Beyond peripersonal boundaries: insights from crossmodal interactions.

We experience our self as a body located in space. However, how information about self-location is integrated into multisensory processes underlying the representation of the peripersonal space (PPS), is still unclear. Prior studies showed that the presence of visual information related to oneself modulates the multisensory processes underlying PPS. Here, we used the crossmodal congruency effect (CCE) to test whether this top-down modulation depends on the spatial location of the body-related visual information. Participants responded to tactile events on their bodies while trying to ignore a visual distractor presented on the mirror reflection of their body (Self) either in the peripersonal space (Near) or in the extrapersonal space (Far). We found larger CCE when visual events were presented on the mirror reflection in the peripersonal space, as compared to the extrapersonal space. These results suggest that top-down modulation of the multisensory bodily self is only possible within the PPS.

Brain-heart interactions are modulated across the respiratory cycle via interoceptive attention.

Respiration and heartbeat continuously interact within the living organism at many different levels, representing two of the main oscillatory rhythms of the body and providing major sources of interoceptive information to the brain. Despite the modulatory effect of respiration on exteroception and cognition has been recently established in humans, its role in shaping interoceptive perception has been scarcely investigated so far. In two independent studies, we investigated the effect of spontaneous breathing on cardiac interoception by assessing the Heartbeat Evoked Potential (HEP) in healthy humans. In Study 1, we compared HEP activity for heartbeats occurred during inhalation and exhalation in 40 volunteers at rest. We found higher HEP amplitude during exhalation, compared to inhalation, over fronto-centro-parietal areas. This suggests increased brain-heart interactions and improved cortical processing of the heartbeats during exhalation. Further analyses revealed that this effect was moderated by heart rate changes. In Study 2, we tested the respiratory phase-dependent modulation of HEP activity in 20 volunteers during Exteroceptive and Interoceptive conditions of the Heartbeat Detection (HBD) task. In these conditions, participants were requested to tap at each heartbeat, either listened to or felt, respectively. Results showed higher HEP activity and higher detection accuracy at exhalation than inhalation in the Interoceptive condition only. Direct comparisons of Interoceptive and Exteroceptive conditions confirmed stronger respiratory phase-dependent modulation of HEP and accuracy when attention was directed towards the interoceptive stimuli. Moreover, HEP changes during the Interoceptive condition were independent of heart physiology, but were positively correlated with higher detection accuracy at exhalation than inhalation. This suggests a link between optimization of cortical processing of cardiac signals and detection of heartbeats across the respiratory cycle. Overall, we provide data showing that respiration shapes cardiac interoception at the neurophysiological and behavioural levels. Specifically, exhalation may allow attentional shift towards the internal bodily states.

Oscillatory Properties of Functional Connections Between Sensory Areas Mediate Cross-Modal Illusory Perception.

The presentation of simple auditory stimuli can significantly impact visual processing and even induce visual illusions, such as the auditory-induced double flash illusion (DFI). These cross-modal processes have been shown to be driven by occipital oscillatory activity within the alpha band. Whether this phenomenon is network specific or can be generalized to other sensory interactions remains unknown. The aim of the current study was to test whether cross-modal interactions between somatosensory-to-visual areas leading to the same (but tactile-induced) DFI share similar properties with the auditory DFI. We hypothesized that if the effects are mediated by the oscillatory properties of early visual areas per se, then the two versions of the illusion should be subtended by the same neurophysiological mechanism (i.e., the speed of the alpha frequency). Alternatively, if the oscillatory activity in visual areas predicting this phenomenon is dependent on the specific neural network involved, then it should reflect network-specific oscillatory properties. In line with the latter, results recorded in humans (both sexes) show a network-specific oscillatory profile linking the auditory DFI to occipital alpha oscillations, replicating previous findings, and tactile DFI to occipital beta oscillations, a rhythm typical of somatosensory processes. These frequency-specific effects are observed for visual (but not auditory or somatosensory) areas and account for auditory-visual connectivity in the alpha band and somatosensory-visual connectivity in the beta band. We conclude that task-dependent visual oscillations reflect network-specific oscillatory properties favoring optimal directional neural communication timing for sensory binding.SIGNIFICANCE STATEMENT We investigated the oscillatory correlates of the auditory- and tactile-induced double flash illusion (DFI), a phenomenon where two interleaved beeps (taps) set within 100 ms apart and paired with one visual flash induce the sensation of a second illusory flash. Results confirm previous evidence that the speed of individual occipital alpha oscillations predict the temporal window of the auditory-induced illusion. Importantly, they provide novel evidence that the tactile-induced DFI is instead mediated by the speed of individual occipital beta oscillations. These task-dependent occipital oscillations are shown to be mediated by the oscillatory properties of the neural network engaged in the task to favor optimal temporal integration between the senses.

Individual Alpha Frequency Predicts Perceived Visuotactile Simultaneity.

Temporal encoding is a key feature in multisensory processing that leads to the integration versus segregation of perceived events over time. Whether or not two events presented at different offsets are perceived as simultaneous varies widely across the general population. Such tolerance to temporal delays is known as the temporal binding window (TBW). It has been recently suggested that individual oscillatory alpha frequency (IAF) peak may represent the electrophysiological correlate of TBW, with IAF also showing a wide variability in the general population (8-12 Hz). In our work, we directly tested this hypothesis by measuring each individual's TBW during a visuotactile simultaneity judgment task while concurrently recording their electrophysiological activity. We found that the individual's TBW significantly correlated with their left parietal IAF, such that faster IAF accounted for narrower TBW. Furthermore, we found that higher prestimulus alpha power measured over the same left parietal regions accounted for more veridical responses of non-simultaneity, which may be explained either by accuracy in perceptual simultaneity or, alternatively, in line with recent proposals by a shift in response bias from more conservative (high alpha power) to more liberal (low alpha power). We propose that the length of an alpha cycle constrains the temporal resolution within which perceptual processes take place.

Altered temporal variance and functional connectivity of BOLD signal is associated with state anxiety during acute systemic inflammation.

Systemic inflammation is accompanied by complex behavioral changes and disturbed emotion regulation that have been related to the pathophysiology of mood disorders including depression and anxiety. However, the causal role of systemic inflammation on mood disorders is still unclear. We herein investigated neural resting state patterns of temporal variance of the amygdala and functional connectivity within the salience network underlying changes in state anxiety during experimentally-induced systemic inflammation. In this randomized, double-blind study, N = 43 healthy men received an intravenous injection of either low-dose lipopolysaccharide (LPS, 0.4 ng/kg body weight) or saline. Resting state functional magnetic resonance imaging was assessed before and 3.5 h after injection. State anxiety, assessed with a standardized questionnaire, and plasma cytokine concentrations were repeatedly measured. LPS administration induced a transient systemic inflammatory response reflected in increases in plasma Interleukin (IL)-6 and Tumor Necrosis Factor (TNF)-α concentration. Compared to placebo, state anxiety and temporal variance in the amygdala significantly increased while functional connectivity in the salience network decreased during LPS-induced systemic inflammation. Together, these data indicate that acute systemic inflammation alters temporal variance of the BOLD signal as well as functional connectivity in brain regions and networks implicated in emotion processing and regulation. These results are of translational importance to encourage further research on the role of inflammatory pathways in the pathophysiology of neuropsychiatric conditions including anxiety disorders.

How action performance affects object perception.

How deep is the linkage between action and perception? Much is known about how object perception impacts on action performance, much less about how action performance affects object perception. Does action performance affect perceptual judgment on object features such as shape and orientation? Answering these questions was the aim of the present study. Participants were asked to reach and grasp a handled mug without any visual feedback before judging whether a visually presented mug was handled or not. Performing repeatedly a grasping action resulted in a perceptual categorization aftereffect as measured by a slowdown in the judgment on a handled mug. We suggest that what people are doing may impact on their perceptual judgments on the surrounding things.

Multisensory integration, body representation and hyperactivity of the immune system.

Multisensory stimuli are integrated over a delimited window of temporal asynchronies. This window is highly variable across individuals, but the origins of this variability are still not clear. We hypothesized that immune system functioning could partially account for this variability. In two experiments, we investigated the relationship between key aspects of multisensory integration in allergic participants and healthy controls. First, we tested the temporal constraint of multisensory integration, as measured by the temporal binding window. Second, we tested multisensory body representation, as indexed by the Rubber Hand Illusion (RHI). Results showed that allergic participants have a narrower temporal binding window and are less susceptible to the RHI than healthy controls. Overall, we provide evidence linking multisensory integration processes and the activity of the immune system. The present findings are discussed within the context of the effect of immune molecules on the brain mechanisms enabling multisensory integration and multisensory body representation.

The role of expectation in multisensory body representation - neural evidence.

Sensory events contribute to body ownership, the feeling that the body belongs to me. However, the encoding of sensory events is not only reactive, but also proactive in that our brain generates prediction about forthcoming stimuli. In previous studies, we have shown that prediction of sensory events is a sufficient condition to induce the sense of body ownership. In this study, we investigated the underlying neural mechanisms. Participants were seated with their right arm resting upon a table just below another smaller table. Hence, the real hand was hidden from the participant's view and a life-sized rubber model of a right hand was placed on the small table in front of them. Participants observed a wooden plank while approaching - without touching - the rubber hand. We measured the phenomenology of the illusion by means of questionnaire. Neural activity was recorded by means of near-infrared spectroscopy (fNIRS). Results showed higher activation of multisensory parietal cortices in the rubber hand illusion induced by touch expectation. Furthermore, such activity was correlated with the subjective feeling of owning the rubber hand. Our results enrich current models of body ownership suggesting that our multisensory brain regions generate prediction on what could be my body and what could not. This finding might have interesting implications in all those cases in which body representation is altered, anorexia, bulimia nervosa and obesity, among others.

Multisensory integration induces body ownership of a handtool, but not any handtool.

Bodily boundaries are computed by integrating multisensory bodily signals and can be experimentally manipulated using bodily illusions. Research on tool use demonstrates that tools alter body representations motorically to account for changes in a user's action repertoire. The present experiment sought to unify perceptual and motoric accounts of tool embodiment using a modified Rubber Hand Illusion (RHI) that also addressed the skill and practice aspects of the tool use literature. In Experiment 1, synchronous multisensory stimulation induced perceptual embodiment of a tool, chopsticks. The embodiment of chopsticks was stronger for more skilled participants, and if the illusion was preceded by tool use. In Experiment 2, the illusion was not elicited with a different type of tool, a teacup, showing that not all objects can be incorporated. This experiment helps to clarify the role of perceptual and motoric embodiment and suggests future avenues for research into tools embodiment using this method.

Intertrial Variability in the Premotor Cortex Accounts for Individual Differences in Peripersonal Space.

We live in a dynamic environment, constantly confronted with approaching objects that we may either avoid or be forced to address. A multisensory and sensorimotor interface, the peripersonal space (PPS), mediates every physical interaction between our body and the environment. Behavioral investigations show high variability in the extension of PPS across individuals, but there is a lack of evidence on the neural underpinnings of these large individual differences. Here, we used approaching auditory stimuli and fMRI to capture the individual boundary of PPS and examine its neural underpinnings. Precisely, we tested the hypothesis that intertrial variability (ITV) in brain regions coding PPS predicts individual differences of its boundary at the behavioral level. Selectively in the premotor cortex, we found that ITV, rather than trial-averaged amplitude, of BOLD responses to far rather than near dynamic stimuli predicts the individual extension of PPS. Our results provide the first empirical support for the relevance of ITV of brain responses for individual differences in human behavior. SIGNIFICANCE STATEMENT: Peripersonal space (PPS) is a multisensory and sensorimotor interface mediating every physical interaction between the body and the environment. A major characteristic of the boundary of PPS in humans is the extremely high variability of its location across individuals. We show that interindividual differences in the extension of the PPS are predicted by variability of BOLD responses in the premotor cortex to far stimuli approaching our body. Our results provide the first empirical support to the relevance of variability of evoked responses for human behavior and its variance across individuals.

Press to grasp: how action dynamics shape object categorization.

Action and object are deeply linked to each other. Not only can viewing an object influence an ongoing action, but motor representations of action can also influence visual categorization of objects. It is tempting to assume that this influence is effector-specific. However, there is indirect evidence suggesting that this influence may be related to the action goal and not just to the effector involved in achieving it. This paper aimed, for the first time, to tackle this issue directly. Participants were asked to categorize different objects in terms of the effector (e.g. hand or foot) typically used to act upon them. The task was delivered before and after a training session in which participants were instructed either just to press a pedal with their foot or to perform the same foot action with the goal of guiding an avatar's hand to grasp a small ball. Results showed that pressing a pedal to grasp a ball influenced how participants correctly identified graspable objects as hand-related ones, making their responses more uncertain than before the training. Just pressing a pedal did not have any similar effect. This is evidence that the influence of action on object categorization can be goal-related rather than effector-specific.

Brain activity modulation during the production of imperative and declarative pointing.

Pointing is a communicative gesture, commonly used for expressing two main intentions: imperative, to obtain a desired object/action from the other, or declarative, to share attention/interest about a referent with the other. Previous neuroimaging research on adults examined pointing almost exclusively as a reaching-like motor act rather than as a communicative gesture. Here, we used fMRI to record brain activity while 16 participants produced either imperative or declarative pointing gestures within a communicative context. A network of regions (the bilateral ventral premotor cortex, anterior midcingulate cortex, middle insula and the right preSMA) showed a preference for the production of declarative pointing as opposed to imperative pointing. The right preSMA also preferred declarative intention during pointing observation. Instead, independently from the intention, the right pMTG was more active during pointing observation than production. In the bilateral posterior parietal reach region we also observed a side (contra>ipsi) effect when the intention was imperative, regardless of the subject's role in the communication. Based on these results, we propose that pointing with declarative intention recruits a network of regions associated with will, motivation, emotional/affective expression and intersubjectivity, whereas pointing with imperative intention recruits regions associated with reaching. The proposal is consistent with the developmental hypothesis that declarative pointing reflects social cognitive abilities more than imperative pointing and establishes a stimulating link for future interdisciplinary research.

The eye in hand: predicting others' behavior by integrating multiple sources of information.

The ability to predict the outcome of other beings' actions confers significant adaptive advantages. Experiments have assessed that human action observation can use multiple information sources, but it is currently unknown how they are integrated and how conflicts between them are resolved. To address this issue, we designed an action observation paradigm requiring the integration of multiple, potentially conflicting sources of evidence about the action target: the actor's gaze direction, hand preshape, and arm trajectory, and their availability and relative uncertainty in time. In two experiments, we analyzed participants' action prediction ability by using eye tracking and behavioral measures. The results show that the information provided by the actor's gaze affected participants' explicit predictions. However, results also show that gaze information was disregarded as soon as information on the actor's hand preshape was available, and this latter information source had widespread effects on participants' prediction ability. Furthermore, as the action unfolded in time, participants relied increasingly more on the arm movement source, showing sensitivity to its increasing informativeness. Therefore, the results suggest that the brain forms a robust estimate of the actor's motor intention by integrating multiple sources of information. However, when informative motor cues such as a preshaped hand with a given grip are available and might help in selecting action targets, people tend to capitalize on such motor cues, thus turning out to be more accurate and fast in inferring the object to be manipulated by the other's hand.

Group membership and social status modulate joint actions.

The ability to form shared task representations is considered a keystone of social cognition. It remains, however, contentious if, and to what extent, social categorization impacts on shared representations. In the present study, we address the possibility of the modulation of action co-representation by social categorization, such as group membership and social status, as indexed by the social Simon effect. Italian participants were requested to perform a social Simon task, along with either an Italian (high-status in-group) or an Albanian (low-status out-group) participant. Results show that Italian participants co-represented the action of their partner when paired with a high-status in-group participant. Conversely, this effect was absent when they performed the task with a low-status out-group participant. Furthermore, the Albanian participants co-represented the action of their partner when paired with an Italian participant. These results suggest that group membership modulates action co-representation through the varying of the groups' relative status. The impact of this issue is boundless given the increasing multicultural nature of our society. Indeed, if multiculturalism fails, modern society does likewise.

Left insular cortex and left SFG underlie prismatic adaptation effects on time perception: evidence from fMRI.

Prismatic adaptation (PA) has been shown to affect left-to-right spatial representations of temporal durations. A leftward aftereffect usually distorts time representation toward an underestimation, while rightward aftereffect usually results in an overestimation of temporal durations. Here, we used functional magnetic resonance imaging (fMRI) to study the neural mechanisms that underlie PA effects on time perception. Additionally, we investigated whether the effect of PA on time is transient or stable and, in the case of stability, which cortical areas are responsible of its maintenance. Functional brain images were acquired while participants (n=17) performed a time reproduction task and a control-task before, immediately after and 30 min after PA inducing a leftward aftereffect, administered outside the scanner. The leftward aftereffect induced an underestimation of time intervals that lasted for at least 30 min. The left anterior insula and the left superior frontal gyrus showed increased functional activation immediately after versus before PA in the time versus the control-task, suggesting these brain areas to be involved in the executive spatial manipulation of the representation of time. The left middle frontal gyrus showed an increase of activation after 30 min with respect to before PA. This suggests that this brain region may play a key role in the maintenance of the PA effect over time.

Being an agent or an observer: different spectral dynamics revealed by MEG.

Several neuroimaging studies reported that a common set of regions is recruited during action observation and execution and it has been proposed that the modulation of the µ rhythm, in terms of oscillations in the alpha and beta bands might represent the electrophysiological correlate of the underlying brain mechanisms. However, the specific functional role of these bands within the µ rhythm is still unclear. Here, we used magnetoencephalography (MEG) to analyze the spectral and temporal properties of the alpha and beta bands in healthy subjects during an action observation and execution task. We associated the modulation of the alpha and beta power to a broad action observation network comprising several parieto-frontal areas previously detected in fMRI studies. Of note, we observed a dissociation between alpha and beta bands with a slow-down of beta oscillations compared to alpha during action observation. We hypothesize that this segregation is linked to a different sequence of information processing and we interpret these modulations in terms of internal models (forward and inverse). In fact, these processes showed opposite temporal sequences of occurrence: anterior-posterior during action (both in alpha and beta bands) and roughly posterior-anterior during observation (in the alpha band). The observed differentiation between alpha and beta suggests that these two bands might pursue different functions in the action observation and execution processes.

The eye contact effect in request and emblematic hand gestures.

Request and emblematic gestures, despite being both communicative gestures, do differ in terms of social valence. Indeed, only the former are used to initiate/maintain/terminate an actual interaction. If such a difference is at stake, a relevant social cue, i.e. eye contact, should have different impacts on the neuronal underpinnings of the two types of gesture. We measured blood oxygen level-dependent signals, using functional magnetic resonance imaging, while participants watched videos of an actor, either blindfolded or not, performing emblems, request gestures, or meaningless control movements. A left-lateralized network was more activated by both types of communicative gestures than by meaningless movements, regardless of the accessibility of the actor's eyes. Strikingly, when eye contact was taken into account as a factor, a right-lateralized network was more strongly activated by emblematic gestures performed by the non-blindfolded actor than by those performed by the blindfolded actor. Such modulation possibly reflects the integration of information conveyed by the eyes with the representation of emblems. Conversely, a wider right-lateralized network was more strongly activated by request gestures performed by the blindfolded than by those performed by the non-blindfolded actor. This probably reflects the effect of the conflict between the observed action and its associated contextual information, in which relevant social cues are missing.

When a laser pen becomes a stick: remapping of space by tool-use observation in hemispatial neglect.

The role of active tool use in the remapping of space in hemispatial neglect patients has been extensively investigated. To date, however, there is no evidence that observing tool use can play a role in the remapping of space in hemispatial neglect patients. In this study, a patient with a severe hemispatial neglect in near but not far space and twelve healthy controls were asked to bisect near and far lines using a laser pen. The task was performed both before and immediately after sessions in which they merely observed the experimenter bisecting near and far lines with a stick. During the observation session, participants were either holding an identical stick or empty-handed. Results, in both the neglect patient and healthy controls, showed that observing the experimenter bisecting line while holding the same tool, produces a remapping of the far space into the near space. This result was particularly evident in the neglect patient where observing line-bisection task extended the spatial deficit from the near to the far space. Our results provide new empirical support to the idea that the space around us is not mapped in merely metrical terms, rather it seems to be deeply impacted by both action observation and execution.

The spatial alignment effect in near and far space: a kinematic study.

The present experiment aimed at verifying whether the spatial alignment effect modifies kinematic parameters of pantomimed reaching-grasping of cups located at reachable and not reachable distance. The cup's handle could be oriented either to the right or to the left, thus inducing a grasp movement that could be either congruent or incongruent with the pantomime. The incongruence/congruence induced an increase/decrease in maximal finger aperture, which was observed when the cup was located near but not far from the body. This effect probably depended on influence of the size of the cup body on pantomime control when, in the incongruent condition, cup body was closer to the grasp hand as compared to the handle. Cup distance (near and far) influenced the pantomime even if it was actually executed in the same peripersonal space. Specifically, arm and hand temporal parameters were affected by actual cup distance as well as movement amplitudes. The results indicate that, when executing a reach-to-grasp pantomime, affordance related to the use of the object was instantiated (and in particular the spatial alignment effect became effective), but only when the object could be actually reached. Cup distance (extrinsic object property) influenced affordance, independently of the possibility to actually reach the target.

Social exclusion modulates pre-reflective interpersonal body representation.

Perception of affordance is enhanced not only when that object is located in one's own peripersonal space, as compared to when it is located within extrapersonal space, but also when the object is located in another person's peripersonal space [as measured by a spatial alignment effect (SAE)]. It has been suggested that this reflects the existence of an interpersonal body representation (IBR) that allows us to represent the perceptual states and action possibilities of others. Here, we address the question of whether IBR can be modulated by higher level/reflective social cognition, such as judgments about one's own social status. Participants responded with either the right or the left hand as soon as a go signal appeared. The go signal screen contained a task-irrelevant stimulus consisting of a 3D scene in which a mug with a left- or right-facing handle was positioned on a table. The mug was positioned either inside or outside the reaching space of the participants. In a third of the trials, the mug was positioned within the reaching space of an avatar seated at the table. Prior to this task we induced an experience of social ostracism in half of the participants by means of a standardized social exclusion condition. The results were that the SAE that normally occurs when the mug is in the avatar's reaching space is extinguished by the induced social exclusion. This indicates that judgments about one's own social status modulate the effect of IBR.