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Visuo-motor adaptation to optical prisms displacing the visual scene (prism adaptation, PA) is a method used for investigating visuo-motor plasticity in healthy individuals and, in clinical settings, for the rehabilitation of unilateral spatial neglect. In the standard paradigm, the adaptation phase involves repeated pointings to visual targets, while wearing optical prisms displacing the visual scene laterally. Here we explored differences in PA, and its aftereffects (AEs), as related to the sensory modality of the target. Visual, auditory, and multisensory - audio-visual - targets in the adaptation phase were used, while participants wore prisms displacing the visual field rightward by 10°. Proprioceptive, visual, visual-proprioceptive, auditory-proprioceptive straightahead shifts were measured. Pointing to auditory and to audio-visual targets in the adaptation phase produces proprioceptive, visual-proprioceptive, and auditoryproprioceptive AEs, as the typical visual targets did. This finding reveals that crossmodal plasticity effects involve both the auditory and the visual modality, and their interactions (Experiment 1). Even a shortened PA phase, requiring only 24 pointings to visual and audio-visual targets (Experiment 2), is sufficient to bring about AEs, as compared to the standard 92-pointings procedure. Finally, pointings to auditory targets cause AEs, although PA with a reduced number of pointings (24) to auditory targets brings about smaller AEs, as compared to the 92-pointings procedure (Experiment 3). Together, results from the three experiments extend to the auditory modality the sensorimotor plasticity underlying the typical AEs produced by PA to visual targets. Importantly, PA to auditory targets appears characterized by less accurate pointings and error correction, suggesting that the auditory component of the PA process may be less central to the building up of the AEs, than the sensorimotor pointing activity per se. These findings highlight both

The stage at which processing of tactile distance occurs is still debated. We addressed this issue by implementing an adaptationaftereffect paradigm with passive touch. We demonstrated the presence of a strong aftereffect, induced by the simultaneous presentation of pairs of tactile stimuli. After adaptation to two different distances, one on each hand, participants systematically perceived a subsequent stimulus delivered to the hand adapted to the smaller distance as being larger. We further investigated the nature of the aftereffects, demonstrating that they are orientation- and skin-region-specific, occur even when just one hand is adapted, do not transfer either contralaterally or across the palm and dorsum, and are defined in a skin-centered, rather than an external, reference frame. These characteristics of tactile distance aftereffects are similar to those of low-level visual aftereffects, supporting the idea that distance perception arises at early stages of tactile processing.

My Doctoral Thesis investigated different aspects of bodily and spatial representations, how they are modulated by multisensory stimulation, and some physiological correlates of their manipulation. Chapter #1, “The space around the body”, reports three studies on prism adaptation, a technique that takes advantage of brain plasticity in the generation and modification of spatial bodily maps. Specifically, Study #1 investigates how the vision of the limb during two different versions of prism adaptation modulates their aftereffects: larger aftereffects take place with a concurrent vision of the limb after prism adaptation achieved through “ecological” visuo-motor activities; conversely, the vision of the very last part of the movement brings about larger aftereffects after the repeated pointing task. Study #2 examines the effect of a multisensory stimulation during prism adaptation, showing that the pointing error reduction is obtained with fewer pointing movements when the target is a visual-acoustic (multisensory), rather than unisensory stimulus. Finally, Study #3, which was performed in a brain-damaged patient, showed that the integrity of the left parieto-cerebellar circuit is required for an appropriate spatial remapping of proprioceptive maps to occur, and that the modulation through transcranial Direct Current Stimulation of these cortical areas temporarily restores the aftereffects. Chapter #2, “The space in the body”, reports studies on the link between bodily spatial representations and homeostatic regulation. Skin temperature has been recently considered as a physiological parameter of disembodiment, and can be modulated by the manipulation of bodily representations. Three experiments in which spatial bodily maps were manipulated by means of different techniques, inducing direction-specific and lateralized effects, are presented. Specifically, Study #4 showed a reduction of hands’ skin temperature after adaptation to right shifting, but not to left shiftin

The physiological regulation of skin temperature can be modulated not only by autonomic brain regions, but also by a network of higher-level cortical areas involved in the maintenance of a coherent representation of the body. In this study we assessed in healthy participants if the sensorimotor changes taking place during motor adaptation to the lateral displacement of the visual scene induced by wearing prismatic lenses (prism adaptation, PA), and the aftereffects, after prisms' removal, on the ability to process spatial coordinates, were associated with skin temperature regulation changes. We found a difference in thermoregulatory control as a function of the direction of the prism-induced displacement of the visual scene, and the subsequent sensorimotor adaptation. After PA to rightward displacing lenses, with leftward aftereffects (the same directional procedure efficaciously used for ameliorating left spatial neglect in right-brain-damaged patients) the hands' temperature decreased. Conversely, after adaptation to neutral lenses, and PA to leftward displacing lenses, with rightward aftereffects, the temperature of both hands increased. These results suggest a lateral asymmetry in the effects of PA on skin temperature regulation, and a relationship between body spatial representations and homeostatic control in humans.

Adaptation to optical prisms displacing the visual scene laterally is a widely investigated instance of visuo-motor plasticity, also because prism adaptation (PA) has been extensively used as a treatment for right-brain-damaged patients suffering from left spatial neglect. The lateral visual displacement brought about by prisms, as indexed by a pointing error in the direction of the displacement, is progressively corrected through repeated pointings: after prism removal, a shift in the direction opposite to the prism-induced deviation occurs in visual, proprioceptive, and visuo-proprioceptive straight-ahead tasks (aftereffects, AEs). The cerebellum and the posterior parietal cortex (PPC) are key components of the bilateral cerebral network subserving the AEs, and the reduction of the pointing error during prism exposure in PA. We report the experimental study of a patient with bilateral occipital and left cerebellar damage, who showed a preserved reduction of the pointing errors to rightward displacing prisms, but not the leftward AEs in the proprioceptive straight-ahead task; instead, visual-proprioceptive and visual AEs were preserved. Anodal transcranial Direct Current Stimulation (tDCS) over the left PPC restored the leftward proprioceptive AEs, and anodal tDCS over the left cerebellum abolished the rightward deviation. Conversely, stimulation over the right PPC or the right cerebellum was ineffective. These results provide novel evidence for neuromodulatory effects of tDCS on defective AEs, through the stimulation over dedicated cortical regions.

Prism adaptation improves a wide range of manifestations of left spatial neglect in right-brain-damaged patients. The typical paradigm consists in repeated pointing movements to visual targets, while patients wear prism goggles that displace the visual scene rightwards. Recently, we demonstrated the efficacy of a novel adaptation procedure, involving a variety of every-day visuo-motor activities. This "ecological" procedure proved to be as effective as the repetitive pointing adaptation task in ameliorating symptoms of spatial neglect, and was better tolerated by patients. However, the absence of adaptation and aftereffects measures for the ecological treatment did not allow for a full comparison of the two procedures. This is important in the light of recent findings showing that the magnitude of prism-induced aftereffects may predict recovery from spatial neglect. Here, we investigated prism-induced adaptation and aftereffects after ecological and pointing adaptation procedures. Forty-eight neurologically healthy participants (young and aged groups) were exposed to rightward shifting prisms while they performed the ecological or the pointing procedures, in separate days. Before and after prism exposure, participants performed proprioceptive, visual, and visual-proprioceptive tasks to assess prism-induced aftereffects. Participants adapted to the prisms during both procedures. Importantly, the ecological procedure induced greater aftereffects in the proprioceptive task (for both the young and the aged groups) and in the visual-proprioceptive task (young group). A similar trend was found for the visual task in both groups. Finally, participants rated the ecological procedure as more pleasant, less monotonous, and more sustainable than the pointing procedure. These results qualify ecological visuo-motor activities as an effective prism-adaptation procedure, suitable for the rehabilitation of spatial neglect.