The peer's point of view: Observing a peer performing an action enhances third-person perspective in adolescents.

Adolescents are particularly prone to peer influence. Since group membership modulates the person's tendency to take someone else's viewpoint, here we decided to investigate whether adolescents are influenced by the presence of a peer when taking another person's perspective. A group of adolescents from upper secondary schools in Naples (Italy) had to observe scenes of an actor (an adolescent or an adult) gazing, grasping, gazing/grasping an object or staying still. When required to judge the spatial location of the object, the adolescent participants adopted the actor's viewpoint (third-person perspective) more frequently when the actor was an adolescent rather than when he was an adult and when the adolescent actor grasped the object. Thus, adolescents seem particularly prone to mentally simulate someone else's actions when the other person is a peer. These findings suggest that modulating motor simulation processes via social environmental factors could influence adolescents' perspective taking.

Path integration in 3D from visual motion cues: A human fMRI study.

While neural correlates of path integration on a yaw plane have been studied extensively, much less is known about path integration in three-dimensions (3D). Here we used fMRI during virtual navigation within tunnels in pseudo-3D. We found that the same visual motion stimuli are encoded differently in the brain depending on whether they represent displacements within the yaw plane or within the pitch plane. The yaw plane is more represented in the hippocampus while the pitch plane is more represented in the angular gyrus (AG) and in the posterior inferior temporal gyrus (pITG), known to be involved in 3D space encoding. In addition, a region in pITG, located just above the previous one, showed two different patterns with multi-voxel analysis, separately coding for the pitch and yaw planes. These results suggest that information encoded within pITG about the yaw plane may be exchanged with the hippocampus, while information about the pitch plane may be exchanged with the AG.

Sound-evoked vestibular stimulation affects the anticipation of gravity effects during visual self-motion.

Humans anticipate the effects of gravity during visually simulated self-motion in the vertical direction. Here we report that an artificial vestibular stimulation consisting of short-tone bursts (STB) suppresses this anticipation. Participants pressed a button upon entering a tunnel during virtual-reality roller coaster rides in downward or forward directions. In different trials, we delivered STB, pulsed white noise (WN), or no sound (NO). In the control conditions (WN, NO), participants responded earlier during downward than forward motion irrespective of true kinematics, consistent with the a priori expectation that downward but not forward motion is accelerated by gravity. STB canceled the difference in response timing between the two directions, without affecting overall task performance. Thus, we argue that vestibular signals play a role in the anticipation of visible gravity effects during self-motion.

Impact of body posture on laterality judgement and explicit recognition tasks performed on self and others' hands.

Judgments on laterality of hand stimuli are faster and more accurate when dealing with one's own than others' hand, i.e. the self-advantage. This advantage seems to be related to activation of a sensorimotor mechanism while implicitly processing one's own hands, but not during explicit one's own hand recognition. Here, we specifically tested the influence of proprioceptive information on the self-hand advantage by manipulating participants' body posture during self and others' hand processing. In Experiment 1, right-handed healthy participants judged laterality of either self or others' hands, whereas in Experiment 2, an explicit recognition of one's own hands was required. In both experiments, the participants performed the task while holding their left or right arm flexed with their hand in direct contact with their chest ("flexed self-touch posture") or with their hand placed on a wooden smooth surface in correspondence with their chest ("flexed proprioceptive-only posture"). In an "extended control posture", both arms were extended and in contact with thighs. In Experiment 1 (hand laterality judgment), we confirmed the self-advantage and demonstrated that it was enhanced when the subjects judged left-hand stimuli at 270° orientation while keeping their left arm in the flexed proprioceptive-only posture. In Experiment 2 (explicit self-hand recognition), instead, we found an advantage for others' hand ("self-disadvantage") independently from posture manipulation. Thus, position-related proprioceptive information from left non-dominant arm can enhance sensorimotor one's own body representation selectively favouring implicit self-hands processing.

Developmental changes of the biomechanical effect in motor imagery.

Motor imagery has been investigated in childhood and early adolescence, but not across adolescence stages; moreover, available evidence did not clarify whether the involvement of motor information in mental rotation of body parts becomes stronger or weaker during development. In the present study, we employed the hand laterality task to assess motor imagery in ninety-seven typically developing adolescents divided into three age groups (i.e., 11-12, 14-15, and 17-18 years); mental rotation of objects and letters were also assessed. As a specific marker of the motor involvement in mental rotation of body parts, we assessed the so-called biomechanical effect, that is, the advantage for judging hand pictures showing physically comfortable positions with respect to hand pictures showing physically impossible or awkward positions. Results demonstrated that the biomechanical effect did not significantly affect early adolescents' performance, whereas it became significant in 14- to 15-year-old participants and even more stronger in 17- to 18-year-old participants; this pattern did not depend on an increase in processing speed to mentally rotate both corporeal and non-corporeal (objects and letters) stimuli. The present findings demonstrated that: (1) motor imagery undergoes a continuous and progressive refinement throughout adolescence, and (2) full exploitation of motor information to mentally transform corporeal stimuli can be attained in late adolescence only.

Self-touch affects motor imagery: a study on posture interference effect.

Several studies showed that mental rotation of body parts is interfered with by manipulation of the subjects' posture. However, the experimental manipulations in such studies, e.g., to hold one arm flexed on one's own chest, activated not only proprioceptive but also self-tactile information. Here, we tested the hypothesis that the combination of self-touch and proprioception is more effective than proprioception alone in interfering with motor imagery. In Experiment 1 right- and left-handers were required to perform the hand laterality task, while holding one arm (right or left) flexed with the hand in direct contact with their chest (self-touch condition, STC) or with the hand placed on a wooden smooth surface in correspondence with their chest (no self-touch condition, NoSTC); in a third neutral condition, subjects kept both arms extended (neutral posture condition, NPC). Right-handers were slower when judging hand laterality in STC with respect to NoSTC and NPC, particularly when the sensory manipulation involved their dominant arm. No posture-related effect was observed in left-handers. In Experiment 2, by applying the same sensory manipulations as above to both arms, we verified that previous results were not due to a conflict between perceived position of the two hands. These data highlighted a complex interaction between body schema and motor imagery, and underlined the role of hand dominance in shaping such interaction.

Sensitivity of occipito-temporal cortex, premotor and Broca's areas to visible speech gestures in a familiar language.

When looking at a speaking person, the analysis of facial kinematics contributes to language discrimination and to the decoding of the time flow of visual speech. To disentangle these two factors, we investigated behavioural and fMRI responses to familiar and unfamiliar languages when observing speech gestures with natural or reversed kinematics. Twenty Italian volunteers viewed silent video-clips of speech shown as recorded (Forward, biological motion) or reversed in time (Backward, non-biological motion), in Italian (familiar language) or Arabic (non-familiar language). fMRI revealed that language (Italian/Arabic) and time-rendering (Forward/Backward) modulated distinct areas in the ventral occipito-temporal cortex, suggesting that visual speech analysis begins in this region, earlier than previously thought. Left premotor ventral (superior subdivision) and dorsal areas were preferentially activated with the familiar language independently of time-rendering, challenging the view that the role of these regions in speech processing is purely articulatory. The left premotor ventral region in the frontal operculum, thought to include part of the Broca's area, responded to the natural familiar language, consistent with the hypothesis of motor simulation of speech gestures.

Numbers are represented in egocentric space: effects of numerical cues and spatial reference frames on hand laterality judgements.

Convergent findings demonstrate that numbers can be represented according to a spatially oriented mental number line. However, it is not established whether a default organization of the mental number line exists (i.e., a left-to-right orientation) or whether its spatial arrangement is only the epiphenomenon of specific task requirements. To address this issue we performed two experiments in which subjects were required to judge laterality of hand stimuli preceded by small, medium or large numerical cues; hand stimuli were compatible with egocentric or allocentric perspectives. We found evidence of a left-to-right number -- hand association in processing stimuli compatible with an egocentric perspective, whereas the reverse mapping was found with hands compatible with an allocentric perspective. These findings demonstrate that the basic left-to-right arrangement of the mental number line is defined with respect to the body-centred egocentric reference frame.

Processing of visual gravitational motion in the peri-sylvian cortex: Evidence from brain-damaged patients.

Rich behavioral evidence indicates that the brain estimates the visual direction and acceleration of gravity quite accurately, and the underlying mechanisms have begun to be unraveled. While the neuroanatomical substrates of gravity direction processing have been studied extensively in brain-damaged patients, to our knowledge no such study exists for the processing of visual gravitational motion. Here we asked 31 stroke patients to intercept a virtual ball moving along the vertical under either natural gravity or artificial reversed gravity. Twenty-seven of them also aligned a luminous bar to the vertical direction (subjective visual vertical, SVV). Using voxel-based lesion-symptom mapping as well as lesion subtraction analysis, we found that lesions mainly centered on the posterior insula are associated with greater deviations of SVV, consistent with several previous studies. Instead, lesions mainly centered on the parietal operculum decrease the ability to discriminate natural from unnatural gravitational acceleration with a timed motor response in the interception task. Both the posterior insula and the parietal operculum belong to the vestibular cortex, and presumably receive multisensory information about the gravity vector. We speculate that an internal model estimating the effects of gravity on visual objects is constructed by transforming the vestibular estimates of mechanical gravity, which are computed in the brainstem and cerebellum, into internalized estimates of virtual gravity, which are stored in the cortical vestibular network. The present lesion data suggest a specific role for the parietal operculum in detecting the mismatch between predictive signals from the internal model and the online visual signals.

Body Constraints on Motor Simulation in Autism Spectrum Disorders.

Developmental data suggested that mental simulation skills become progressively dissociated from overt motor activity across development. Thus, efficient simulation is rather independent from current sensorimotor information. Here, we tested the impact of bodily (sensorimotor) information on simulation skills of adolescents with Autism Spectrum Disorders (ASD). Typically-developing (TD) and ASD participants judged laterality of hand images while keeping one arm flexed on chest or while holding both arms extended. Both groups were able to mentally simulate actions, but this ability was constrained by body posture more in ASD than in TD adolescents. The strong impact of actual body information on motor simulation implies that simulative skills are not fully effective in ASD individuals.

Transcranial Electrical Stimulation over Dorsolateral Prefrontal Cortex Modulates Processing of Social Cognitive and Affective Information.

Recent neurofunctional studies suggested that lateral prefrontal cortex is a domain-general cognitive control area modulating computation of social information. Neuropsychological evidence reported dissociations between cognitive and affective components of social cognition. Here, we tested whether performance on social cognitive and affective tasks can be modulated by transcranial direct current stimulation (tDCS) over dorsolateral prefrontal cortex (DLPFC). To this aim, we compared the effects of tDCS on explicit recognition of emotional facial expressions (affective task), and on one cognitive task assessing the ability to adopt another person's visual perspective. In a randomized, cross-over design, male and female healthy participants performed the two experimental tasks after bi-hemispheric tDCS (sham, left anodal/right cathodal, and right anodal/left cathodal) applied over DLPFC. Results showed that only in male participants explicit recognition of fearful facial expressions was significantly faster after anodal right/cathodal left stimulation with respect to anodal left/cathodal right and sham stimulations. In the visual perspective taking task, instead, anodal right/cathodal left stimulation negatively affected both male and female participants' tendency to adopt another's point of view. These findings demonstrated that concurrent facilitation of right and inhibition of left lateral prefrontal cortex can speed-up males' responses to threatening faces whereas it interferes with the ability to adopt another's viewpoint independently from gender. Thus, stimulation of cognitive control areas can lead to different effects on social cognitive skills depending on the affective vs. cognitive nature of the task, and on the gender-related differences in neural organization of emotion processing.

"Put Myself Into Your Place": Embodied Simulation and Perspective Taking in Autism Spectrum Disorders.

Embodied cognition theories hold that cognitive processes are grounded in bodily states. Embodied processes in autism spectrum disorders (ASD) have classically been investigated in studies on imitation. Several observations suggested that unlike typical individuals who are able of copying the model's actions from the model's position, individuals with ASD tend to reenact the model's actions from their own egocentric perspective. Here, we performed two behavioral experiments to directly test the ability of ASD individuals to adopt another person's point of view. In Experiment 1, participants had to explicitly judge the left/right location of a target object in a scene from their own or the actor's point of view (visual perspective taking task). In Experiment 2, participants had to perform left/right judgments on front-facing or back-facing human body images (own body transformation task). Both tasks can be solved by mentally simulating one's own body motion to imagine oneself transforming into the position of another person (embodied simulation strategy), or by resorting to visual/spatial processes, such as mental object rotation (nonembodied strategy). Results of both experiments showed that individual with ASD solved the tasks mainly relying on a nonembodied strategy, whereas typical controls adopted an embodied strategy. Moreover, in the visual perspective taking task ASD participants had more difficulties than controls in inhibiting other-perspective when directed to keep one's own point of view. These findings suggested that, in social cognitive tasks, individuals with ASD do not resort to embodied simulation and have difficulties in cognitive control over self- and other-perspective.

The role of embodied simulation in mental transformation of whole-body images: evidence from Parkinson's disease.

It has been repeatedly demonstrated that mentally performing an action and mentally transforming body-parts entail simulation of one's own body movements, consistent with predictions of embodied cognition theories. However, the involvement of embodied simulation in mental transformation of whole-body images is still disputed. Here, we assessed own body transformation in Parkinson's disease (PD) patients with symptoms most affecting the left or the right body side. PD patients were required to perform left-right judgments on front-facing or back-facing human figures, and a letter rotation task. Results demonstrated that PD patients were selectively impaired in judging the side of back-facing human figures corresponding to their own most affected side, but performed as well as healthy subjects on mental transformation of front-facing bodies and on letter rotation. These findings demonstrate a parallel impairment between motor and mental simulation mechanisms in PD patients, thus highlighting the specific contribution of embodied cognition to mental transformation of whole-body images.

Brain systems for visual perspective taking and action perception.

Taking another person's viewpoint and making sense of their actions are key processes that guide social behavior. Previous neuroimaging investigations have largely studied these processes separately. The current study used functional magnetic resonance imaging to examine how the brain incorporates another person's viewpoint and actions into visual perspective judgments. Participants made a left-right judgment about the location of a target object from their own (egocentric) or an actor's visual perspective (altercentric). Actor location varied around a table and the actor was either reaching or not reaching for the target object. Analyses examined brain regions engaged in the egocentric and altercentric tasks, brain regions where response magnitude tracked the orientation of the actor in the scene and brain regions sensitive to the action performed by the actor. The blood oxygen level-dependent (BOLD) response in dorsomedial prefrontal cortex (dmPFC) was sensitive to actor orientation in the altercentric task, whereas the response in right inferior frontal gyrus (IFG) was sensitive to actor orientation in the egocentric task. Thus, dmPFC and right IFG may play distinct but complementary roles in visual perspective taking (VPT). Observation of a reaching actor compared to a non-reaching actor yielded activation in lateral occipitotemporal cortex, regardless of task, showing that these regions are sensitive to body posture independent of social context. By considering how an observed actor's location and action influence the neural bases of visual perspective judgments, the current study supports the view that multiple neurocognitive "routes" operate during VPT.

Motor imagery in Asperger syndrome: testing action simulation by the hand laterality task.

Asperger syndrome (AS) is a neurodevelopmental condition within the Autism Spectrum Disorders (ASD) characterized by specific difficulties in social interaction, communication and behavioural control. In recent years, it has been suggested that ASD is related to a dysfunction of action simulation processes, but studies employing imitation or action observation tasks provided mixed results. Here, we addressed action simulation processes in adolescents with AS by means of a motor imagery task, the classical hand laterality task (to decide whether a rotated hand image is left or right); mental rotation of letters was also evaluated. As a specific marker of action simulation in hand rotation, we assessed the so-called biomechanical effect, that is the advantage for judging hand pictures showing physically comfortable versus physically awkward positions. We found the biomechanical effect in typically-developing participants but not in participants with AS. Overall performance on both hand laterality and letter rotation tasks, instead, did not differ in the two groups. These findings demonstrated a specific alteration of motor imagery skills in AS. We suggest that impaired mental simulation and imitation of goal-less movements in ASD could be related to shared cognitive mechanisms.

Observation of another's action but not eye gaze triggers allocentric visual perspective.

In the present paper, we investigated whether observation of bodily cues-that is, hand action and eye gaze-can modulate the onlooker's visual perspective taking. Participants were presented with scenes of an actor gazing at an object (or straight ahead) and grasping an object (or not) in a 2 × 2 factorial design and a control condition with no actor in the scene. In Experiment 1, two groups of subjects were explicitly required to judge the left/right location of the target from their own (egocentric group) or the actor's (allocentric group) point of view, whereas in Experiment 2 participants did not receive any instruction on the point of view to assume. In both experiments, allocentric coding (i.e., the actor's point of view) was triggered when the actor grasped the target, but not when he gazed towards it, or when he adopted a neutral posture. In Experiment 3, we demonstrate that the actor's gaze but not action affected participants' attention orienting. The different effects of others' grasping and eye gaze on observers' behaviour demonstrated that specific bodily cues convey distinctive information about other people's intentions.

Judging hand laterality from my or your point of view: interactions between motor imagery and visual perspective.

Motor imagery tasks (hand laterality judgment) are usually performed with respect to a self-body (egocentric) representation, but manipulations of stimulus features (hand orientation) can induce a shift to other's body (allocentric) reference frame. Visual perspective taking tasks are also performed in self-body perspective but a shift to an allocentric frame can be triggered by manipulations of context features (e.g., another person present in the to-be-judged scene). Combining hand laterality task and visual perspective taking, we demonstrated that both stimulus and context features can modulate motor imagery performance. In Experiment 1, participants judged laterality of a hand embedded in a human or non-human silhouette. Results showed that observing a human silhouette interfered with judgments on "egocentric hand stimuli" (right hand, fingers up). In Experiment 2, participants were explicitly required to judge laterality of a hand embedded in a human silhouette from their own (egocentric group) or from the silhouette's perspective (allocentric group). Consistent with previous results, the egocentric group was significantly faster than the allocentric group in judging fingers-up right hand stimuli. These findings showed that concurrent activation of egocentric and allocentric frames during mental transformation of body parts impairs participants' performance due to a conflict between motor and visual mechanisms.