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Temporal Binding Window (TBW) represents a reliable index of efficient multisensory integration process, which allows individuals to infer which sensory inputs from different modalities pertain to the same event. TBW alterations have been reported in some neurological and neuropsychiatric disorders and seem to negatively affects cognition and behavior. So far, it is still unknown whether deficits of multisensory integration, as indexed by an abnormal TBW, are present even in Multiple Sclerosis. We addressed this issue by testing 25 participants affected by relapsing-remitting Multiple Sclerosis (RRMS) and 30 age-matched healthy controls. Participants completed a simultaneity judgment task (SJ2) to assess the audio-visual TBW; two unimodal SJ2 versions were used as control tasks. Individuals with RRMS showed an enlarged audio-visual TBW (width range = from - 166 ms to + 198 ms), as compared to healthy controls (width range = - 177/ + 66 ms), thus showing an increased tendency to integrate temporally asynchronous visual and auditory stimuli. Instead, simultaneity perception of unimodal (visual or auditory) events overall did not differ from that of controls. These results provide first evidence of a selective deficit of multisensory integration in individuals affected by RRMS, besides the well-known motor and cognitive impairments. The reduced multisensory temporal acuity is likely caused by a disruption of the neural interplay between different sensory systems caused by multiple sclerosis., (© 2022. The Author(s).)

Motor learning interacts with and shapes experience-dependent cerebral plasticity. In stroke patients with paresis of the upper limb, motor recovery was proposed to reflect a process of re-learning the lost/impaired skill, which interacts with rehabilitation. However, to what extent stroke patients with hemiparesis may retain the ability of learning with their affected limb remains an unsolved issue, that was addressed by this study. Nineteen patients, with a cerebrovascular lesion affecting the right or the left hemisphere, underwent an explicit motor learning task (finger tapping task, FTT), which was performed with the paretic hand. Eighteen age-matched healthy participants served as controls. Motor performance was assessed during the learning phase (i.e., online learning), as well as immediately at the end of practice, and after 90 min and 24 h (i.e., retention). Results show that overall, as compared to the control group, stroke patients, regardless of the side (left/right) of the hemispheric lesion, do not show a reliable practice-dependent improvement; consequently, no retention could be detected in the long-term (after 90 min and 24 h). The motor learning impairment was associated with subcortical damage, predominantly affecting the basal ganglia; conversely, it was not associated with age, time elapsed from stroke, severity of upper-limb motor and sensory deficits, and the general neurological condition. This evidence expands our understanding regarding the potential of post-stroke motor recovery through motor practice, suggesting a potential key role of basal ganglia, not only in implicit motor learning as previously pointed out, but also in explicit finger tapping motor tasks., (© 2021. The Author(s).)

Right-brain-damaged patients with left unilateral spatial neglect typically set the mid-point of horizontal lines to the right of the objective center. By contrast, healthy participants exhibit a reversed bias (pseudoneglect). The same effect has been described also when bisecting orthographic strings. In particular, for this latter kind of stimulus, some recent studies have shown that visuo-perceptual characteristics, like stimulus length, may contribute to both the magnitude and the direction bias of the bisection performance (Arduino et al. in Neuropsychologia 48:2140-2146, 2010). Furthermore, word stress was shown to modulate reading performances in both healthy participants, and patients with left spatial neglect and neglect dyslexia (Cubelli and Beschin in Brain Lang 95:319-326, 2005; Rusconi et al. in Neuropsychology 18:135-140, 2004). In Experiment I, 22 right-brain-damaged patients (11 with left visuo-spatial neglect) and 11 matched neurologically unimpaired control participants were asked to set the subjective mid-point of word letter strings, and of lines of comparable length. Most patients exhibited an overall disproportionate rightward bias, sensitive to stimulus length, and similar for words and lines. Importantly, in individual patients, biases differed according to stimulus type (words vs. lines), indicating that at least partly different mechanisms may be involved. In Experiment II, the putative effects on the bisection bias of ortho-phonological information (i.e., word stress endings), arising from the non-neglected right hand side of the stimulus were investigated. The orthographic cue induced a rightward shift of the perceived mid-point in both patients and controls, with short words stressed on the antepenultimate final sequence inducing a smaller rightward deviation with respect to short words stressed on the penultimate final sequence. In conclusion, partly different mechanisms, including both visuo-spatial and lexical factors, may support line and word bisection performance of right-brain-damaged patients with left spatial neglect, and healthy participants.

Crossmodal illusions clearly show how perception, rather than being a modular and self-contained function, can be dramatically altered by interactions between senses. Here, we provide evidence for a novel crossmodal "physiological" illusion, showing that sounds can boost visual cortical responses in such a way to give rise to a striking illusory visual percept. In healthy participants, a single-pulse transcranial magnetic stimulation (sTMS) delivered to the occipital cortex evoked a visual percept, i.e., a phosphene. When sTMS is accompanied by two auditory beeps, the second beep induces in neurologically unimpaired participants the perception of an illusory second phosphene, namely the sound-induced phosphene illusion. This perceptual "fission" of a single phosphene, due to multiple beeps, is not matched by a "fusion" of double phosphenes due to a single beep, and it is characterized by an early auditory modulation of the TMS-induced visual responses (~80 ms). Multiple beeps also induce an illusory feeling of multiple TMS pulses on the participants' scalp, consistent with an audio-tactile fission illusion. In conclusion, an auditory stimulation may bring about a phenomenological change in the conscious visual experience produced by the transcranial stimulation of the occipital cortex, which reveals crossmodal binding mechanisms within early stages of visual processing.

Numbers induce shifts of spatial attention on the left or the right sides of external space as a function of their magnitude. However, whether this number-space association is restricted to the linear horizontal extensions, or extends to the whole visual scene, is still an open question. This study investigates, by means of a cancellation paradigm, the influence of numerical magnitude during scanning tasks in which participants freely explore complex visual scenes unconstrained towards either the horizontal or the vertical unidimensional axes. Five cancellation tasks were adapted in which Arabic digits were used as targets or distracters, in structured (lines and columns) or unstructured visual displays, with a smaller (2 or 3 types of distracters) or larger (10 or more types of distracters) sets of stimuli. Results show that the participants' hits distribution was a function of number magnitude: shifted on the left for small and on the right for large numbers. This effect was maximised when numerical cues were sparse, randomly arranged and, critically, irrelevant to the task. Overall, this study provides novel evidence from visuo-spatial exploratory cancellation tasks for an attentional shift induced by number magnitude.

Distracting gaze has been shown to elicit automatic gaze following. However, it is still debated whether the effects of perceived gaze are a simple automatic spatial orienting response or are instead sensitive to the context (i.e. goals and task demands). In three experiments, we investigated the conditions under which gaze following occurs. Participants were instructed to saccade towards one of two lateral targets. A face distracter, always present in the background, could gaze towards: (a) a task-relevant target--("matching" goal-directed gaze shift)--congruent or incongruent with the instructed direction, (b) a task-irrelevant target, orthogonal to the one instructed ("non-matching" goal-directed gaze shift), or (c) an empty spatial location (no-goal-directed gaze shift). Eye movement recordings showed faster saccadic latencies in correct trials in congruent conditions especially when the distracting gaze shift occurred before the instruction to make a saccade. Interestingly, while participants made a higher proportion of gaze-following errors (i.e. errors in the direction of the distracting gaze) in the incongruent conditions when the distracter's gaze shift preceded the instruction onset indicating an automatic gaze following, they never followed the distracting gaze when it was directed towards an empty location or a stimulus that was never the target. Taken together, these findings suggest that gaze following is likely to be a product of both automatic and goal-driven orienting mechanisms.

Numerical magnitude is believed to be represented along a mental number line (MNL), and there is evidence to suggest that the activation of the MNL affects the perception and representation of external space. In the present study, we investigated whether a spatial motor task affects numerical processing in the auditory modality. Blindfolded participants were presented with a numerical interval bisection task, while performing a tapping task with either their left or right hand, either in the fronto-central, fronto-left, or fronto-right peripersonal space. Results showed that tapping significantly influenced the participants' numerical bisection, with tapping in the left side of space increasing the original tendency to err leftward, and tapping to the right reducing such bias. Importantly, the effect depended on the side of space in which the tapping activity was performed, regardless of which hand was used. Tapping with either the left or right hand in the fronto-central space did not affect the participants' bias. These findings offer novel support for the existence of bidirectional interactions between external and internal representations of space.

Neglect dyslexia (ND) is reviewed, based on published single-patient and group studies. ND is frequently associated with right hemispheric damage and unilateral spatial neglect (USN), and typically involves the left side of the letter string. Left-brain-damaged patients showing ND, ipsilateral (left) or contralateral (right) to the side of the left-sided hemispheric lesion, have also been reported, as well as a few patients with bilateral damage, with more frequently left than right ND. As USN, ND is temporarily ameliorated by lateralized stimulations (vestibular caloric, visual prism adaptation). ND may occur independent of USN, suggesting the damage to specific visuospatial representational/attentional systems, supporting reading. ND errors comprise omission, substitution, and, less frequently, addition of letters on one side of the stimulus, resulting in words or nonwords, also with reference to the stimulus' linguistic features. Patients with ND may show preserved lexical-morphological effects and implicit processing, up to the semantic level, of the misread string. This preserved processing is a feature of ND, shared with the USN syndrome. The mechanisms modulating error type and lexical-morphological effects are partly independent of each other. Different levels of representation of the letter string may be affected, giving rise to egocentric, stimulus-centred, and word-centred patterns of impairment. The anatomical correlates of ND include the temporo-parieto-occipital regions.

A review of published brain-damaged patients showing delusional beliefs concerning the contralesional side of the body (somatoparaphrenia) is presented. Somatoparaphrenia has been reported, with a few exceptions, in right-brain-damaged patients, with motor and somatosensory deficits, and the syndrome of unilateral spatial neglect. Somatoparaphrenia, most often characterized by a delusion of disownership of left-sided body parts, may however occur without associated anosognosia for motor deficits, and personal neglect. Also somatosensory deficits may not be a core pathological mechanism of somatoparaphrenia, and visual field disorders may be absent. Deficits of proprioception, however, may play a relevant role. Somatoparaphrenia is often brought about by extensive right-sided lesions, but patients with posterior (parietal-temporal), and insular damage are on record, as well as a few patients with subcortical lesions. Possible pathological factors include a deranged representation of the body concerned with ownership, mainly right-hemisphere-based, and deficits of multisensory integration. Finally, the rubber hand illusion, that brings about a bodily misattribution in neurologically unimpaired participants, as somatoparaphrenia does in brain-damaged patients, is briefly discussed.

This study investigated the relationships between numerical and spatial representations by means of a bisection task, exploring the view that the core representation of number meaning is spatially organized as a mental number line. In Experiment nos. 1 (bisection of digit strings) and 2 (bisection of flanked lines) spatial biases towards the larger digit were found to be related only to processing of relative magnitude. Experiment nos. 3 (bisection of an unfilled space) and 4 (bisection of flanked lines/unfilled spaces) aimed at disclosing perceptual, attentional, and numerical constraints on the bias induced by the position of the larger digit. This effect is interpreted in terms of a cognitive illusion of length, whereby a spatial bias compensates for the numerical disparity. This seems to operate in a categorical fashion ("small/large"), and to be congruent with the assumption that relatively large numbers are associated with the right side of a mental representational space.

Both a neuropsychological syndrome (unilateral spatial neglect) and a visual illusion of length (the Brentano version of the Müller-Lyer illusion) bring about a misjudgement of the subjective centre of a horizontal line, with a unilateral shift. In experiment 1 we investigated, in patients with left unilateral neglect, illusory effects of horizontal length, with the aim of exploring the functional and neural basis of horizontal space perception, and the role of visual processing in shaping the patients' bisection performance. Fourteen right-brain-damaged patients with left spatial unilateral neglect, seven with and seven without left visual half-field deficits (assessed by confrontation, perimetry, and visual event-related potentials), entered this study. Two conditions of manual line bisection were assessed: setting the mid-point of a horizontal line, and of the shaft of the Brentano-Müller-Lyer illusion, with either a left- or a right-sided expansion. Both groups of patients set the subjective midpoint to the right of the objective centre of the line, consistent with the presence of left neglect. Patients with neglect and left hemianopia showed no illusory effects and a greater bisection error. The effects of the illusion, by contrast, were fully present in neglect patients without hemianopia, in both illusory conditions, adding to, or subtracting from, the rightward bisection bias. Anatomoclinical correlations revealed an association of damage to the occipital regions with the lack of illusory effects. Conversely, more anterior damage, sparing these regions, did not disrupt the illusion, revealing a dissociation between visual and spatial processing of extension. These findings suggest that processing of the Müller-Lyer illusion of length is likely to occur in the occipital cortex, at a retinotopic level of representation. In neglect patients with left homonymous hemianopia the visual deficit adds to the spatial bias, yielding a greater error in line bisection, but not in other visual exploratory tasks, such as cancellation, where the contribution of retinotopic frames is likely to be comparatively minor. Experiment 2 showed preserved illusory effects in patients with homonymous visual field defects without spatial unilateral neglect, suggesting that preserved spatial processing may compensate for unilateral visual field defects.

The spatial location of an object can be represented in the brain with respect to different classes of reference frames, either relative to or independent of the subject's position. We used functional magnetic resonance imaging to identify regions of the healthy human brain subserving mainly egocentric or allocentric (object-based) coordinates by asking subjects to judge the location of a visual stimulus with respect to either their body or an object. A color-judgement task, matched for stimuli, difficulty, motor and oculomotor responses, was used as a control. We identified a bilateral, though mainly right-hemisphere based, fronto-parietal network involved in egocentric processing. A subset of these regions, including a much less extensive unilateral, right fronto-parietal network, was found to be active during object-based processing. The right-hemisphere lateralization and the partial superposition of the egocentric and the object-based networks is discussed in the light of neuropsychological findings in brain-damaged patients with unilateral spatial neglect and of neurophysiological studies in the monkey.

Spatial orientation is based on coordinates referring to the subject's body. A fundamental principle is the mid-sagittal plane, which divides the body and space into the left and right sides. Its neural bases were investigated by functional magnetic resonance imaging (fMRI). Seven normal subjects pressed a button when a vertical bar, moving horizontally, crossed the subjective mid-sagittal plane. In the control condition, the subjects' task was to press a button when the direction of the bar movement changed, at the end of each leftward or rightward movement. The task involving the computation of the mid-sagittal plane yielded increased signal in posterior parietal and lateral frontal premotor regions, with a more extensive activation in the right cerebral hemisphere. This direct evidence in normal human subjects that a bilateral, mainly right hemisphere-based, cortical network is active during the computation of the egocentric reference is consistent with neuropsychological studies in patients with unilateral cerebral lesions. Damage to the right hemisphere, more frequently to the posterior-inferior parietal region, may bring about a neglect syndrome of the contralesional, left side of space, including a major rightward displacement of the subjective mid-sagittal plane. The existence of a posterior parietal-lateral premotor frontal network concerned with egocentric spatial reference frames is also in line with neurophysiological studies in the monkey.

Right brain-damaged patients with left visuospatial neglect were required to bisect a line placed in front of them in two different body positions (upright and supine) and two different light conditions (light and dark). The neglect patients, unlike right brain-damaged patients without neglect, strongly reduced their rightward directional error in the supine compared with the upright position. No systematic changes were produced by the light-dark manipulation. The present result cannot be explained with an attentional interpretation of hemispatial neglect. We suggest that the present data provide further evidence that hemineglect is the consequence of a mismatch between different afferent information integrated into an egocentric space representation. According to this model, the presence of a lateralized brain lesion produces asymmetries in some intermediate spatial representations (eye-head, head-trunk, body-environment) but not in the retinotopic one. Any experimental manipulation that reduces the asymmetry of the intermediate representation such as the reduction of gravitational inputs may improve the dynamic integration of the egocentric coordinates.

The cerebral representation of space depends on the integration of many different sensory inputs. The vestibular system provides one such input and its dysfunction can cause profound spatial disorientation. Using positron emission tomography (PET), we measured regional cerebral perfusion with various vestibular stimulations to map central vestibular projections and to investigate the cerebral basis of spatial disorientation. We showed that the temporoparietal cortex, the insula, the putamen, and the anterior cingulate cortex are the cerebral projections of the vestibular system in man and that the spatial disorientation caused by unilateral vestibular stimulation is associated with their asymmetric activation.