual” Gravity

Gravity is a physical constraint all terrestrial species have adapted to through evolution. Indeed, gravity effects are taken into account in many forms of interaction with the environment, from the seemingly simple task of maintaining balance to the complex motor skills performed by athletes and dancers. Graviceptors, primarily located in the vestibular otolith organs, feed the Central Nervous System with information related to the gravity acceleration vector. This information is integrated with signals from semicircular canals, vision, and proprioception in an ensemble of interconnected brain areas, including the vestibular nuclei, cerebellum, thalamus, insula, retroinsula, parietal operculum, and temporo-parietal junction, in the so-called vestibular network. Classical views consider this stage of multisensory integration as instrumental to sort out conflicting and/or ambiguous information from the incoming sensory signals. However, there is compelling evidence that it also contributes to an internal representation of gravity effects based on prior experience with the environment. This a priori knowledge could be engaged by various types of information, including sensory signals like the visual ones, which lack a direct correspondence with physical gravity. Indeed, the retinal accelerations elicited by gravitational motion in a visual scene are not invariant, but scale with viewing distance. Moreover, the “visual” gravity vector may not be aligned with physical gravity, as when we watch a scene on a tilted monitor or in weightlessness. This review will discuss experimental evidence from behavioral, neuroimaging (connectomics, fMRI, TMS), and patients’ studies, supporting the idea that the internal model estimating the effects of gravity on visual objects is constructed by transforming the vestibular estimates of physical gravity, which are computed in the brainstem and cerebellum, into internalized estimates of virtual gravity, stored in the vestibular cortex. The integration of the internal model of gravity with visual and non-visual signals would take place at multiple levels in the cortex and might involve recurrent connections between early visual areas engaged in the analysis of spatio-temporal features of the visual stimuli and higher visual areas in temporo-parietal-insular regions.

Mental imagery of object motion in weightlessness

Mental imagery represents a potential countermeasure for sensorimotor and cognitive dysfunctions due to spaceflight. It might help train people to deal with conditions unique to spaceflight. Thus, dynamic interactions with the inertial motion of weightless objects are only experienced in weightlessness but can be simulated on Earth using mental imagery. Such training might overcome the problem of calibrating fine-grained hand forces and estimating the spatiotemporal parameters of the resulting object motion. Here, a group of astronauts grasped an imaginary ball, threw it against the ceiling or the front wall, and caught it after the bounce, during pre-flight, in-flight, and post-flight experiments. They varied the throwing speed across trials and imagined that the ball moved under Earth’s gravity or weightlessness. We found that the astronauts were able to reproduce qualitative differences between inertial and gravitational motion already on ground, and further adapted their behavior during spaceflight. Thus, they adjusted the throwing speed and the catching time, equivalent to the duration of virtual ball motion, as a function of the imaginary 0 g condition versus the imaginary 1 g condition. Arm kinematics of the frontal throws further revealed a differential processing of imagined gravity level in terms of the spatial features of the arm and virtual ball trajectories. We suggest that protocols of this kind may facilitate sensorimotor adaptation and help tuning vestibular plasticity in-flight, since mental imagery of gravitational motion is known to engage the vestibular cortex.

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.

CENTRAL VERTIGO

Central vertigo is a symptom characterized by a feeling of changes in body position or environment as a result of diseases originating from the central nervous system. Central vertigo is caused by a disease that extend from vestibular nuclei in medulla oblongata to ocular motor nuclei and integration system in mesencephalon to vestibulocerebellum, thalamus and vestibular cortex in temporoparietal and the neuronal pathway which mediate VOR (vestibulo-ocular reflex). The diseases can be vestibular migrain, TIA (Transient Ischemic Attack), Vertebrobasilar ischemic stroke, multiple sclerosis, tumor in cerebelopontine angle and congenital malformation like Dandy Walker Syndrome. Central vertigo can be diagnosed by performing several special tests. This examination can also distinguish central vertigo from its differential diagnosis, namely peripheral vertigo. Management of central vertigo can be in the form of acute attack management and specific management according to the cause.

Vestibular Stimulation May Drive Multisensory Processing: Principles for Targeted Sensorimotor Therapy (TSMT)

At birth, the vestibular system is fully mature, whilst higher order sensory processing is yet to develop in the full-term neonate. The current paper lays out a theoretical framework to account for the role vestibular stimulation may have driving multisensory and sensorimotor integration. Accordingly, vestibular stimulation, by activating the parieto-insular vestibular cortex, and/or the posterior parietal cortex may provide the cortical input for multisensory neurons in the superior colliculus that is needed for multisensory processing. Furthermore, we propose that motor development, by inducing change of reference frames, may shape the receptive field of multisensory neurons. This, by leading to lack of spatial contingency between formally contingent stimuli, may cause degradation of prior motor responses. Additionally, we offer a testable hypothesis explaining the beneficial effect of sensory integration therapies regarding attentional processes. Key concepts of a sensorimotor integration therapy (e.g., targeted sensorimotor therapy (TSMT)) are also put into a neurological context. TSMT utilizes specific tools and instruments. It is administered in 8-weeks long successive treatment regimens, each gradually increasing vestibular and postural stimulation, so sensory-motor integration is facilitated, and muscle strength is increased. Empirically TSMT is indicated for various diseases. Theoretical foundations of this sensorimotor therapy are discussed.

The human vestibular cortex: functional anatomy, connectivity and the effect of vestibular disease

Area OP2 in the posterior peri-sylvian cortex has been proposed to be the core human vestibular cortex. We defined the functional anatomy of OP2 using spatially constrained independent component analysis of functional MRI data from the Human Connectome Project. Ten distinct subregions were identified. Most subregions showed significant connectivity to other areas with vestibular function: the parietal opercula, the primary somatosensory cortex, the supracalcarine cortex, the left inferior parietal lobule and the anterior cingulate cortex. OP2 responses to vestibular and visual-motion were analysed in 17 controls and 17 right-sided unilateral vestibular lesion patients (vestibular neuritis) who had previously undergone caloric and optokinetic stimulation during functional MRI. In controls, a posterior part of right OP2 showed: (a) direction-selective responses to visual motion; and (b) activation during caloric stimulation that correlated positively with perceived self-motion, and negatively with visual dependence. Patients showed abnormal OP2 activity, with an absence of visual or caloric activation of the healthy ear and no correlations with dizziness or visual dependence despite normal brainstem responses to caloric stimulation (slow-phase nystagmus velocity). A lateral part of right OP2 showed activity that correlated with chronic dizziness (situational vertigo) in patients. Our results define the functional anatomy of OP2 in health and disease. A posterior subregion of right OP2 shows strong functional connectivity to other vestibular regions and a visuo-vestibular profile that becomes profoundly disrupted after vestibular disease. In vestibular patients, a lateral subregion of right OP2 shows responses linked to the challenging long-term symptoms which define poorer clinical outcomes.

Altered Gray Matter Volume and Functional Connectivity in Patients With Vestibular Migraine

SubjectsVestibular migraine (VM) is the most common neurological cause of vertigo in adults. Previous neuroimaging studies have reported structural alterations in areas associated with pain and vestibular processing. However, it is unclear whether altered resting-state functional connectivity (FC) exists in brain regions with structural abnormalities in patients with VM.MethodsResting-state functional magnetic resonance imaging (MRI) and three-dimensional T1-weighed MRI were performed in 30 patients with VM and 30 healthy controls (HCs). Patients underwent an evaluation of migraine and dizziness severity. FC and voxel-based morphometry (VBM) were performed using DPABI 4.3 and CAT12, respectively. The association between changes in gray matter (GM) volume or FC and clinical parameters was also analyzed.ResultsCompared with HCs, patients with VM demonstrated a reduced GM volume in the bilateral parietoinsular vestibular cortex (PIVC), right middle frontal gyrus, and precuneus. The GM volume of the left PIVC was negatively associated with Dizziness Handicap Inventory score in patients with VM. Taking this region as a seed region, we further observed increased FC between the left primary somatosensory cortex (S1)/inferior parietal lobule (IPL) and the left PIVC in patients with VM.ConclusionFC between regions with a decline in GM volume (the PIVC and S1/IPL) is altered in patients with VM, suggesting that abnormalities in vestibular cortical network could be useful for understanding the underlying mechanisms of VM.

Vestibular bone thickness of the mandible in relation to the mandibular canal in a population from Bosnia and Herzegovina

Introduction: Dental implantology is the branch of dentistry that is gaining greater significance because a larger number of patients come with requests of implant placements. During dental implant placements, with patients with whom operation is carried out in the mandible, very frequently nervus alveolaris inferior can be injured. The nerve injury may occur during the implant placement, but the nerve may also be injured in case of harvesting of intraoral bone graft. During the bone graft harvesting, but also during any other procedure in the dentistry that entails working on vestibular side of corpus of the mandible, in order not to injure the nervus alveolaris inferior, it is important to familiarize oneself with the distance of the nerve from the outer vestibular cortex of the mandible. The objective of the study was to assess the vestibular bone thickness of the mandible in relation to the mandibular canal with the help of analysis of cone-beam computed tomography (CBCT) images.Methods: It was accessed the database of CBCT images taken at the School of Dental Medicine at the University of Sarajevo, where out of 700 reviewed CBCT images, an analysis of 322 CBCT images was conducted that satisfied inclusion criteria of the study. CBCT images were taken using of ORTHOPHOS SLX imaging unit. The measurement was conducted by Sidexis program on cross-section of CBCT image. The measurement of vestibular bone thickness was performed, by measuring the distance from the lateral wall of the mandibular canal to buccal mandibular compact bone, in the region of the second premolar, of the first and the second molar.Results: There were statistically significant differences in vestibular bone thickness between men and women on both sides in the region of the second premolar (p < 0.001) and first molar (p = 0.016 right, p = 0.018 left). T-test demonstrated no statistically significant difference in the vestibular bone thickens between men and women on either side in the case of vestibular bone thickness of the center of the second molar (p = 0.397 right, p = 0.743 left).Conclusion: Values of vestibular thickness of the mandible are larger with men than with women in all measuring points; however, statistically more significant differences between genders have been detected in the second premolar and center of the first molar.