Vestibular contribution to spatial encoding.

Determining the spatial relation between objects and our location in the surroundings is essential for survival. Vestibular inputs provide key information about the position and movement of our head in the three-dimensional space, contributing to spatial navigation. Yet, their role in encoding spatial localisation of environmental targets remains to be fully understood. We probed the accuracy and precision of healthy participants' representations of environmental space by measuring their ability to encode the spatial location of visual targets (Experiment 1). Participants were asked to detect a visual light and then walk towards it. Vestibular signalling was artificially disrupted using stochastic galvanic vestibular stimulation (sGVS) applied selectively during encoding targets' location. sGVS impaired the accuracy and precision of locating the environmental visual targets. Importantly, this effect was specific to the visual modality. The location of acoustic targets was not influenced by vestibular alterations (Experiment 2). Our findings indicate that the vestibular system plays a role in localising visual targets in the surrounding environment, suggesting a crucial functional interaction between vestibular and visual signals for the encoding of the spatial relationship between our body position and the surrounding objects.

The vestibular system modulates the contributions of head and torso to egocentric spatial judgements.

Egocentric representations allow us to describe the external world as experienced from an individual's bodily location. We recently developed a novel method of quantifying the weight given to different body parts in egocentric judgments (the Misalignment Paradigm). We found that both head and torso contribute to simple alter-egocentric spatial judgments. We hypothesised that artificial stimulation of the vestibular system would provide a head-related signal, which might affect the weighting given to the head in egocentric spatial judgments. Bipolar Galvanic Vestibular Stimulation (GVS) was applied during the Misalignment Paradigm. A Sham stimulation condition was also included to control for non-specific effects. Our data show that the weight given to the head was increased during left anodal and right cathodal GVS, compared to the opposite GVS polarity (right anodal and left cathodal GVS) and Sham stimulation. That is, the polarity of GVS, which preferentially activates vestibular areas in the right cerebral hemisphere, influenced the relative weightings of head and torso in egocentric spatial judgments.

Dissociating contributions of head and torso to spatial reference frames: The misalignment paradigm.

When we represent someone's view of a scene as egocentrically structured, where do we represent the origin of the reference frame? By analysing responses in a spatial perspective-taking task as a function of spatial location with respect to both head and torso, we isolated the respective contribution of each part to spatial judgments. Both the head and the torso contributed to judgements, though with greater contributions from the torso. A second experiment manipulating visual contrast of the torso showed that this does not reflect low-level differences in visual salience between body parts. Our results demonstrate that spatial perspective-taking relies on a weighted combination of reference frames centred on different parts of the body.

Systematic underestimation of human hand weight.

According to Newton's laws, the weight of a body part is equal to its mass times gravitational acceleration. Our experience of body part weight, however, is constructed by the central nervous system. No sensory receptors directly specify the weight of body parts, and the factors influencing perceived weight remain unknown. The perceived weight of held objects has been linked to sensations of the magnitude of central motor commands sent to the muscles, what Helmholtz called the effort of will and has subsequently been called the sense of effort1. The link between the sense of effort and the perceived weight of objects is shown by studies demonstrating that held weights feel heavier when muscles are weakened by fatigue1, anaesthesia2, and following brain damage3. Similar drive to muscles is required to counteract the force of gravity on the limbs themselves, though few studies have investigated the perceived weight of body parts4. Stroke patients with hemiplegia frequently comment that their limbs feel heavy5, an effect linked to fatigue in the affected limb6. Similarly, amputees commonly complain of the weight of prosthetic limbs7, despite these typically weighing less than actual limbs. Here we report that healthy adult humans systematically underestimate hand weight. We used a psychophysical matching task to measure the experience of hand weight, which was underestimated on average by 49.4%. We further found that experimental induction of hand fatigue causes a systematic increase in perceived hand weight. Our results demonstrate that humans fail to experience the full weight of their body.

Gravitational and retinal reference frames shape spatial memory.

When reproducing the remembered location of dots within a circle, judgments are biased toward the center of imaginary quadrants formed by imaginary vertical and horizontal axes. This effect may result from the heightened precision in the visual system for these orientations in a retinotopic reference frame, or alternately on the internal representation of gravity. We dissociated reference frames defined by the retina and by gravity by having participants locate dots from memory in a circle when their head was upright (aligned with gravity) versus tilted 30° to the left (misaligned with gravity). We mapped the structure of spatial prototypes in a data-driven way using a novel "imaging" procedure. We calculated the rotation of the prototype maps which maximized the similarity between postures, letting us quantify the contribution of each reference frame. Spatial categories are determined by a combination of reference frames, with clear contributions from both gravitational and retinal factors. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Spatial Sensory References for Vestibular Self-Motion Perception.

While navigating through the surroundings, we constantly rely on inertial vestibular signals for self-motion along with visual and acoustic spatial references from the environment. However, the interaction between inertial cues and environmental spatial references is not yet fully understood. Here we investigated whether vestibular self-motion sensitivity is influenced by sensory spatial references. Healthy participants were administered a Vestibular Self-Motion Detection Task in which they were asked to detect vestibular self-motion sensations induced by low-intensity Galvanic Vestibular Stimulation. Participants performed this detection task with or without an external visual or acoustic spatial reference placed directly in front of them. We computed the d prime ( d ' ) as a measure of participants' vestibular sensitivity and the criterion as an index of their response bias. Results showed that the visual spatial reference increased sensitivity to detect vestibular self-motion. Conversely, the acoustic spatial reference did not influence self-motion sensitivity. Both visual and auditory spatial references did not cause changes in response bias. Environmental visual spatial references provide relevant information to enhance our ability to perceive inertial self-motion cues, suggesting a specific interaction between visual and vestibular systems in self-motion perception.

Pharmacological and non-pharmacological countermeasures to Space Motion Sickness: a systematic review.

Introduction: Space Motion Sickness (SMS) is a syndrome that affects around 70% of astronauts and includes symptoms of nausea, dizziness, fatigue, vertigo, headaches, vomiting, and cold sweating. Consequences range from discomfort to severe sensorimotor and cognitive incapacitation, which might cause potential problems for mission-critical tasks and astronauts and cosmonauts' well-being. Both pharmacological and non-pharmacological countermeasures have been proposed to mitigate SMS. However, their effectiveness has not been systematically evaluated. Here we present the first systematic review of published peer-reviewed research on the effectiveness of pharmacological and non-pharmacological countermeasures to SMS. Methods: We performed a double-blind title and abstract screening using the online Rayyan collaboration tool for systematic reviews, followed by a full-text screening. Eventually, only 23 peer-reviewed studies underwent data extraction. Results: Both pharmacological and non-pharmacological countermeasures can help mitigate SMS symptoms. Discussion: No definitive recommendation can be given regarding the superiority of any particular countermeasure approach. Importantly, there is considerable heterogeneity in the published research methods, lack of a standardized assessment approach, and small sample sizes. To allow for consistent comparisons between SMS countermeasures in the future, standardized testing protocols for spaceflight and ground-based analogs are needed. We believe that the data should be made openly available, given the uniqueness of the environment in which it is collected. Systematic review registration: https://www.crd.york.ac.uk/prospero/display_record.php?ID=CRD42021244131.

Vestibular modulation of somatosensory perception.

Functional imaging studies show that vestibular and somatosensory projections overlap in the human brain. However, it remains unclear whether and how vestibular inputs affect somatosensory function. To address this issue, we studied the effects of left caloric vestibular stimulation (CVS) on detection of near-threshold somatosensory stimuli delivered to the left and right hands of healthy volunteers. To investigate whether these effects were somatosensory specific, or supramodal, we also tested CVS modulation of visual contrast detection. Signal detection analyses showed increased somatosensory perceptual sensitivity immediately after CVS, both ipsilaterally and contralaterally. No statistically reliable effects on visual contrast sensitivity were found. These findings suggest that vestibular stimulation has a specific facilitatory effect on somatosensory detection, distinct from non-specific arousal and spatial attentional effects of CVS. Thus, the overlap in brain activations for vestibular and somatosensory inputs is not simply an anatomical curiosity, but may reflect a functional cross-modal perceptual interaction.

Reducing Cybersickness in 360-Degree Virtual Reality.

Despite the technological advancements in Virtual Reality (VR), users are constantly combating feelings of nausea and disorientation, the so-called cybersickness. Cybersickness symptoms cause severe discomfort and hinder the immersive VR experience. Here we investigated cybersickness in 360-degree head-mounted display VR. In traditional 360-degree VR experiences, translational movement in the real world is not reflected in the virtual world, and therefore self-motion information is not corroborated by matching visual and vestibular cues, which may trigger symptoms of cybersickness. We evaluated whether a new Artificial Intelligence (AI) software designed to supplement the 360-degree VR experience with artificial six-degrees-of-freedom motion may reduce cybersickness. Explicit (simulator sickness questionnaire and Fast Motion Sickness (FMS) rating) and implicit (heart rate) measurements were used to evaluate cybersickness symptoms during and after 360-degree VR exposure. Simulator sickness scores showed a significant reduction in feelings of nausea during the AI-supplemented six-degrees-of-freedom motion VR compared to traditional 360-degree VR. However, six-degrees-of-freedom motion VR did not reduce oculomotor or disorientation measures of sickness. No changes were observed in FMS and heart rate measures. Improving the congruency between visual and vestibular cues in 360-degree VR, as provided by the AI-supplemented six-degrees-of-freedom motion system considered, is essential for a more engaging, immersive and safe VR experience, which is critical for educational, cultural and entertainment applications.

Shared contributions of the head and torso to spatial reference frames across spatial judgments.

Egocentric frames of reference take the body as the point of origin of a spatial coordinate system. Bodies, however, are not points, but extended objects, with distinct parts that can move independently of one another. We recently developed a novel paradigm to probe the use of different body parts in simple spatial judgments, what we called the misalignment paradigm. In this study, we applied the misalignment paradigm in a perspective-taking task to investigate whether the weightings given to different body parts are shared across different spatial judgments involving different spatial axes. Participants saw birds-eye images of a person with their head rotated 45° relative to the torso. On each trial, a ball appeared and participants made judgments either of whether the ball was to the person's left or right, or whether the ball was in front of the person or behind them. By analysing the pattern of responses with respect to both head and torso, we quantified the contribution of each body part to the reference frames underlying each judgment. For both judgment types we found clear contributions of both head and torso, with more weight being given on average to the torso. Individual differences in the use of the two body parts were correlated across judgment types indicating the use of a shared set of weightings used across spatial axes and judgments. Moreover, retesting of participants several months later showed high stability of these weightings, suggesting that they are stable characteristics of people.

Distortions of perceived volume and length of body parts.

We experience our body as a 3D, volumetric object in the world. Measures of our conscious body image, in contrast, have investigated the perception of body size along one or two dimensions at a time. There is, thus, a discrepancy between existing methods for measuring body image and our subjective experience of having 3D body. Here we assessed in a sample of healthy adults the perception of body size in terms of its 1D length and 3D volume. Participants were randomly assigned to two groups using different measuring units (other body part and non-body object). They estimated how many units would fit in a perceived size of body segments and the whole body. The patterns of length and volume misperception across judged segments were determined as their perceived size proportional to their actual size. The pattern of volume misperception paints the representation of 3D body proportions resembling those of a somatosensory homunculus. The body parts with a smaller actual surface area relative to their volume were underestimated more. There was a tendency for body parts underestimated in volume to be overestimated in length. Perceived body proportions thus changed as a function of judgement type while showing a similarity in magnitude of the absolute estimation error, be it an underestimation of volume or overestimation of length. The main contribution of this study is assessing the body image as a 3D body representation, and thus extending beyond the conventional 'allocentric' focus to include the body on the inside. Our findings highlight the value of studying the perceptual distortions "at the baseline", i.e., in healthy population, so as to advance the understanding of the nature of perceptual distortions in clinical conditions.

Disentangling the visual, motor and representational effects of vestibular input.

The body midline provides a basic reference for egocentric representation of external space. Clinical observations have suggested that vestibular information underpins egocentric representations. Here we aimed to clarify whether and how vestibular inputs contribute to egocentric representation in healthy volunteers. In a psychophysical task, participants were asked to judge whether visual stimuli were located to the left or to the right of their body midline. Artificial vestibular stimulation was applied to stimulate the vestibular organs. We found that artificial stimulation of the vestibular system biased body midline perception. Importantly, no effect was found on motor effector selection. We also ruled out additional explanations based on allocentric visual representations and on potential indirect effects caused by vestibular-driven movements of the eyes, head and body. Taken together our data suggest that vestibular information contributes to computation of egocentric representations by affecting the internal representation of the body midline.

Cortical inhibitory function in cervical dystonia.

OBJECTIVE: To assess the specificity of cortical inhibitory deficits in cervical dystonia patients. METHODS: A systematic test battery was developed to assess spatial and temporal aspects of cortical inhibition, in motor and somatosensory systems of the hand. We tested 17 cervical dystonia (CD) patients and 19 controls assessing somatosensory spatial inhibition (grating orientation test, interdigital feedforward subliminal inhibition), somatosensory temporal inhibition (temporal discrimination threshold, feedforward subliminal inhibition), motor spatial inhibition (surround inhibition), and motor temporal inhibition (short interval intracortical inhibition). RESULTS: A significant deficit in CD was observed in both measures of somatosensory spatial inhibition, with a trend in the same direction in our measure of motor spatial inhibition. We found no significant group differences in temporal inhibition measures. Importantly, statistical comparison of effect sizes across the different measures showed that deficits in tests of spatial inhibition were greater than those in tests of temporal inhibition. CONCLUSION: Our results suggest that CD is associated with abnormal function of local inhibitory cortical circuits subserving spatial sensory processing. Importantly, this abnormality relates to the somatotopic representation of an unaffected body part. SIGNIFICANCE: These results clarify the nature of deficits in cortical inhibitory function in dystonia.

Feedforward somatosensory inhibition is normal in cervical dystonia.

BACKGROUND: Insufficient cortical inhibition is a key pathophysiological finding in dystonia. Subliminal sensory stimuli were reported to transiently inhibit somatosensory processing. Here we investigated whether such subliminal feedforward inhibition is reduced in patients with cervical dystonia. METHODS: Sixteen cervical dystonia patients and 16 matched healthy controls performed a somatosensory detection task. We measured the drop in sensitivity to detect a threshold-level digital nerve shock when it was preceded by a subliminal conditioning shock, compared to when it was not. RESULTS: Subliminal conditioning shocks reduced sensitivity to threshold stimuli to a similar extent in both patients and controls, suggesting that somatosensory subliminal feedforward inhibition is normal in cervical dystonia. CONCLUSION: Somatosensory feedforward inhibition was normal in this group of cervical dystonia patients. Our results qualify previous concepts of a general dystonic deficit in sensorimotor inhibitory processing.

Galvanic vestibular stimulation increases novelty in free selection of manual actions.

Making optimal choices in changing environments implies the ability to balance routine, exploitative patterns of behavior with novel, exploratory ones. We investigated whether galvanic vestibular stimulation (GVS) interferes with the balance between exploratory and exploitative behaviors in a free action selection task. Brief right-anodal and left-cathodal GVS or left-anodal and right-cathodal GVS were delivered at random to activate sensorimotor circuits in the left and right hemisphere, respectively. A sham stimulation condition was included. Participants endogenously generated sequences of possible actions, by freely choosing successive movements of the index or middle finger of the left or right hand. Left-anodal and right-cathodal GVS, which preferentially activates the vestibular projections in the right cerebral hemisphere, increased the novelty in action sequences, as measured by the number of runs in the sequences. In contrast, right-anodal and left-cathodal GVS decreased the number of runs. There was no evidence of GVS-induced spatial bias in action choices. Our results confirm previous reports showing a polarity-dependent effect of GVS on the balance between novel and routine responses, and thus between exploratory and exploitative behaviors.

The balance of feelings: vestibular modulation of bodily sensations.

The vestibular system processes information about head movement and orientation. No unimodal vestibular cortex has been identified in the mammalian brain. Rather, vestibular inputs are combined with many other sensory signals in the cortex. This arrangement suggests that vestibular input could influence processing in other sensory modalities. Here we show that vestibular stimulation differentially modulates two submodalities of the somatosensory system, increasing sensitivity to tactile input, and independently reducing sensitivity to nociceptive input. These modulations of touch and pain can clearly be distinguished from supramodal attentional effects of vestibular stimulation, because they are bilateral and operate in different directions. Outside the artificial conditions of laboratory stimulation, the vestibular system codes movements of the head, indicating a new relation between the body and the external world. We suggest the vestibular system participates in a form of sensory signal management, changing the balance between the various sensory systems as the relation between the body and the external environment changes. This sensory rebalancing may be a crucial element in the brain's capacity to reorient towards novel or salient features in the environment.

Vestibular modulation of spatial perception.

Vestibular inputs make a key contribution to the sense of one's own spatial location. While the effects of vestibular stimulation on visuo-spatial processing in neurological patients have been extensively described, the normal contribution of vestibular inputs to spatial perception remains unclear. To address this issue, we used a line bisection task to investigate the effects of galvanic vestibular stimulation (GVS) on spatial perception, and on the transition between near and far space. Brief left-anodal and right-cathodal GVS or right-anodal and left-cathodal GVS were delivered. A sham stimulation condition was also included. Participants bisected lines of different lengths at six distances from the body using a laser pointer. Consistent with previous results, our data showed an overall shift in the bisection bias from left to right as viewing distance increased. This pattern suggests leftward bias in near space, and rightward bias in far space. GVS induced strong polarity dependent effects in spatial perception, broadly consistent with those previously reported in patients: left-anodal and right-cathodal GVS induced a leftward bisection bias, while right-anodal and left-cathodal GVS reversed this effect, and produced bisection bias toward the right side of the space. Interestingly, the effects of GVS were comparable in near and far space. We speculate that vestibular-induced biases in space perception may optimize gathering of information from different parts of the environment.

How the vestibular system interacts with somatosensory perception: a sham-controlled study with galvanic vestibular stimulation.

The vestibular system has widespread interactions with other sensory modalities. Here we investigate whether vestibular stimulation modulates somatosensory function, by assessing the ability to detect faint tactile stimuli to the fingertips of the left and right hand with or without galvanic vestibular stimulation (GVS). We found that left anodal and right cathodal GVS, significantly enhanced sensitivity to mild shocks on either hand, without affecting response bias. There was no such effect with either right anodal and left cathodal GVS or sham stimulation. Further, the enhancement of somatosensory sensitivity following GVS does not strongly depend on the duration of GVS, or the interval between GVS and tactile stimulation. Vestibular inputs reach the somatosensory cortex, increasing the sensitivity of perceptual circuitry.

An anatomical account of somatoparaphrenia.

Somatoparaphrenia is a delusional belief whereby a patient feels that a paralyzed limb does not belong to his body; the symptom is typically associated with unilateral neglect and most frequently with anosognosia for hemiplegia. This association of symptoms makes anatomical inference based on single case studies not sufficiently specific. On the other hand, the only three anatomical group studies on somatoparaphrenia are contradictory: the right posterior insula, the supramarginal gyrus and the posterior corona radiata, or the right medial or orbito-frontal regions were all proposed as specific lesional correlates. We compared 11 patients with and 11 without somatoparaphrenia matched for the presence and severity of other associated symptoms (neglect, motor deficits and anosognosia). To take into account the frequent association of SP and neglect and hemiplegia, patients with and without somatoparaphrenia were also compared with a group of fifteen right brain damage patients without neglect and hemiplegia. We found a lesion pattern involving a fronto-temporo-parietal network typically associated with spatial neglect, hemiplegia and anosognosia. Somatoparaphrenic patients showed an additional lesion pattern primarily involving white matter and subcortical grey structures (thalamus, basal ganglia and amygdala). Further cortical damage was present in the middle and inferior frontal gyrus, postcentral gyrus and hippocampus. We propose that somatoparaphrenia occurs providing that a distributed cortical lesion pattern is present together with a subcortical lesion load that prevents most sensory input from being processed in neocortical structures; involvement of deep cortical and subcortical grey structures of the temporal lobe may contribute to reduce the sense of familiarity experienced by somatoparaphrenic patients for their paralyzed limb.