Ilaria Bufalari

The Motor Somatotopy of Speech Perception

Listening to speech recruits a network of fronto-temporo-parietal cortical areas. Classical models consider anterior (motor) sites to be involved in speech production whereas posterior sites are considered to be involved in comprehension. This functional segregation is challenged by action-perception theories suggesting that brain circuits for speech articulation and speech perception are functionally dependent. Although recent data show that speech listening elicits motor activities analogous to production, its still debated whether motor circuits play a causal contribution to the perception of speech. Here we administered transcranial magnetic stimulation (TMS) to motor cortex controlling lips and tongue during the discrimination of lip- and tongue-articulated phonemes. We found a neurofunctional double dissociation in speech sound discrimination, supporting the idea that motor structures provide a specific functional contribution to the perception of speech sounds. Moreover, our findings show a fine-grained motor somatotopy for speech comprehension. We discuss our results in light of a modified motor theory of speech perception according to which speech comprehension is grounded in motor circuits not exclusively involved in speech production.

Stimulus-driven modulation of motor-evoked potentials during observation of others' pain

Empathy may allow interindividual sharing not only of emotions (e.g., joy, sadness, disgust) but also of sensations (e.g., touch, itching, pain). Although empathy for pain may rely upon both sensory and affective components of the pain experience, neuroimaging studies indicate that only the affective component of the pain matrix is involved in empathy for pain. By using transcranial magnetic stimulation (TMS), we highlighted the sensorimotor side of empathy for pain by showing a clear motor inhibition during the mere observation of needles penetrating body parts of a human model. Here, we explored stimulus-specific and instruction-specific influences on this inhibition by manipulating task instructions (request to adopt first- or third-person perspective vs. passive observation) and painfulness of the experimental stimuli (presentation of videos of needles deeply penetrating or simply pinpricking a hand). We found a significant reduction in amplitudes of motor-evoked potentials (MEPs) specific to the muscle the subjects observed being penetrated that correlated with the intensity of the pain attributed to the model. Crucially, this motor inhibition was present during observation of penetrating but not of pinpricking needles. Moreover, no MEPs modulation contingent upon different task instructions was found. Results suggest that the motor inhibition elicited by the observation of flesh and bone pain stimuli is more stimulus-driven than instruction-driven.

My face in yours: Visuo-tactile facial stimulation influences sense of identity

Abstract Self-face recognition is crucial for sense of identity and self-awareness. Finding self-face recognition disorders mainly in neurological and psychiatric diseases suggests that modifying sense of identity in a simple, rapid way remains a “holy grail” for cognitive neuroscience. By touching the face of subjects who were viewing simultaneous touches on a partners face, we induced a novel illusion of personal identity that we call “enfacement”: The partners facial features became incorporated into the representation of the participants own face. Subjects reported that morphed images of themselves and their partner contained more self than other only after synchronous, but not asynchronous, stroking. Therefore, we modified self-face recognition by means of a simple psychophysical manipulation. While accommodating gradual change in ones own face is an important form of representational plasticity that may help maintaining identity over time, the surprisingly rapid changes induced by our procedure suggest that sense of facial identity may be more malleable than previously believed. “Enfacement” correlated positively with the participants empathic traits and with the physical attractiveness the participants attributed to their partners. Thus, personality variables modulate enfacement, which may represent a marker of the tendency to be social and may be absent in subjects with defective empathy.

The pain of a model in the personality of an onlooker: influence of state- reactivity and personality traits on embodied empathy for pain

The study of inter-individual differences at behavioural and neural levels represents a new avenue for neuroscience. The response to socio-emotional stimuli varies greatly across individuals. For example, identification with the feelings of a movie character may be total for some people or virtually absent for others. Inter-individual differences may reflect both the on-line effect (state) of the observed stimuli and more stable personal characteristics (trait). Here we show that somatomotor mirror responses when viewing others pain are modulated by both state- and trait-differences in empathy. We recorded motor-evoked potentials (MEPs) induced by Transcranial Magnetic Stimulation (TMS) in healthy individuals observing needles penetrating a models hand. We found a reduction of corticospinal excitability that was specific for the muscle that subjects observed being penetrated. This inhibition correlated with sensory qualities of the pain ascribed to the model. Moreover, it was greater in subjects with high trait-cognitive empathy and lower in subjects with high trait-personal distress and in those with high aversion for the observed movies. Results indicate that somatomotor responses to others pain are influenced by specific onlookers personality traits and self-oriented emotional reactions. Our findings suggest that multiple distinct mechanisms shape mirror mapping of others pain.

Tongue corticospinal modulation during attended verbal stimuli: Priming and coarticulation effects

Humans perceive continuous speech through interruptions or brief noise bursts cancelling entire phonemes. This robust phenomenon has been classically associated with mechanisms of perceptual restoration. In parallel, recent experimental evidence suggests that the motor system may actively participate in speech perception, even contributing to phoneme discrimination. In the present study we intended to verify if the motor system has a specific role in speech perceptual restoration as well. To this aim we recorded tongue corticospinal excitability during phoneme expectation induced by contextual information. Results showed that phoneme expectation determines an involvement of the individuals motor system specifically implicated in the production of the attended phoneme, exactly as it happens during actual listening of that phoneme, suggesting the presence of a speech imagery-like process. Very interestingly, this motoric phoneme expectation is also modulated by subtle coarticulation cues of which the listener is not consciously aware. Present data indicate that the rehearsal of a specific phoneme requires the contribution of the motor system exactly as it happens during the rehearsal of actions executed by the limbs, and that this process is abolished when an incongruent phonemic cue is presented, as similarly occurs during observation of anomalous hand actions. We propose that altogether these effects indicate that during speech listening an attentional-like mechanism driven by the motor system, based on a feed-forward anticipatory mechanism constantly verifying incoming information, is working allowing perceptual restoration.

Vocal Pitch Discrimination in the Motor System.

Speech production can be broadly separated into two distinct components: Phonation and Articulation. These two aspects require the efficient control of several phono-articulatory effectors. Speech is indeed generated by the vibration of the vocal-folds in the larynx (F0) followed by filtering by articulators, to select certain resonant frequencies out of that wave (F1, F2, F3, etc.). Recently it has been demonstrated that the motor representation of articulators (lips and tongue) participates in the discrimination of articulatory sounds (lips- and tongue-related speech sounds). Here we investigate whether the results obtained on articulatory sounds discrimination could be extended to phonation by applying a dual-pulse TMS protocol while subjects had to discriminate F0-shifted vocal utterances [a]. Stimulation over the larynx motor representation, compared to the control site (tongue/lips motor cortex), induced a reduction in RT for stimuli including a subtle pitch shift. We demonstrate that vocal pitch discrimination, in analogy with the articulatory component, requires the contribution of the motor system and that this effect is somatotopically organized.

Enfacing others but only if they are nice to you

Experiencing tactile facial stimulation while seeing synchronous stimuli on the face of another individual induces “enfacement,” i.e., the subjective illusory experience of ownership of the others face (explicit measure) and the attribution of the others facial features to ones own face (implicit measure). Here we expanded previous knowledge by investigating if the tendency to include the other into ones own representation is influenced by positive or negative interpersonal attitudes derived either from consolidated socio-cultural stereotypes or from newly acquired, short-term individual interactions with a specific person. To this aim, we tested in Caucasian white participants the enfacement with a white and a black confederate, before and after an experimental procedure inducing a positive or negative perception of each of them. The results show that the subjective experience of enfacement with in- and out-group others before and after the manipulation is similar. The bias in attributing others facial features to ones own face after synchronous stroking was, instead, dependent on whether the other person was positively perceived, independently of his/her ethnicity. Thus, we show that realistic positive face-to-face interactions are more effective than consolidated racial biases in influencing the strength of self-attribution of another persons facial features in the context of multisensory illusions. Results suggest that positive interpersonal interactions might powerfully change the plasticity of self-other representations.

Interpersonal Multisensory Stimulation reduces the overwhelming distracting power of self-gaze: psychophysical evidence for ‘engazement’

Ones own face and gaze are never seen directly but only in a mirror. Yet, these stimuli capture attention more powerfully than others face and gaze, suggesting the self is special for brain and behavior. Synchronous touches felt on ones own and seen on the face of others induce the sensation of including others in ones own face (enfacement). We demonstrate that enfacement may also reduce the overwhelming distracting power of self-gaze. This effect, hereafter called ‘engazement, depends on the perceived physical attractiveness and inner beauty of the pair partner. Thus, we highlight for the first time the close link between enfacement and engazement by showing that changes of the self-face representation induced by facial visuo-tactile stimulation extend to gaze following, a separate process likely underpinned by different neural substrates. Moreover, although gaze following is a largely automatic, engazement is penetrable to the influence of social variables, such as positive interpersonal perception.

Motor imagery beyond the joint limits: a transcranial magnetic stimulation study.

The processes and neural bases used for motor imagery are also used for the actual execution of correspondent movements. Humans, however, can imagine movements they cannot perform. Here we explored whether plausibility of movements is mapped on the corticospinal motor system and whether the process is influenced by visuomotor vs. kinesthetic-motor first person imagery strategy. Healthy subjects imagined performing possible or biomechanically impossible right index finger movements during single pulse TMS of the left motor cortex. We found an increase of corticospinal excitability during motor imagery which was higher for impossible than possible movements and specific for the muscle involved in the actual execution of the imagined movement. We expand our previous action observation studies, suggesting that the plausibility of a movement is computed in regions upstream the primary motor cortex, and that motor imagery is a higher-order process not fully constrained by the rules that govern motor execution.

Illusory and Veridical Mapping of Tactile Objects in the Primary Somatosensory and Posterior Parietal Cortex

While several behavioral and neuroscience studies have explored visual, auditory, and cross-modal illusions, information about the phenomenology and neural correlates of somatosensory illusions is meager. By combining psychophysics and somatosensory evoked potentials, we explored in healthy humans the neural correlates of 2 compelling tactuo-proprioceptive illusions, namely Aristotle (1 object touching the contact area between 2 crossed fingers is perceived as 2 lateral objects) and Reverse illusions (2 lateral objects are perceived as 1 between crossed-fingers object). These illusions likely occur because of the tactuo-proprioceptive conflict induced by fingers being crossed in a non-natural posture. We found that different regions in the somatosensory stream exhibit different proneness to the illusions. Early electroencephalographic somatosensory activity (at 20 ms) originating in the primary somatosensory cortex (S1) reflects the phenomenal rather than the physical properties of the stimuli. Notably, later activity (around 200 ms) originating in the posterior parietal cortex is higher when subjects resist the illusions. Thus, while S1 activity is related to illusory perception, PPC acts as a conflict resolver that recodes tactile events from somatotopic to spatiotopic frames of reference and ultimately enables veridical perception.