Hemispheric competence for auditory spatial representation

Sound localization relies on the analysis of interaural time and intensity differences, as well as attenuation patterns by the outer ear. We investigated the relative contributions of interaural time and intensity difference cues to sound localization by testing 60 healthy subjects: 25 with focal left and 25 with focal right hemispheric brain damage. Group and single-case behavioural analyses, as well as anatomo-clinical correlations, confirmed that deficits were more frequent and much more severe after right than left hemispheric lesions and for the processing of interaural time than intensity difference cues. For spatial processing based on interaural time difference cues, different error types were evident in the individual data. Deficits in discriminating between neighbouring positions occurred in both hemispaces after focal right hemispheric brain damage, but were restricted to the contralesional hemispace after focal left hemispheric brain damage. Alloacusis (perceptual shifts across the midline) occurred only after focal right hemispheric brain damage and was associated with minor or severe deficits in position discrimination. During spatial processing based on interaural intensity cues, deficits were less severe in the right hemispheric brain damage than left hemispheric brain damage group and no alloacusis occurred. These results, matched to anatomical data, suggest the existence of a binaural sound localization system predominantly based on interaural time difference cues and primarily supported by the right hemisphere. More generally, our data suggest that two distinct mechanisms contribute to: (i) the precise computation of spatial coordinates allowing spatial comparison within the contralateral hemispace for the left hemisphere and the whole space for the right hemisphere; and (ii) the building up of global auditory spatial representations in right temporo-parietal cortice

Investigating the interplay of the human attentional and vestibular systems using transcranial magnetic stimulation

The aim of this doctoral thesis was to investigate the relationship between the processing of vestibular information, on the one hand, and higher cognitive functions such as visual (spatial) attention and perceptual decision making, on the other. In order to draw causal inference about the role of specific cortical regions in this interplay, two experimental studies were conducted which combined psychophysical task designs using verticality judgment tasks with transcranial magnetic stimulation (TMS). The first study employed a simultaneous TMS-EEG approach to examine the role of the right intraparietal sulcus (IPS) within the dorsal parietal cortex in verticality judgments – a cortical area that has repeatedly been associated with both the visual attention and vestibular systems. Facilitatory effects of right IPS TMS on the bias of verticality perception were reported and mirrored by EEG results, which pointed to a normalization of individual perceptual biases reflected in a fronto-central ERP component following the stimulation. In contrast, no effects of left IPS TMS on either behavioural or electrophysiological measures were observed and right IPS TMS did not modulate performance in a control task that used the same set of stimuli (vertical Landmark task). These findings point to a causal role of the right IPS in the neuronal implementation of upright perception and strengthen the notion of vestibular-attentional coupling. In the second study verticality judgments had to be made under different levels of perceptual demand to address the question of how perceptual decision making interacts with vestibular processing. Stimuli adapted from those used in the first study were presented in a visual search setting, which required perceptual and response switches, in a way that varied attentional demands. This task was combined with offline theta-burst TMS applied to the dorsal medial frontal cortex (dMFC). The dMFC has been found to crucially contribute to perceptual decision making and is connected to core parts of the vestibular cortical network. Analysis of distinct features of behavioural performance before as compared to following dMFC TMS revealed a specific involvement of the dMFC in establishing the precision and accuracy of verticality judgments, particularly under conditions of high perceptual load. In summary, the results of the two studies support the idea of a functional link between the processing of vestibular information, (spatial) attention, and perceptual decision making, giving rise to higher vestibular cognition. Moreover, they suggest that on a cortical level this interplay is achieved within a network of multimodal processing regions such as the parietal and frontal cortices

Distinct neural networks underlie encoding of categorical versus coordinate spatial relations during active navigation

It has been proposed that spatial relations are encoded either categorically, such that the relative positions of objects are defined in prepositional terms; or in terms of visual coordinates, such that the precise distances between objects are represented. In humans, it has been assumed that a left hemisphere neural network sub-serves categorical representations, and that coordinate representations are right lateralised. However, evidence in support of this distinction has been garnered exclusively from tasks that involved static, two-dimensional (2D) arrays. We used functional magnetic resonance imaging (fMRI) to identify neural circuits underlying categorical and coordinate representations during active spatial navigation. Activity in the categorical condition was significantly greater in the parietal cortex, whereas the coordinate condition revealed greater activity in medial temporal cortex and dorsal striatum. In addition, activity in the categorical condition was greater in parietal cortex within the left hemisphere than within the right Our findings are consistent with analogous studies in rodents, and support the suggestion of distinct neural circuits underlying categorical and coordinate representations during active spatial navigation. The findings also support the claim of a left hemispheric preponderance for the processing of categorical spatial relations. (C) 2012 Elsevier Inc. All rights reserved

Bodily awareness and novel multisensory features

According to the decomposition thesis, perceptual experiences resolve without remainder into their different modality-specific components. Contrary to this view, I argue that certain cases of multisensory integration give rise to experiences representing features of a novel type. Through the coordinated use of bodily awareness—understood here as encompassing both proprioception and kinaesthesis—and the exteroceptive sensory modalities, one becomes perceptually responsive to spatial features whose instances couldn’t be represented by any of the contributing modalities functioning in isolation. I develop an argument for this conclusion focusing on two cases: 3D shape perception in haptic touch and experiencing an object’s egocentric location in crossmodally accessible, environmental space

From sensory perception to spatial cognition

To interact with the environmet, it is crucial to have a clear space representation. Several findings have shown that the space around our body is split in several portions, which are differentially coded by the brain. Evidences of such subdivision have been reported by studies on people affected by neglect, on space near (peripersonal) and far (extrapersonal) to the body position and considering space around specific different portion of the body. Moreover, recent studies showed that sensory modalities are at the base of important cognitive skills. However, it is still unclear if each sensory modality has a different role in the development of cognitive skills in the several portions of space around the body. Recent works showed that the visual modality is crucial for the development of spatial representation. This idea is supported by studies on blind individuals showing that visual information is fundamental for the development of auditory spatial representation. For example, blind individuals are not able to perform the spatial bisection task, a task that requires to build an auditory spatial metric, a skill that sighted children acquire around 6 years of age. Based these prior researches, we hypothesize that if different sensory modalities have a role on the devlopment of different cognitive skills, then we should be able to find a clear correlation between availability of the sensory modality and the cognitive skill associated. In particular we hypothesize that the visual information is crucial for the development of auditory space represnetation; if this is true, we should find different spatial skill between front and back spaces. In this thesis, I provide evidences that spaces around our body are differently influenced by sensory modalities. Our results suggest that visual input have a pivotal role in the development of auditory spatial representation and that this applies only to the frontal space. Indeed sighted people are less accurated in spatial task only in space where vision is not present (i.e. the back), while blind people show no differences between front and back spaces. On the other hand, people tend to report sounds in the back space, suggesting that the role of hearing in allertness could be more important in the back than frontal spaces. Finally, we show that natural training, stressing the integration of audio motor stimuli, can restore spatial cognition, opening new possibility for rehabilitation programs. Spatial cognition is a well studied topic. However, we think our findings fill the gap regarding how the different availibility of sensory information, across spaces, causes the development of different cognitive skills in these spaces. This work is the starting point to understand the strategies that the brain adopts to maximize its resources by processing, in the more efficient way, as much information as possible

Mental Imagery for Full and Upper Human Bodies: Common Right Hemisphere Activations and Distinct Extrastriate Activations

The processing of human bodies is important in social life and for the recognition of another person's actions, moods, and intentions. Recent neuroimaging studies on mental imagery of human body parts suggest that the left hemisphere is dominant in body processing. However, studies on mental imagery of full human bodies reported stronger right hemisphere or bilateral activations. Here, we measured functional magnetic resonance imaging during mental imagery of bilateral partial (upper) and full bodies. Results show that, independently of whether a full or upper body is processed, the right hemisphere (temporo-parietal cortex, anterior parietal cortex, premotor cortex, bilateral superior parietal cortex) is mainly involved in mental imagery of full or partial human bodies. However, distinct activations were found in extrastriate cortex for partial bodies (right fusiform face area) and full bodies (left extrastriate body area). We propose that a common brain network, mainly on the right side, is involved in the mental imagery of human bodies, while two distinct brain areas in extrastriate cortex code for mental imagery of full and upper bodie

Balancing Interoception and Exteroception: Vestibular and Spatial Contributions to the Bodily Self

Experiencing the body as a coherent, stable, entity involves the dynamic integration of information from several internal (i.e. interoceptive) and external (i.e. exteroceptive) sensory sources, to produce a feeling that the body is mine (sense of body ownership), that I am in control (sense of agency) and I am aware of its movements (motor awareness). However, the exact contribution of these different sensory sources to self-consciousness, as well as the context in which we experience them, is still a matter of debate. This thesis aimed to investigate the neurocognitive mechanisms of body ownership, agency and motor awareness, including interoceptive (via affective touch), proprioceptive, exteroceptive (visuo-spatial) and vestibular contributions to body representation, in both healthy subjects and brain damaged patients. To examine the role of the vestibular and interoceptive systems in body ownership, a series of studies in healthy subjects was devised, using multisensory illusions (i.e. the rubber hand illusion; RHI), that involve the integration of interoceptive and exteroceptive sensory sources, and using electrical stimulation of the vestibular system (i.e. Galvanic Vestibular Stimulation; GVS). To investigate ownership, agency and motor awareness in neuropsychological patients with disorders of ownership and/or unawareness of motor deficits, behavioural manipulation of body ownership (via a rubber hand) and visual perspective (via a mirror) were tested. Finally, to explore underlying mechanisms of awareness of one’s own performance (i.e. meta-cognition), two studies were carried out in healthy subjects using behavioural manipulations of spatial reference frames (either centred on the subject, i.e. egocentric, or world-centred, i.e. allocentric). The results of these studies indicate that the vestibular system balances vision and proprioception according to contextual relevance: when there is no tactile stimulation, visual cues are stronger than proprioceptive ones (i.e. proprioceptive drifts are greater); when touch is delivered synchronously, this effect is enhanced (even more when touch is affective rather than neutral). However, when touch is only felt but not seen, the vestibular system downregulates vision in favour of proprioception (i.e. proprioceptive drifts are smaller), whilst the opposite happens when touch is only vicariously perceived via vision. Nevertheless, when the rubber hand is positioned in a non-biomechanically possible fashion, there appears to be no difference in proprioceptive drifts in comparison with anatomically plausible positions, suggesting that such rebalancing may be more related to basic multisensory integration processes underlying body representation. In patients with disorders of the self, visual cues seem to dominate over proprioceptive ones, leading to strong feelings of ownership of a rubber hand following mere exposure to it; however, the same is not true for agency, which seems to be more susceptible to changes in the environment (i.e. presence or absence of a visual feedback following attempted movement). Moreover, manipulating visual perspective using a mirror (from 1st to 3rd) seem to lead to a temporary remission of dis-ownership but not motor unawareness, suggesting that awareness may not be influenced by online changes in visual perspectives. Finally, when judging their own performance in a visuo-proprioceptive task from an egocentric rather than an allocentric perspective, healthy subjects appear less objective prospectively rather than during the task (i.e. their belief updating is biased when judging their ability to complete a task egocentrically). In sum, the work described above adds to the evidence that the sense of self derives from a complex integration of several sensory modalities, flexibly adjusting to the environment. Following brain damage, such flexibility may be impaired, even though it can be influenced by spatial perspective. Similarly, the point of reference from which we perceive stimuli affects the way we judge our own perceptual choices. Hence, the way we represent our bodily self is a dynamic process, constantly updated by exteroceptive and interoceptive incoming stimuli, regulated by the vestibular system. These findings could provide new avenues in rehabilitating disorders of the self (such as unawareness and dis-ownership)

Handedness effects of imagined fine motor movements

Previous studies of movement imagery have found inter-individual differences in the ability to imagine whole-body movements. The majority of these studies have used subjective scales to measure imagery ability, which may be confounded by other factors related to effort. Madan and Singhal [2013. Introducing TAMI: An objective test of ability in movement imagery. Journal of Motor Behavior, 45(2), 153–166. doi:10.1080/00222895.2013.763764] developed the Test of Ability in Movement Imagery (TAMI) to address these confounds by using a multiple-choice format with objectively correct responses. Here we developed a novel movement imagery questionnaire targeted at assessing movement imagery of fine-motor hand movements. This questionnaire included two subscales: Functionally-involved Movement (i.e., tool-related) and Isolated Movement (i.e., hand-only). Hand-dominance effects were observed, such that right-handed participants were significantly better at responding to right-hand questions compared to left-hand questions for both imagery types. A stronger handedness effect was observed for Functionally-involved Movement imagery, and it did not correlate with the Edinburgh Handedness Inventory. We propose that the Functionally-involved Movement imagery subscale provides an objective hand imagery test that induces egocentric spatial processing and a greater involvement of memory processes, potentially providing a better skill-based measure of handedness