Benefits of active listening during 3D sound localization.

In everyday life, sound localization entails more than just the extraction and processing of auditory cues. When determining sound position in three dimensions, the brain also considers the available visual information (e.g., visual cues to sound position) and resolves perceptual ambiguities through active listening behavior (e.g., spontaneous head movements while listening). Here, we examined to what extent spontaneous head movements improve sound localization in 3D-azimuth, elevation, and depth-by comparing static vs. active listening postures. To this aim, we developed a novel approach to sound localization based on sounds delivered in the environment, brought into alignment thanks to a VR system. Our system proved effective for the delivery of sounds at predetermined and repeatable positions in 3D space, without imposing a physically constrained posture, and with minimal training. In addition, it allowed measuring participant behavior (hand, head and eye position) in real time. We report that active listening improved 3D sound localization, primarily by ameliorating accuracy and variability of responses in azimuth and elevation. The more participants made spontaneous head movements, the better was their 3D sound localization performance. Thus, we provide proof of concept of a novel approach to the study of spatial hearing, with potentials for clinical and industrial applications.

Associative learning in peripersonal space: fear responses are acquired in hand-centered coordinates.

Space coding affects perception of stimuli associated to negative valence: threatening stimuli presented within the peripersonal space (PPS) speed up behavioral responses compared with nonthreatening events. However, it remains unclear whether the association between stimuli and their negative valence is acquired in a body part-centered reference system, a main feature of the PPS coding. Here we test the hypothesis that associative learning takes place in hand-centered coordinates and can therefore remap according to hand displacement. In two experiments, we used a Pavlovian fear-learning paradigm to associate a visual stimulus [light circle, the conditioned stimulus (CS)] with an aversive stimulus (electrocutaneous shock) applied on the right hand only when the CS was displayed close (CS+) but when not far from it (CS-). Measuring the skin conductance response (SCR), we observed successful fear conditioning, with increased anticipatory fear responses associated with CS+. Crucially, experiment I showed a remapping of these responses following hand displacement, with a generalization to both types of CS. Experiment II corroborated and further extended our findings by ruling out the novelty of the experimental context as a driving factor of such modulations. Indeed, fear responses were present only for stimuli within the PPS but not for new stimuli displayed outside the PPS. By revealing a hand-centered (re)mapping of the conditioning effect, these findings indicate that associative learning can arise in hand-centered coordinates. They further suggest that the threatening valence of an object also depends on its basic spatial relationship with our body.NEW & NOTEWORTHY Associative fear learning takes place in hand-centered coordinates. Using a Pavlovian fear-learning paradigm, we show that the anticipatory skin conductance response indicating the association between the negative value and an initially neutral stimulus is acquired and then remapped in space when the stimulated body part moves to a different position. These results demonstrate the relationship between the representation of peripersonal space and the encoding of threatening stimuli. Hypotheses concerning the underlying neural network are discussed.

Peripersonal and reaching space differ: Evidence from their spatial extent and multisensory facilitation pattern.

Peripersonal space (PPS) is a multisensory representation of the space near body parts facilitating interactions with the close environment. Studies on non-human and human primates agree in showing that PPS is a body part-centered representation that guides actions. Because of these characteristics, growing confusion surrounds peripersonal and arm-reaching space (ARS), that is the space one's arm can reach. Despite neuroanatomical evidence favoring their distinction, no study has contrasted directly their respective extent and behavioral features. Here, in five experiments (N = 140) we found that PPS differs from ARS, as evidenced both by participants' spatial and temporal performance and by its modeling. We mapped PPS and ARS using both their respective gold standard tasks and a novel multisensory facilitation paradigm. Results show that: (1) PPS is smaller than ARS; (2) multivariate analyses of spatial patterns of multisensory facilitation predict participants' hand locations within ARS; and (3) the multisensory facilitation map shifts isomorphically following hand positions, revealing hand-centered coding of PPS, therefore pointing to a functional similarity to the receptive fields of monkeys' multisensory neurons. A control experiment further corroborated these results and additionally ruled out the orienting of attention as the driving mechanism for the increased multisensory facilitation near the hand. In sharp contrast, ARS mapping results in a larger spatial extent, with undistinguishable patterns across hand positions, cross-validating the conclusion that PPS and ARS are distinct spatial representations. These findings show a need for refinement of theoretical models of PPS, which is relevant to constructs as diverse as self-representation, social interpersonal distance, and motor control.

Eye dominance modulates visuospatial attention.

Visuospatial attention has an inherent asymmetry: the leftward bias called pseudoneglect. In typical line bisection tasks, healthy individuals tend to judge the center of a line leftward of the true center, an effect attributed to the right hemisphere dominance in visuospatial attention. Since it has been shown that information perceived by the dominant eye strongly activates the ipsilateral visual cortex, we hypothesized that eye dominance may modulate visuospatial attention bias. Because activation of the left hemisphere induced by left eye dominance should mitigate the right hemisphere dominance in attention, we predicted that right-handed individuals with left dominant eye would show smaller amount of pseudoneglect than right-handed individuals with right dominant eye. We compared the performance at both the perceptual (Landmark) and manual line bisection task of forty right-handed healthy individuals, half of whom had a right dominant eye and the other half a left dominant eye. As predicted, the left eyed dominant group showed smaller, actually not significant pseudoneglect, which was thus greater in the right eye dominant group. The influence of eye dominance on visuospatial attention was present in the Landmark but not the manual line bisection task, in which the amount of visuospatial bias correlated with participants' degree of (right) handedness. This is the first report of the effect of eye dominance on visuospatial attention within a right-handed population. This finding, by showing the influence of eye dominance on visuospatial cognition, not only helps in better defining intact visuospatial cognition mechanism but also encourages further research to pinpoint the neural basis of such interaction.

The Attentional Fields of Visual Search in Simultanagnosia and Healthy Individuals: How Object and Space Attention Interact.

Simultanagnosia is a deficit in which patients are unable to perceive multiple objects simultaneously. To date, it remains disputed whether this deficit results from disrupted object or space perception. We asked both healthy participants as well as a patient with simultanagnosia to perform different visual search tasks of variable difficulty. We also modulated the number of objects (target and distracters) presented. For healthy participants, we found that each visual search task was performed with a specific "attentional field" depending on the difficulty of visual object processing but not on the number of objects falling within this "working space." This was demonstrated by measuring the cost in reaction times using different gaze-contingent visible window sizes. We found that bilateral damage to the superior parietal lobule impairs the spatial integration of separable features (within-object processing), shrinking the attentional field in which a target can be detected, but causing no deficit in processing multiple objects per se.

Effects of structural and functional cerebellar lesions on sensorimotor adaptation of saccades.

The cerebellum is critically involved in the adaptation mechanisms that maintain the accuracy of goal-directed acts such as saccadic eye movements. Two categories of saccades, each relying on different adaptation mechanisms, are defined: reactive (externally triggered) saccades and voluntary (internally triggered) saccades. The contribution of the medio-posterior part of the cerebellum to reactive saccades adaptation has been clearly demonstrated, but the evidence that other parts of the cerebellum are also involved is limited. Moreover, the cerebellar substrates of voluntary saccades adaptation have only been marginally investigated. Here, we addressed these two questions by investigating the adaptive capabilities of patients with cerebellar or pre-cerebellar stroke. We recruited three groups of patients presenting focal lesions located, respectively, in the supero-anterior cerebellum, the infero-posterior cerebellum and the lateral medulla (leading to a Wallenberg syndrome including motor dysfunctions similar to those resulting from lesion of the medio-posterior cerebellum). Adaptations of reactive saccades and of voluntary saccades were tested during separate sessions in all patients and in a group of healthy participants. The functional lesion of the medio-posterior cerebellum in Wallenberg syndrome strongly impaired the adaptation of both reactive and voluntary saccades. In contrast, patients with lesion in the supero-anterior part of the cerebellum presented a specific adaptation deficit of voluntary saccades. Finally, patients with an infero-posterior cerebellar lesion showed mild adaptation deficits. We conclude that the medio-posterior cerebellum is critical for the adaptation of both saccade categories, whereas the supero-anterior cerebellum is specifically involved in the adaptation of voluntary saccades.

The Rubber Hand Illusion: two's a company, but three's a crowd.

On the one hand, it is often assumed that the Rubber Hand Illusion (RHI) is constrained by a structural body model so that one cannot implement supernumerary limbs. On the other hand, several recent studies reported illusory duplication of the right hand in subjects exposed to two adjacent rubber hands. The present study tested whether spatial constraints may affect the possibility of inducing the sense of ownership to two rubber hands located side by side to the left of the subject's hand. We found that only the closest rubber hand appeared both objectively (proprioceptive drift) and subjectively (ownership rating) embodied. Crucially, synchronous touch of a second, but farther, rubber hand disrupted the objective measure of the RHI, but not the subjective one. We concluded that, in order to elicit a genuine RHI for multiple rubber hands, the two rubber hands must be at the same distance from the subject's hand/body.

When action is not enough: tool-use reveals tactile-dependent access to Body Schema.

Proper motor control of our own body implies a reliable representation of body parts. This information is supposed to be stored in the Body Schema (BS), a body representation that appears separate from a more perceptual body representation, the Body Image (BI). The dissociation between BS for action and BI for perception, originally based on neuropsychological evidence, has recently become the focus of behavioural studies in physiological conditions. By inducing the rubber hand illusion in healthy participants, Kammers et al. (2009) showed perceptual changes attributable to the BI to which the BS, as indexed via motor tasks, was immune. To more definitively support the existence of dissociable body representations in physiological conditions, here we tested for the opposite dissociation, namely, whether a tool-use paradigm would induce a functional update of the BS (via a motor localization task) without affecting the BI (via a perceptual localization task). Healthy subjects were required to localize three anatomical landmarks on their right arm, before and after using the same arm to control a tool. In addition to this classical task-dependency approach, we assessed whether preferential access to the BS could also depend upon the way positional information about forearm targets is provided, to subsequently execute the same task. To this aim, participants performed either verbally or tactually driven versions of the motor and perceptual localization tasks. Results showed that both the motor and perceptual tasks were sensitive to the update of the forearm representation, but only when the localization task (perceptual or motor) was driven by a tactile input. This pattern reveals that the motor output is not sufficient per se, but has to be coupled with tactually mediated information to guarantee access to the BS. These findings shade a new light on the action-perception models of body representations and underlie how functional plasticity may be a useful tool to clarify their operational definition.

Parietal damage dissociates saccade planning from presaccadic perceptual facilitation.

A well-known theory in the field of attention today is the premotor theory of attention which suggests that the mechanisms involved in eye movements are the same as those for spatial attention shifts. We tested a parietal damaged patient with unilateral optic ataxia and 4 controls on a dual saccade/attentional task and show a dissociation between saccadic eye movements and presaccadic perceptual enhancement at the saccade goal. Remarkably, though the patient was able to make the appropriate saccades to the left, impaired visual field (undistinguishable from saccades to his right, intact visual field), he was unable to discriminate the letter at the saccade goal (whereas his performance was like controls for letter discrimination in his right visual field). This suggests that saccade planning and presaccadic perceptual facilitation are separable--planning a saccade to a location does not necessitate that the processing of this location is enhanced. Based on these results, we suggest that the parietal cortex is necessary for the coupling between saccade planning and presaccadic perceptual facilitation.

Separate neural substrates in the human cerebellum for sensory-motor adaptation of reactive and of scanning voluntary saccades.

Sensory-motor adaptation processes are critically involved in maintaining accurate motor behavior throughout life. Yet their underlying neural substrates and task-dependency bases are still poorly understood. We address these issues here by studying adaptation of saccadic eye movements, a well-established model of sensory-motor plasticity. The cerebellum plays a major role in saccadic adaptation but it has not yet been investigated whether this role can account for the known specificity of adaptation to the saccade type (e.g., reactive versus voluntary). Two patients with focal lesions in different parts of the cerebellum were tested using the double-step target paradigm. Each patient was submitted to two separate sessions: one for reactive saccades (RS) triggered by the sudden appearance of a visual target and the second for scanning voluntary saccades (SVS) performed when exploring a more complex scene. We found that a medial cerebellar lesion impaired adaptation of reactive-but not of voluntary-saccades, whereas a lateral lesion affected adaptation of scanning voluntary saccades, but not of reactive saccades. These findings provide the first evidence of an involvement of the lateral cerebellum in saccadic adaptation, and extend the demonstrated role of the cerebellum in RS adaptation to adaptation of SVS. The double dissociation of adaptive abilities is also consistent with our previous hypothesis of the involvement in saccadic adaptation of partially separated cerebellar areas specific to the reactive or voluntary task (Alahyane et al. Brain Res 1135:107-121 (2007)).

Optic ataxia errors depend on remapped, not viewed, target location.

Optic ataxia is a disorder associated with posterior parietal lobe lesions, in which visually guided reaching errors typically occur for peripheral targets. It has been assumed that these errors are related to a faulty sensorimotor transformation of inputs from the 'ataxic visual field'. However, we show here that the errors observed in the contralesional field in optic ataxia depend on a dynamic gaze-centered internal representation of reach space.