The role of preSMA and STS in face recognition: A transcranial magnetic stimulation (TMS) study.

Current models propose that facial recognition is mediated by two independent yet interacting anatomo-functional systems: one processing facial features mainly mediated by the Fusiform Face Area and the other involved in the extraction of dynamic information from faces, subserved by Superior Temporal Sulcus (STS). Also, the pre-Supplementary Motor Area (pre-SMA) is implicated in facial expression processing as it is involved in its motor mimicry. However, the literature only shows evidence of the implication of STS and preSMA for facial expression recognition, without relating it to face recognition. In addition, the literature shows a facilitatory role of facial motion in the recognition of unfamiliar faces, particularly for poor recognizers. The present study aimed at studying the role of STS and preSMA in unfamiliar face recognition in people with different face recognition skills. 34 healthy participants received repetitive transcranial magnetic stimulation over the right posterior STS, pre-SMA and as sham during a task of matching of faces encoded through: facial expression, rigid head movement or as static (i.e., absence of any facial or head motion). All faces were represented without emotional content. Results indicate that STS has a direct role in recognizing identities through rigid head movement and an indirect role in facial expression processing. This dissociation represents a step forward with respect to current face processing models suggesting that different types of motion involve separate brain and cognitive processes. PreSMA interacts with face recognition skills, increasing the performance of poor recognizers and decreasing that of good recognizers in all presentation conditions. Together, the results suggest the use of at least partially different mechanisms for face recognition in poor and good recognizers and a different role of STS and preSMA in face recognition.

Transcranial magnetic stimulation on the right dorsal attention network modulates the center-surround profile of the attentional focus.

Psychophysical observations indicate that the spatial profile of visuospatial attention includes a central enhancement around the attentional focus, encircled by a narrow zone of reduced excitability in the immediate surround. This inhibitory ring optimally amplifies relevant target information, likely stemming from top-down frontoparietal recurrent activity modulating early visual cortex activations. However, the mechanisms through which neural suppression gives rise to the surrounding attenuation and any potential hemispheric specialization remain unclear. We used transcranial magnetic stimulation to evaluate the role of two regions of the dorsal attention network in the center-surround profile: the frontal eye field and the intraparietal sulcus. Participants performed a psychophysical task that mapped the entire spatial attentional profile, while transcranial magnetic stimulation was delivered either to intraparietal sulcus or frontal eye field on the right (Experiment 1) and left (Experiment 2) hemisphere. Results showed that stimulation of right frontal eye field and right intraparietal sulcus significantly changed the center-surround profile, by widening the inhibitory ring around the attentional focus. The stimulation on the left frontal eye field, but not left intraparietal sulcus, induced a general decrease in performance but did not alter the center-surround profile. Results point to a pivotal role of the right dorsal attention network in orchestrating inhibitory spatial mechanisms required to limit interference by surrounding distractors.

Age-related differences in the statistical learning of target selection and distractor suppression.

In recent years, the use of implicit mechanisms based on statistical learning (SL) has emerged as a strong factor in biasing visuospatial attention, so that target selection is improved at frequently attended locations and distractor filtering is facilitated at frequently suppressed locations. Although these mechanisms have been consistently described in younger adults, similar evidence in healthy aging is scarce. Therefore, we studied the learning and persistence of SL of target selection and distractor suppression in younger and older adults in visual search tasks where the frequency of target (Experiment 1) or distractor (Experiment 2) was biased across spatial locations. The results show that SL of target selection was preserved in the older adults so, similar to their younger counterparts, they showed a strong and persistent advantage in target selection at locations more frequently attended. However, unlike young adults, they did not benefit from implicit SL of distractor suppression, so that distractor interference was maintained throughout the experiment independently of the contingencies associated with distractor locations. Taken together, these results provide novel evidence of distinct developmental patterns for SL of task-relevant and task-irrelevant visual information, likely reflecting differences in the implementation of proactive suppression attentional mechanisms between younger and older adults. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

How to Test the Association Between Baseline Performance Level and the Modulatory Effects of Non-Invasive Brain Stimulation Techniques.

Behavioral effects of non-invasive brain stimulation techniques (NIBS) can dramatically change as a function of different factors (e.g., stimulation intensity, timing of stimulation). In this framework, lately there has been a growing interest toward the importance of considering the inter-individual differences in baseline performance and how they are related with behavioral NIBS effects. However, assessing how baseline performance level is associated with behavioral effects of brain stimulation techniques raises up crucial methodological issues. How can we test whether the performance at baseline is predictive of the effects of NIBS, when NIBS effects themselves are estimated with reference to baseline performance? In this perspective article, we discuss the limitations connected to widely used strategies for the analysis of the association between baseline value and NIBS effects, and review solutions to properly address this type of question.

Exploring the Effects of Brain Stimulation on Musical Taste: tDCS on the Left Dorso-Lateral Prefrontal Cortex-A Null Result.

Humans are the only species capable of experiencing pleasure from esthetic stimuli, such as art and music. Neuroimaging evidence suggests that the left dorsolateral prefrontal cortex (DLPFC) plays a critical role in esthetic judgments, both in music and in visual art. In the last decade, non-invasive brain stimulation (NIBS) has been increasingly employed to shed light on the causal role of different brain regions contributing to esthetic appreciation. In Experiment #1, musician (N = 20) and non-musician (N = 20) participants were required to judge musical stimuli in terms of "liking" and "emotions". No significant differences between groups were found, although musicians were slower than non-musicians in both tasks, likely indicating a more analytic judgment, due to musical expertise. Experiment #2 investigated the putative causal role of the left dorsolateral pre-frontal cortex (DLPFC) in the esthetic appreciation of music, by means of transcranial direct current stimulation (tDCS). Unlike previous findings in visual art, no significant effects of tDCS were found, suggesting that stimulating the left DLPFC is not enough to affect the esthetic appreciation of music, although this conclusion is based on negative evidence,.

"Help! I Need Somebody": Music as a Global Resource for Obtaining Wellbeing Goals in Times of Crisis.

Music can reduce stress and anxiety, enhance positive mood, and facilitate social bonding. However, little is known about the role of music and related personal or cultural (individualistic vs. collectivistic) variables in maintaining wellbeing during times of stress and social isolation as imposed by the COVID-19 crisis. In an online questionnaire, administered in 11 countries (Argentina, Brazil, China, Colombia, Italy, Mexico, the Netherlands, Norway, Spain, the UK, and USA, N = 5,619), participants rated the relevance of wellbeing goals during the pandemic, and the effectiveness of different activities in obtaining these goals. Music was found to be the most effective activity for three out of five wellbeing goals: enjoyment, venting negative emotions, and self-connection. For diversion, music was equally good as entertainment, while it was second best to create a sense of togetherness, after socialization. This result was evident across different countries and gender, with minor effects of age on specific goals, and a clear effect of the importance of music in people's lives. Cultural effects were generally small and surfaced mainly in the use of music to obtain a sense of togetherness. Interestingly, culture moderated the use of negatively valenced and nostalgic music for those higher in distress.

Thematic Reanalysis in the Left Posterior Parietal Sulcus: A TMS Study.

Understanding who does what to whom is at the core of sentence comprehension. The actors that contribute to the verb meaning are labeled thematic roles. We used transcranial magnetic stimulation (TMS) to verify the possible impact of verb semantics on the thematic role encoding process that has been shown to involve the posterior portion of the left posterior parietal sulcus (PPS; Finocchiaro et al., 2015). Sixteen participants underwent TMS and sham stimulation sessions while performing an agent-decision task, in which they had to decide by key press which of the two arguments was the agent of visually presented sentences or pseudo-sentences. The (pseudo)sentences were all reversible and were presented in the active or passive diathesis. Double pulse TMS was delivered to the posterior part of the intraparietal sulcus in an event-related fashion, at two different time windows: 200-400 ms (T1) or 600-800 ms (T2) time-locked to the presentation of the (pseudo)sentence. Results showed that TMS increased accuracy on passive sentences and pseudo-sentences as compared to active sentences and to the baseline, sham condition. Indeed, the presence of a verb with a full semantic representation was not a necessary precondition for the TMS-induced facilitation of passive (pseudo)sentences. Stimulation timing had no effect on accuracy for sentences vs. pseudo-sentences. These observations support the idea that the posterior parietal site is recruited when the correct interpretation of a sentence requires reanalysis of temporarily encoded thematic roles (as in reversible passive sentences) even when the verb is not an entry in the lexicon and hence does not have a semantic representation. Results are consistent with previous evidence and deserve further investigation in larger experimental samples. Increasing the number and variety of stimulus sentences, and administering TMS to additional control sites will be key to further articulate the conclusions allowed by these initial findings.

Modulating the influence of recent trial history on attentional capture via transcranial magnetic stimulation (TMS) of right TPJ.

In visual search, salient yet task-irrelevant distractors in the stimulus array interfere with target selection. This is due to the unwanted shift of attention towards the salient stimulus-the so-called attentional capture effect, which delays deployment of attention onto the target. Although powerful and automatic, attentional capture by a salient distractor is nonetheless antagonized by distractor-filtering mechanisms and is further modulated by cross-trial contingencies: The distractor cost is typically more robust when no distraction has been experienced in the immediate past, compared to when a distractor was present on the immediately preceding trial. Here, we used transcranial magnetic stimulation (TMS) to shed light on the causal role of two crucial nodes of the ventral attention network, namely the Temporo-Parietal Junction (TPJ) and the Middle Frontal Gyrus (MFG), in the exogenous control of attention (i.e., attentional capture) and its history-dependent modulation. Participants were asked to discriminate the direction of a target arrow while ignoring a task-irrelevant salient distractor, when present. Immediately after display onset, 10 Hz triple-pulse TMS was delivered either to TPJ or MFG on the right hemisphere. Results demonstrated that stimulation of right TPJ-but not of right MFG, strongly modulated attentional capture as a function of the type of previous trial, by somewhat enhancing the distractor-related cost when the preceding trial was a distractor-absent trial and significantly decreasing the cost when the preceding trial was a distractor-present trial. These findings indicate that TMS of right TPJ exacerbates the effect of the recent history, likely reflecting enhanced updating of the predictive model that dynamically governs proactive distractor-filtering mechanisms. More generally, the results attest to a role of TPJ in mediating the history-dependent modulation of attentional capture.

The Topography of Visually Guided Grasping in the Premotor Cortex: A Dense-Transcranial Magnetic Stimulation (TMS) Mapping Study.

Visuomotor transformations at the cortical level occur along a network where posterior parietal regions are connected to homologous premotor regions. Grasping-related activity is represented in a diffuse, ventral and dorsal system in the posterior parietal regions, but no systematic causal description of a premotor counterpart of a similar diffuse grasping representation is available. To fill this gap, we measured the kinematics of right finger movements in 17 male and female human participants during grasping of three objects of different sizes. Single-pulse transcranial magnetic stimulation was applied 100 ms after visual presentation of the object over a regular grid of 8 spots covering the left premotor cortex (PMC) and 2 Sham stimulations. Maximum finger aperture during reach was used as the feature to classify object size in different types of classifiers. Classification accuracy was taken as a measure of the efficiency of visuomotor transformations for grasping. Results showed that transcranial magnetic stimulation reduced classification accuracy compared with Sham stimulation when it was applied to 2 spots in the ventral PMC and 1 spot in the medial PMC, corresponding approximately to the ventral PMC and the dorsal portion of the supplementary motor area. Our results indicate a multifocal representation of object geometry for grasping in the PMC that matches the known multifocal parietal maps of grasping representations. Additionally, we confirm that, by applying a uniform spatial sampling procedure, transcranial magnetic stimulation can produce cortical functional maps independent of a priori spatial assumptions.SIGNIFICANCE STATEMENT Visually guided actions activate a large frontoparietal network. Here, we used a dense grid of transcranial magnetic stimulation spots covering the whole premotor cortex (PMC), to identify with accurate spatial mapping the functional specialization of the human PMC during grasping movement. Results corroborate previous findings about the role of the ventral PMC in preshaping the fingers according to the size of the target. Crucially, we found that the medial part of PMC, putatively covering the supplementary motor area, plays a direct role in object grasping. In concert with findings in nonhuman primates, these results indicate a multifocal representation of object geometry for grasping in the PMC and expand our understanding of how our brain integrates visual and motor information to perform visually guided actions.

Instrumental expertise and musical timbre modulate the spatial representation of pitch.

Humans show a tendency to represent pitch in a spatial format. A classical finding supporting this spatial representation is the Spatial-Musical Association of Response Codes (SMARC) effect, reflecting faster responses to low tones when pressing a left/bottom-side key and to high tones when pressing a right/top-side key. Despite available evidence suggesting that the horizontal and vertical SMARC effect may be differently modulated by instrumental expertise and musical timbre, no study has so far directly explored this hypothesis in a unified framework. Here, we investigated this possibility by comparing the performance of professional pianists, professional clarinettists and non-musicians in an implicit timbre judgement task, in both horizontal and vertical response settings. Results showed that instrumental expertise significantly modulates the SMARC effect: whereas in the vertical plane a comparable SMARC effect was observed in all groups, in the horizontal plane the SMARC effect was significantly modulated by the specific instrumental expertise, with pianists showing a stronger pitch-space association compared to clarinettists and non-musicians. Moreover, the influence of pitch along the horizontal dimension was stronger in those pianists who started the instrumental training at a younger age. Results also showed an influence of musical timbre in driving the horizontal, but not the vertical, SMARC effect, with only piano notes inducing a pitch-space association. Taken together, these findings suggest that sensorimotor experience due to instrumental training and musical timbre affect the mental representation of pitch on the horizontal space, whereas the one on the vertical space would be mainly independent from musical practice.

Two Distinct Systems Represent Contralateral and Ipsilateral Sensorimotor Processes in the Human Premotor Cortex: A Dense TMS Mapping Study.

Animal brains contain behaviorally committed representations of the surrounding world, which integrate sensory and motor information. In primates, sensorimotor mechanisms reside in part in the premotor cortex (PM), where sensorimotor neurons are topographically clustered according to functional specialization. Detailed functional cartography of the human PM is still under investigation. We explored the topographic distribution of spatially dependent sensorimotor functions in healthy volunteers performing left or right, hand or foot, responses to visual cues presented in the left or right hemispace, thus combining independently stimulus side, effector side, and effector type. Event-related transcranial magnetic stimulation was applied to single spots of a dense grid of 10 points on the participants' left hemiscalp, covering the whole PM. Results showed: (1) spatially segregated hand and foot representations, (2) focal representations of contralateral cues and movements in the dorsal PM, and (3) distributed representations of ipsilateral cues and movements in the ventral and dorso-medial PM. The present novel causal information indicates that (1) the human PM is somatotopically organized and (2) the left PM contains sensory-motor representations of both hemispaces and of both hemibodies, but the hemispace and hemibody contralateral to the PM are mapped on a distinct, nonoverlapping cortical region compared to the ipsilateral ones.

Probing the Neural Mechanisms for Distractor Filtering and Their History-Contingent Modulation by Means of TMS.

In visual search, the presence of a salient, yet task-irrelevant, distractor in the stimulus array interferes with target selection and slows down performance. Neuroimaging data point to a key role of the frontoparietal dorsal attention network in dealing with visual distractors; however, the respective roles of different nodes within the network and their hemispheric specialization are still unresolved. Here, we used transcranial magnetic stimulation (TMS) to evaluate the causal role of two key regions of the dorsal attention network in resisting attentional capture by a salient singleton distractor: the frontal eye field (FEF) and the cortex within the intraparietal sulcus (IPS). The task of the participants (male/female human volunteers) was to discriminate the pointing direction of a target arrow while ignoring a task-irrelevant salient distractor. Immediately after stimulus onset, triple-pulse 10 Hz TMS was delivered either to IPS or FEF on either side of the brain. Results indicated that TMS over the right FEF significantly reduced the behavioral cost engendered by the salient distractor relative to left FEF stimulation. No such effect was obtained with stimulation of IPS on either side of brain. Interestingly, this FEF-dependent reduction in distractor interference interacted with the contingent trial history, being maximal when no distractor was present on the previous trial relative to when there was one. Our results provide direct causal evidence that the right FEF houses key mechanisms for distractor filtering, pointing to a pivotal role of the frontal cortex of the right hemisphere in limiting interference from an irrelevant but attention-grabbing stimulus.SIGNIFICANCE STATEMENT Visually conspicuous stimuli attract our attention automatically and interfere with performance by diverting resources away from the main task. Here, we applied transcranial magnetic stimulation over four frontoparietal cortex locations (frontal eye field and intraparietal sulcus in each hemisphere) to identify regions of the dorsal attention network that help limit interference from task-irrelevant, salient distractors. Results indicate that the right FEF participates in distractor-filtering mechanisms that are recruited when a distracting stimulus is encountered. Moreover, right FEF implements adjustments in distraction-filtering mechanisms following recent encounters with distractors. Together, these findings indicate a different hemispheric contribution of the left versus right dorsal frontal cortex to distraction filtering. This study expands our understanding of how our brains select relevant targets in the face of task-irrelevant, salient distractors.

A walk on the dark side: TMS over the right inferior frontal gyrus (rIFG) disrupts behavioral responses to infant stimuli.

Infant signals, including infant sounds and facial expressions, play a critical role in eliciting parental proximity and care. Processing of infant signals in the adulthood brain is likely to recruit emotional empathy neural circuits, including the inferior frontal gyrus (IFG). Here, we used transcranial magnetic stimulation (TMS) to test the role of right IFG (rIFG) in behavioral responses to infant signals. Specifically, a group of nulliparous women were asked to perform a handgrip dynamometer task and an Approach-Avoidance Task (AAT) after receiving TMS over the right IFG or over a control site (vertex). Suppressing activity in the rIFG affected the modulation of handgrip force in response to infant crying. Moreover, the AAT showed that participants tend to avoid the sad infant face after Vertex stimulation, and this bias was counteracted by rIFG stimulation. Our results suggest a causal role of rIFG in sensitive responding towards sad infants and point to the rIFG as a critical node in the neural network underlying the innate releasing mechanism for feelings of love, affection and caring of sad infants.

Auditory prediction cues motor preparation in the absence of movements.

There is increasing evidence for integrated representation of sensory and motor information in the brain, and that seeing or hearing action-related stimuli may automatically cue the movements required to respond to or produce them. In this study we tested whether anticipation of tones in a known melody automatically activates corresponding motor representations in a predictive way, in preparation for potential upcoming movements. Therefore, we trained 20 non-musicians (8 men, 12 women) to play a simple melody. Then, while they passively listened to the learned or unlearned melodies, we applied single pulse transcranial magnetic stimulation (TMS) over M1 to measure motor evoked potentials from the associated finger muscle either preceding or following the onset of individual tones. Our results show that listening to the learned melody increased corticospinal excitability for specific finger muscles before tone onset. This demonstrates that predictable auditory information can activate motor representations in an anticipatory muscle-specific manner, even in the absence of intention to move. This suggests that the motor system is involved in the prediction of sensory events, likely based on auditory-parietal-prefrontal feedforward/feedback loops that automatically prepare predictable sound-related actions independent of actual execution and the associated auditory feedback. Overall, we propose that multimodal forward models of upcoming sounds and actions support motor preparation, facilitate error detection and correction, and guide perception.

The Spatial Musical Association of Response Codes does not depend on a normal visual experience: A study with early blind individuals.

Converging evidence suggests that the perception of auditory pitch exhibits a characteristic spatial organization. This pitch-space association can be demonstrated experimentally by the Spatial Musical Association of Response Codes (SMARC) effect. This is characterized by faster response times when a low-positioned key is pressed in response to a low-pitched tone, and a high-positioned key is pressed in response to a high-pitched tone. To investigate whether the development of this pitch-space association is mediated by normal visual experience, we tested a group of early blind individuals on a task that required them to discriminate the timbre of different instrument sounds with varying pitch. Results revealed a comparable pattern in the SMARC effect in both blind participants and sighted controls, suggesting that the lack of prior visual experience does not prevent the development of an association between pitch height and vertical space.

Testing the Role of Dorsal Premotor Cortex in Auditory-Motor Association Learning Using Transcranical Magnetic Stimulation (TMS).

Interactions between the auditory and the motor systems are critical in music as well as in other domains, such as speech. The premotor cortex, specifically the dorsal premotor cortex (dPMC), seems to play a key role in auditory-motor integration, and in mapping the association between a sound and the movement used to produce it. In the present studies we tested the causal role of the dPMC in learning and applying auditory-motor associations using 1 Hz repetitive Transcranical Magnetic Stimulation (rTMS). In this paradigm, non-musicians learn a set of auditory-motor associations through melody training in two contexts: first when the sound to key-press mapping was in a conventional sequential order (low to high tones mapped onto keys from left to right), and then when it was in a novel scrambled order. Participant's ability to match the four pitches to four computer keys was tested before and after the training. In both experiments, the group that received 1 Hz rTMS over the dPMC showed no significant improvement on the pitch-matching task following training, whereas the control group (who received rTMS to visual cortex) did. Moreover, in Experiment 2 where the pitch-key mapping was novel, rTMS over the dPMC also interfered with learning. These findings suggest that rTMS over dPMC disturbs the formation of auditory-motor associations, especially when the association is novel and must be learned rather explicitly. The present results contribute to a better understanding of the role of dPMC in auditory-motor integration, suggesting a critical role of dPMC in learning the link between an action and its associated sound.

Melodic Priming of Motor Sequence Performance: The Role of the Dorsal Premotor Cortex.

The purpose of this study was to determine whether exposure to specific auditory sequences leads to the induction of new motor memories and to investigate the role of the dorsal premotor cortex (dPMC) in this crossmodal learning process. Fifty-two young healthy non-musicians were familiarized with the sound to key-press mapping on a computer keyboard and tested on their baseline motor performance. Each participant received subsequently either continuous theta burst stimulation (cTBS) or sham stimulation over the dPMC and was then asked to remember a 12-note melody without moving. For half of the participants, the contour of the melody memorized was congruent to a subsequently performed, but never practiced, finger movement sequence (Congruent group). For the other half, the melody memorized was incongruent to the subsequent finger movement sequence (Incongruent group). Hearing a congruent melody led to significantly faster performance of a motor sequence immediately thereafter compared to hearing an incongruent melody. In addition, cTBS speeded up motor performance in both groups, possibly by relieving motor consolidation from interference by the declarative melody memorization task. Our findings substantiate recent evidence that exposure to a movement-related tone sequence can induce specific, crossmodal encoding of a movement sequence representation. They further suggest that cTBS over the dPMC may enhance early offline procedural motor skill consolidation in cognitive states where motor consolidation would normally be disturbed by concurrent declarative memory processes. These findings may contribute to a better understanding of auditory-motor system interactions and have implications for the development of new motor rehabilitation approaches using sound and non-invasive brain stimulation as neuromodulatory tools.

A TMS investigation on the role of the cerebellum in pitch and timbre discrimination.

BACKGROUND: Growing neuroimaging and clinical evidence suggests that the cerebellum plays a critical role in perception. In the auditory domain, the cerebellum seems to be important in different aspects of music and sound processing. Here we investigated the possible causal role of the cerebellum in two auditory tasks, a pitch discrimination and a timbre discrimination task. Specifically, participants performed a pitch and a timbre discrimination task prior and after receiving offline low frequency transcranical magnetic stimulation (TMS) over their (right) cerebellum. RESULTS: Suppressing activity in the right cerebellum by means of inhibitory 1 Hz TMS affected participants' ability to discriminate pitch but not timbre. CONCLUSION: These findings point to a causal role of the cerebellum in at least certain aspects of sound processing and are important in a clinical perspective helping understanding the impact of cerebellar lesions on sensory functions.

Grasping the sound: Auditory pitch influences size processing in motor planning.

Growing evidence shows that individuals consistently match auditory pitch with visual size. For instance, high-pitched sounds are perceptually associated with smaller visual stimuli, whereas low-pitched sounds with larger ones. The present study explores whether this crossmodal correspondence, reported so far for perceptual processing, also modulates motor planning. To address this issue, we carried out a series of kinematic experiments to verify whether actions implying size processing are affected by auditory pitch. Experiment 1 showed that grasping movements toward small/large objects were initiated faster in response to high/low pitches, respectively, thus extending previous findings in the literature to more complex motor behavior. Importantly, auditory pitch influenced the relative scaling of the hand preshaping, with high pitches associated with smaller grip aperture compared with low pitches. Notably, no effect of auditory pitch was found in case of pointing movements (no grasp implied, Experiment 2), as well as when auditory pitch was irrelevant to the programming of the grip aperture, that is, in case of grasping an object of uniform size (Experiment 3). Finally, auditory pitch influenced also symbolic manual gestures expressing "small" and "large" concepts (Experiment 4). In sum, our results are novel in revealing the impact of auditory pitch on motor planning when size processing is required, and shed light on the role of auditory information in driving actions.