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When neurons do not mirror the agent's intentions: Sex differences in neural coding of goal-directed actions

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Abstract

In the current research we used a neurolinguistic paradigm (N400), in which brain activation to semantically congruent and incongruent actions (with respect to the context) was compared in men and women. Twenty-three right-handed students viewed 260 pictures portraying people engaged in simple actions. Perception of plausible and understandable actions (e.g., smiling couple clinking glasses of champagne) was contrasted with that of implausible and unintelligible actions (e.g., businesswoman balancing on one foot in desert). ERP data showed early processing of the action's purpose in the female brain, with a larger parietal N200 to understandable behavior. Source reconstruction (swLORETA) located the neural generators of this effect in the inferior/parietal, left inferior/frontal, left and right premotor areas, right cingulate cortex, right superior/temporal and extra-striate cortex belonging to the so-called "human mirror-neuron system (MNS)". Anterior N400 discriminative response (implausible-plausible) was greater in women than men. The data suggest that congruent/incongruent actions are processed differently from the two sexes, with a prevalence of limbic and cingulate activation in women, and orbito/frontal one in men, along with a right STG activation of comparable amplitude in men and women.

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... These data suggest that action coding was automatically performed and that only skilled players detected the violation of basketball rules. These finding are consistent with previous results on the coding of spontaneous actions (e.g., tool manipulation or goal directed behavior), in contrast with the coding of incoherent and purposeless behavior1013141516. Perception of the latter type of scenes elicited an anterior N400 response, reflecting a difficulty to integrate incoming visual information with sensorimotor related knowledge. ...
... Neural generators (investigated by swLORETA inverse solution) would reveal which brain areas represented ineffective and purposeless actions according to a specific system of sport rules. Indeed, previous ERP literature1013141516 has shown that neural circuits are able to discriminate comprehensible from meaningless actions and that this activity is reflected through a modulation of the N400 response in ERPs. For example, Proverbio and coworkers10 showed that purposeless actions (e.g., a young woman cutting jewelry on a plate with a fork and knife, a man splashing his face with pebbles, or a surgeon dissecting a book) elicited a larger anterior negativity (N400) compared to comprehensible actions (e.g., a woman shopping or doing the laundry). ...
... The mean area amplitude of the N400 response was measured at prefrontal (FP1 FP2), anterior frontal (AF7 AF8), and frontal sites (AFF5h, AFF6h) in the 450–530 ms time window. Both time window and electrode locations are consistent with previous ERP literature on action recognition1013141516. The amplitude of late positivity (LP) was also measured over the same sites in the 900–1000 ms time window. ...
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While the existence of a mirror neuron system (MNS) representing and mirroring simple purposeful actions (such as reaching) is known, neural mechanisms underlying the representation of complex actions (such as ballet, fencing, etc.) that are learned by imitation and exercise are not well understood. In this study, correct and incorrect basketball actions were visually presented to professional basketball players and naïve viewers while their EEG was recorded. The participants had to respond to rare targets (unanimated scenes). No category or group differences were found at perceptual level, ruling out the possibility that correct actions might be more visually familiar. Large, anterior N400 responses of event-related brain potentials to incorrectly performed basketball actions were recorded in skilled brains only. The swLORETA inverse solution for incorrect-correct contrast showed that the automatic detection of action ineffectiveness/incorrectness involved the fronto/parietal MNS, the cerebellum, the extra-striate body area, and the superior temporal sulcus.
... A goal-directed action semantic involves comprehension of the object and its corresponding actions with respect to the context. In previous studies of action semantics, participants were mostly asked to determine the compatibility of the actions in a given situation [1][2][3][4][5][6][7][8][9][10][11][12]. Presentations of conditions violating action-semantics have often involved orientation or functional mismatch to properly execute the functions of the tools [1,2], illogical tool substitution (e.g. ...
... Presentations of conditions violating action-semantics have often involved orientation or functional mismatch to properly execute the functions of the tools [1,2], illogical tool substitution (e.g. cutting bread or playing cello with a saw instead of a bread knife or a bow [3][4][5][6][7]), or inappropriate body movements in a given context (e.g., a woman who's looking at her watch, and carrying a suitcase while walking on a treadmill [4]). Concurrent event-related brain potentials (ERPs) have been analyzed to reveal the neural bases of the cognitive processes of action semantics [1][2][3][4][5][6][7][8][9][10][11][12]. ...
... Presentations of conditions violating action-semantics have often involved orientation or functional mismatch to properly execute the functions of the tools [1,2], illogical tool substitution (e.g. cutting bread or playing cello with a saw instead of a bread knife or a bow [3][4][5][6][7]), or inappropriate body movements in a given context (e.g., a woman who's looking at her watch, and carrying a suitcase while walking on a treadmill [4]). Concurrent event-related brain potentials (ERPs) have been analyzed to reveal the neural bases of the cognitive processes of action semantics [1][2][3][4][5][6][7][8][9][10][11][12]. ...
Article
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Background Action semantics have been investigated in relation to context violation but remain less examined in relation to the meaning of gestures. In the present study, we examined tool–gesture incongruity by event-related potentials (ERPs) and hypothesized that the component N400, a neural index which has been widely used in both linguistic and action semantic congruence, is significant for conditions of incongruence. Methods Twenty participants performed a tool–gesture judgment task, in which they were asked to judge whether the tool–gesture pairs were correct or incorrect, for the purpose of conveying functional expression of the tools. Online electroencephalograms and behavioral performances (the accuracy rate and reaction time) were recorded. ResultsThe ERP analysis showed a left centro-parieto-temporal N300 effect (220–360 ms) for the correct condition. However, the expected N400 (400–550 ms) could not be differentiated between correct/incorrect conditions. After 700 ms, a prominent late negative complex for the correct condition was also found in the left centro-parieto-temporal area. Conclusions The neurophysiological findings indicated that the left centro-parieto-temporal area is the predominant region contributing to neural processing for tool–gesture incongruity in right-handers. The temporal dynamics of tool–gesture incongruity are: (1) firstly enhanced for recognizable tool–gesture using patterns, (2) and require a secondary reanalysis for further examination of the highly complicated visual structures of gestures and tools. The evidence from the tool–gesture incongruity indicated altered brain activities attributable to the N400 in relation to lexical and action semantics. The online interaction between gesture and tool processing provided minimal context violation or anticipation effect, which may explain the missing N400.
... According to the authors, these findings suggest that actions are semantically processed in early and later stages in a similar manner to linguistic stimuli. In a subsequent study, Proverbio et al. (2010) replicated these results and further showed that the N400 for actions could be modulated by gender, with larger amplitudes for women compared to men. Taken together, the reviewed evidence suggests that daily actions elicit a more frontally distributed N400 with a bias, in some cases, toward the right hemisphere (West and Holcomb, 2002; Reid and Striano, 2008). ...
... Based on the scalp-recorded and the intracranial activity, three ERP studies have recently attempted to determine the neural sources of the action-elicited N400 effect. In the first study (Proverbio et al., 2010), source reconstruction using swLORETA (Palmero-Soler et al., 2007 ) located the generators of this effect in the left inferior, left middle, and right superior temporal regions (BA 20, 21) parietal areas (AG, BA 39), frontopolar regions (BA 10), bilateral premotor areas (BA 6), right posterior cingulate cortex, and extrastriate cortex. In the second study (Van Elk et al., 2010a ) the stronger N400 effect for meaningful actions compared to meaningless actions was localized in the left premotor area (BA 6). ...
... As shown by the reviewed studies, the negative activity elicited by action-related anomalous stimuli begins early, approximately at 250–300 ms post-stimulus onset; perhaps reflecting the rapid access that realistic visual images have to semantic memory networks (West and Holcomb, 2002; Sitnikova et al., 2003 Sitnikova et al., , 2008 Mudrik et al., 2010). Nevertheless, other relevant literature, which also includes early components modulation without reporting the N400 (Hauk and Pulvermuller, 2004; Kiefer et al., 2007; Hauk et al., 2008Rorden and Brett, 2000) and the spherical regions of interest (ROIs) (5 mm) displayed in the picture were taken from the MEG, fMRI, ERP, and intracranial studies reviewed in this article (please see Halgren et al., 2002; Proverbio et al., 2010; Ibáñez et al., 2012a). Please note that overlapping activations (in pink) in frontal, temporal, and parietal areas are common to both N400s, while motor and premotor regions are activated only during the processing of action-related material. ...
Article
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Converging neuroscientific evidence suggests the existence of close links between language and sensorimotor cognition. Accordingly, during the comprehension of meaningful actions, our brain would recruit semantic-related operations similar to those associated with the processing of language information. Consistent with this view, electrophysiological findings show that the N400 component, traditionally linked to the semantic processing of linguistic material, can also be elicited by action-related material. This review outlines recent data from N400 studies that examine the understanding of action events. We focus on three specific domains, including everyday action comprehension, co-speech gesture integration, and the semantics involved in motor planning and execution. Based on the reviewed findings, we suggest that both negativities (the N400 and the action-N400) reflect a common neurocognitive mechanism involved in the construction of meaning through the expectancies created by previous experiences and current contextual information. To shed light on how this process is instantiated in the brain, a testable contextual fronto-temporo-parietal model is proposed.
... According to the authors, these findings suggest that actions are semantically processed in early and later stages in a similar manner to linguistic stimuli. In a subsequent study, Proverbio et al. (2010) replicated these results and further showed that the N400 for actions could be modulated by gender, with larger amplitudes for women compared to men. ...
... Based on the scalp-recorded and the intracranial activity, three ERP studies have recently attempted to determine the neural sources of the action-elicited N400 effect. In the first study (Proverbio et al., 2010), source reconstruction using swLORETA (Palmero-Soler et al., 2007) located the generators of this effect in the left inferior, left middle, and right superior temporal regions (BA 20, 21) parietal areas (AG, BA 39), frontopolar regions (BA 10), bilateral premotor areas (BA 6), right posterior cingulate cortex, and extrastriate cortex. In the second study (Van Elk et al., 2010a) the stronger N400 effect for meaningful actions compared to meaningless actions was localized in the left premotor area (BA 6). ...
... Lateral view of the left hemisphere showing the N400 neural sources for words (in blue) and for actions (in red). The figure was computed using the MRIcron software (Rorden and Brett, 2000) and the spherical regions of interest (ROIs) (5 mm) displayed in the picture were taken from the MEG, fMRI, ERP, and intracranial studies reviewed in this article (please see Halgren et al., 2002;Proverbio et al., 2010;Ibáñez et al., 2012a). Please note that overlapping activations (in pink) in frontal, temporal, and parietal areas are common to both N400s, while motor and premotor regions are activated only during the processing of action-related material. ...
... We hypothesized that presenting a verbal description of an emotional or physiological state would activate the conceptual representation of corresponding body language (because of resonating empathic systems), and that the presentation of a picture representing a person actually experiencing the same or totally different feeling would stimulate a congruent (''same'') vs. incongruent (''different'') neural response. Electric neuroimaging literature have identified such a response as a negative deflection peaking at about 400 ms (but generally more anterior than linguistic N400) indexing the automatic detection of an incongruence between incoming visual information about an action being performed, and previous knowledge (about the action's goal, intention, appropriateness, procedure, context of use, etc.): [16,[21][22][23][24][25]. ...
... Previous studies of congruent/incongruent actions (e.g., [23,24,28]) have not reported posterior P3 and LP responses. This discrepancy is most likely due to methodological differences. ...
... Previous ERP literature has revealed which neural circuits are involved in the recognition of purposeful versus purposeless behavior. It is thought that the activities of these circuits are reflected on the modulation of the anterior N400 response [18,[21][22][23]27]. For example, Proverbio & Riva [22] provided evidence of that incongruent actions (e.g., a surgeon dissecting a book) elicit larger anterior negativities (i.e., N400) than do congruent actions (e.g., a woman doing the laundry), especially at inferior frontal sites (F1, F2). ...
Article
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In this study, the neural mechanism subserving the ability to understand people's emotional and mental states by observing their body language (facial expression, body posture and mimics) was investigated in healthy volunteers. ERPs were recorded in 30 Italian University students while they evaluated 280 pictures of highly ecological displays of emotional body language that were acted out by 8 male and female Italian actors. Pictures were briefly flashed and preceded by short verbal descriptions (e.g., "What a bore!") that were incongruent half of the time (e.g., a picture of a very attentive and concentrated person shown after the previous example verbal description). ERP data and source reconstruction indicated that the first recognition of incongruent body language occurred 300 ms post-stimulus. swLORETA performed on the N400 identified the strongest generators of this effect in the right rectal gyrus (BA11) of the ventromedial orbitofrontal cortex, the bilateral uncus (limbic system) and the cingulate cortex, the cortical areas devoted to face and body processing (STS, FFA EBA) and the premotor cortex (BA6), which is involved in action understanding. These results indicate that face and body mimics undergo a prioritized processing that is mostly represented in the affective brain and is rapidly compared with verbal information. This process is likely able to regulate social interactions by providing on-line information about the sincerity and trustfulness of others.
... Cognitive task performance present significant differences between genders (Ingalhalikar et al., 2014;Satterthwaite et al., 2014b). ERP studies using the oddball task as stimuli also documented the gender effect (Karakaş et al., 2006), while using emotional stimuli present even more pronounced gender effect (Arnone et al., 2011;Hung and Cheng, 2014;Proverbio et al., 2010). Inconsistently, the opposing statement supporting that there are no gender differences in the ERP components has been also reported (Fallgatter et al., 1999;Kasai et al., 2002). ...
... Kemmotsu et al. (2012) found a reduction of N400 with aging in left prefrontal area, correlated to slower response time, as well, whereas, Wilkinson et al. (2013) described a reduced N400 to the young group when they performed a task-related shift. Last but not least, genderrelated differences in N400 are also documented (Daltrozzo et al., 2007;Proverbio et al., 2010;Steffensen et al., 2008;Wirth et al., 2007). According to these data semantic analysis and language processing elicit an earlier and larger N400 in women (Daltrozzo et al., 2007;Wirth et al., 2007). ...
... In spite of the existing literature about gender differences in MMN (Fan et al., 2013;Hung and Cheng, 2014;Matsubayashi et al., 2008), P300 (Arnone et al., 2011;Glaser et al., 2012;Huster et al., 2011;Schiff et al., 2008) and N400 (Daltrozzo et al., 2007;Proverbio et al., 2010;Steffensen et al., 2008), no gender-related differences were found in this study, for the latency and amplitude of any of the three investigated ERP sub-components, for the simple auditory oddball paradigm (see Table 2 and Fig. 3). This inconsistency to the literature maybe is task-related. ...
Article
The localization of neuronal generators during an ERP study, using a high-density electroencephalogram (HD-EEG) equipment was made on three Evoked Related Potential (ERP) components, i.e., the Mismatch Negativity (MMN), the P300 and the N400. Furthermore, the ERP characteristics, their field distribution and the area of their maximum field intensity were extracted and compared between young and elderly, as well as between females and males. A two tone oddball experiment was conducted, involving 27 young adults and 18 elderly, healthy and right handed, and HD-EEG data were acquired. These data were then subjected to auditory ERPs extraction and thorough statistical analysis. The derived experimental results revealed significant age-related differences to both the latencies and the amplitudes of the MMN and the P300 and the topographic distribution of the HD-EEG amplitudes. Additionally, a shift in the maximum intensities from frontal to temporal lobe with aging appeared in the case of the P300, whereas no effect was observed for the MMN component. No statistical significant differences (p>0.05) due to age was found in N400 characteristics. Finally, gender-related differences were significant in the response time of the subjects, finding males response faster. The level and the location of the maximum intensity of sources also differed between genders, especially in young subjects. These findings justify the enhanced potential of HD-EEG data to accurately reflect the age and gender dependencies at the three components of simple auditory ERPs and pave the way for the investigation of neurodegenerative pathologies, such as the Alzheimer’s disease.
... Finally, it should be pointed out how several studies strongly suggest that females activate motor-related areas more than males during the processing of social stimuli (including faces, bodies and body parts). Indeed, the activation of the mirror neuron system is greater in females than in males during several tasks, such as the observation of leg and hand movements (Cheng et al. 2006(Cheng et al. , 2007(Cheng et al. , 2008, the observation of body parts in both painful and non-painful situations (Yang et al. 2009), the observation of affective scenes depicting humans (Proverbio et al. 2009(Proverbio et al. , 2010, the mental rotation of hands (Seurinck et al. 2004), and emotional perspective taking (Derntl et al. 2010;Schulte-Rüther et al. 2008). Compared to males, females also exhibit more pronounced and more congruent facial electromyographic reactions to observed facial expressions (Dimberg and Lundqvist 1990;Lundqvist 1995;Sonnby-Borgström et al. 2008), greater embodiment during spatial perspective taking (Kessler and Wang 2012), and better action and emotion recognition from point-light figures (Alaerts et al. 2011). ...
... Past research suggests that females make use of embodied strategies more than males during the processing of social stimuli such as bodies and body parts (e.g., Cheng et al. 2006Cheng et al. , 2007Cheng et al. , 2008Proverbio et al. 2009Proverbio et al. , 2010Seurinck et al. 2004;Yang et al. 2009) and thus we assessed whether the effects of posture were stronger in females than in males. Moreover, given the hypothesized greater contribution of motor representations in right-handers than in left-handers during action simulation (Conson et al. 2011;Gentilucci et al. 1998;Ionta and Blanke 2009;Nì Choisdealbha et al. 2011;Marzoli et al. 2011aMarzoli et al. , b, 2013b, we chose to examine only right-handed participants (a decision legitimized by our interest in the effects of posture rather than in differences between right-and left-handers). ...
... This account would be in line with several embodied cognition studies indicating that proprioceptive information arising from body posture, facial expression and movements facilitates congruent responses (e.g., Bach and Tipper 2007;Duclos et al. 1989;Förster 2003;Marzoli et al. 2013a;Schubert and Koole 2009). Moreover, the sex difference observed is consistent with the greater use of embodied strategies by females than by males during the processing of bodies and body parts (e.g., Cheng et al. 2006Cheng et al. , 2007Cheng et al. , 2008Proverbio et al. 2009Proverbio et al. , 2010Seurinck et al. 2004;Yang et al. 2009; see also Marzoli et al. 2011b). The fact that hand shape did not modulate the effects of posture suggests that, compared with shaping the hand behind the back as an open hand, shaping the hand as a fist did not increase substantially the degree of muscle contraction (in this regard, it should be pointed out that the experimenter instructed participants to shape their hand as a fist without soliciting them to clench their fist vigorously). ...
Article
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In a series of previous studies, we found that when participants were required to imagine another person performing a manual action, they imagined a significantly higher proportion of actions performed with their dominant rather than non-dominant hand, which indicates that shared motor representations between the self and the other are involved also during the imagination of others' actions. Interestingly, the activation of lateralized body-specific motor representations (as indexed by the congruence between the participant's handedness and the imagined person's handedness) appeared to be affected by the visual perspective adopted and participants' handedness. Given that past literature indicates that incongruent or unnatural postures interfere with motor imagery, we tested 480 right-handed participants to investigate whether subjects holding their right hand behind their back would have imagined right-handed actions less frequently than those holding their left hand behind their back. Moreover, we examined the effects of participant's sex, action category (simple or complex) and hand shape (open or fist). Contrary to our prediction, female participants holding their right hand behind their back imagined right-handed actions more frequently than those holding their left hand behind their back, whereas no significant effect was observed in male participants. We propose that the muscle contraction needed to keep a hand behind the back could activate the motor representations of that hand so as to increase the likelihood of imagining an action performed with the corresponding hand. Moreover, the sex difference observed is consistent with the greater use of embodied strategies by females than by males.
... It has also provided critical information regarding the time courses of action coding. For example, a N400 response was previously exhibited during the observation of actions that ended unexpectedly (Reid and Striano, 2008) or in an anomalous way (Sitnikova et al., 2003(Sitnikova et al., , 2008, actions that violated semantic world-knowledge or those that are incomprehensible/implausible (Proverbio et al., 2010). Proverbio et al. (2012) identified a larger N400 in professional basketball players (compared with nonplayers) during the observation of incorrect/ineffective (compared with correct) game actions, which suggests a role of learned motor skills in automatically perceiving a violation of the rules of the game (Proverbio et al., 2012). ...
... These features of the stimuli, in addition to the implicit nature of the task, may explain the centro-parietal topographic distribution identified for the N400 response. Despite substantial evidence showing a more frontal than parietal distributed negativity elicited by pictures and videos that display incorrect/incongruent human actions (Amoruso et al., 2013;Proverbio et al., 2010;Reid and Striano, 2008;Sitnikova et al., 2003Sitnikova et al., , 2008, there are few studies showing a centro-parietal distribution of N400 (Amoruso et al., 2014;Bach et al., 2009;Shibata et al., 2009), in accordance with our finding. In the present study, it is possible to hypothesize that a resonance response involved, to a greater extent, more posterior (visuomotor and somatosensory) than anterior (motor) regions in the brains of the dancers. ...
... A strong activity was also found in the left uncus (BA 28), the forepart of the parahippocampal gyrus (PHG), and the right posterior cingulate cortex (PCC, BA 23). A modulation of these regions (i.e., PHG and CC) was previously shown on the base of motor expertise level with the displayed body actions (Calvo-Merino et al., 2005;Cross et al., 2006;Kim et al., 2011;Petrini et al., 2014), the observation strategy (Decety et al., 1997), and the required task (Suchan et al., 2008), as well as indexing an emotional response (Proverbio et al., 2010; to the perception of an incongruence (e.g., the image of a girl ironing her hair). Moreover, several areas of the AON were active during the observation of deviance, such as the fusiform gyrus (FG,BA 19,20/37), the inferior temporal gyrus (ITG, BA 20) and the middle temporal gyrus (MTG, BA 21). ...
Article
Motor resonance processes that occur when observing an individual perform an action may be modulated by acquired visuomotor expertise. We used the event-related potential (EEG/ERP) technique to investigate the ability to automatically recognize a subtle difference between very similar novel contemporary dance movements. Twelve professional dancers and twelve non-dancers were shown 212 pairs of videos of complex whole-body movements that lasted 3seconds. The second of each pair was the repetition of the previous movement or a slight variation of it (deviance). The participants were engaged in a secondary attentional task. Modulation of a larger centro-parietal N400 effect and a reduction of the Late Positivity amplitude (repetition suppression effect) were identified in response to deviant stimuli only in the dancers. Source reconstruction (swLORETA) showed activations in biological motion, body and face processing related areas, and fronto-parietal and limbic systems. The current findings provide evidence that acquired dance expertise modifies the ability to visually code whole-body complex movements.
... correlation has opposite direction for men compared to women; Gur et al., 2004), sex may impact on links between N200 and symptom severity. Indeed sex differences are seen in the relationship between N200 and goal directed behaviour in the general population, thought to reflect OFC-medial temporal activity (Proverbio et al., 2010). ...
... Given that these regions have been shown to contribute to the composite N200 and P300, such volumetric differences could be reflected in the current findings. Indeed, sex differences are also seen in associations between N200 amplitude and goal-directed behaviour, supposedly underpinned by these frontal regions, in the general population (Proverbio et al., 2010). Such studies, and others that show sex differences in neurocognitive strategies (Sumich et al., 2012a), are also in line with the presently reported sex differences in the association between excitement and anterior ERP amplitudes. ...
... Investigating the socio-perceptual processes preceding helping behavior in children could help to disambiguate conflicting findings in this domain. Systematic gender differences in social perception have been observed in studies with adult participants (Proverbio et al., 2008(Proverbio et al., , 2010, and also in studies that include children in their scope . These reports found that female participants show greater brain activation for social stimuli vs. non-social stimuli compared to male participants of different ages. ...
... The observed differences between younger boys and girls might, however, be explained within the larger framework of differences between men and women in processing of socially relevant content. This has already been discussed with regard to socially relevant stimuli, such as biological motion or social scenes (Proverbio et al., 2008 and processing of purposeful actions in particular (Proverbio et al., 2010). It could be that girls allocate a higher amount of attention to the socially relevant content of the presented visual stimuli from an early age on, while for boys the relevance of the stimuli' social content increases gradually with age. ...
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The exploratory study presented here examines children's ability to recognize another person's need-of-help. This social perception process necessarily precedes the decision to actively help others. Fifty-eight children aged between 5 and 13 completed three experimental paradigms. They were asked to look at black-and-white drawings and to indicate which ones showed somebody in need of help. A control task requiring children to differentiate between pictures of humans and birds measured general categorization abilities. This experimental design enabled us to consider confounding effects of children's developmental status and motivation and to distinguish them from specific need-of-help recognition abilities. As gender and age have been shown to influence social perception as well as helping behavior, we explored whether these factors also have an impact on need-of-help recognition. Children's response accuracies and response times (RTs) were analyzed. We observed clearly higher accuracy rates for younger girls compared to younger boys specifically in the need-of-help recognition tasks. For boys, an age-related performance improvement was found. Younger girls performed at a similarly high level as older girls and boys. No gender differences were observed for children aged over nine. This report provides first evidence that the developmental trajectory of children's ability to recognize another person's need-of-help differs for girls and boys.
... In some cases tool action (Lewis, Brefczynski, Phinney, Janik, & DeYoe, 2005), everyday activities (Koehler et al., 2012), or communicative gesture (Andric & Small, 2012) were considered. In other cases complex actions were represented, but the focus of analysis was toward the relationship between object and context of action production (Proverbio, Riva, & Zani, 2010) or between functional or instrumental features of object (Balconi & Caldiroli, 2011;Sitnikova, Holcomb, Kiyonaga, & Kuperberg, 2008;Van Elk, van Schie, & Bekkering, 2010). ...
... The present results are also consistent with previous studies that showed how an object-related action may implicate the activation of the posterior areas in the case of a semantic task (i.e., representation of congruence of gesture-object relationship; Balconi & Caldiroli, 2011;Balconi, Canavesio, & Vitaloni 2014;Ganis & Kutas, 2003;Sitnikova et al., 2008). It was observed that a specific processing of identification of an object may implicate the activation of the temporal and parietal lobe, in addition to the anterior sites (Proverbio et al., 2010). Indeed this cortical area could be implicated in action of visual scene schemata stored (Ganis & Kutas, 2003), where the semantic coherence object-action-context is crucial to induce an object use representation. ...
Article
The authors explored cortical correlates of action execution and observation, directly comparing control condition condition and execution-observation, using functional near-infrared spectroscopy. Transitive actions (meaningful gestures produced in presence of an object) or intransitive actions (meaningful gestures produced in absence of an object) were performed. Increased oxygenated hemoglobin levels were revealed for both action execution and action observation in premotor cortex, and sensorimotor cortex compared to control condition. However, a higher activity in motor areas was observed for action execution than motor observation. In contrast the posterior parietal cortex was similarly activated in case of both execution and observation task. Finally, it was shown that action execution and observation of transitive more than intransitive gestures was supported by similar parietal posterior areas. These findings support the hypothesis of a partial common network for observation and execution of action, and significant implications related to action types (transitive vs. intransitive).
... The p-value associated with mediation are for the path "c-c " or "a*b." be more engaged in interpreting social interactions (Proverbio et al., 2010), an attribute related to their superiority in empathy (Baron-Cohen and Wheelwright, 2004). It has been posited that the mirror-neuron system, involving the premotor and inferior parietal cortex, provides the basic mechanism for social cognition (Rizzolatti and Craighero, 2004;Cross et al., 2009). ...
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Social interaction is critical to emotional well-being. Previous studies have suggested sex differences in the perception of social interaction. However, the findings depend on the nature of interactions and whether it involves facial emotions. Here, we explored sex differences in neural responses to the perception of social interaction using the Human Connectome Project data. Participants (n = 969, 505 women) were engaged in a social cognition task with geometric objects moving and colliding to simulate social interaction. Behaviorally, men relative to women demonstrated higher accuracy in perceiving social vs. random interactions. Men vs. women showed higher activation in the right superior temporal gyrus, bilateral occipital and posterior cingulate cortex and precuneus, and women vs. men showed higher activation in the right inferior frontal cortex, during exposure to social vs. random interactions. In whole-brain regressions, the differences in accuracy rate in identifying social vs. random interactions (AR SOC - AR RAN ) were associated with higher activation in the paracentral lobule (PCL) and lower activation in bilateral anterior insula (AI), pre-supplementary motor area (preSMA), and left middle frontal gyrus (MFG) in men and women combined, lower activation in bilateral AI, preSMA and left MFG in men alone, and higher activation in the PCL and the medial orbitofrontal cortex in women alone. The latter sex differences were confirmed by slope tests. Further, the PCL activity mediated the correlation between an internalizing syndromal score, as assessed by the Achenbach Self-Report, and (AR SOC - AR RAN ) across all subjects. These findings highlighted sex differences in the behavioral and neural processes underlying the perception of social interaction, as well as the influence of internalizing traits on these processes.
... The N400 is a negative component that appears around 400 ms after the presentation of semantically unrelated information between two words or between a context and a word. Although this component was first studied in the linguistic field, recent studies have extended previous results to richer action sequences and pictorial stimuli (sometimes called N350 or N400-like), such as congruent-incongruent pictures or videos of gestures, actions and motor events (Aravena et al., 2010;Cornejo et al., 2009;Proverbio et al., 2010;Ibañez et al., 2010bIbañez et al., , 2011Guerra et al., 2009;Sitnikova et al., 2003). Although spatial resolution provided by ERP does not allow a precise localization of N400 neural generators, evidence from lesion studies, MEG and intracranial recordings converge to implicate temporal areas (left superior/middle temporal gyrus, the anterior-medial temporal lobe, the PHC and anterior fusiform gyrus) as the possible sources of N400 (Van Petten and Luka, 2006). ...
... The N400 component, a large negative deflection peaking at about 400 ms, is usually largest at central and parietal channels and is sensitive to semantic integration in the verbal domain (Brown & Hagoort, 1993) as well as to lexical decision processes (Federmeier, 2007;Kutas & Federmeier, 2000;Kutas & Hillyard, 1980;Lau, Phillips, & Poeppel, 2008). Recent works have indicated that the N400 does not specifically reflect a linguistic processing, but rather contextual comprehension and the integration process of incoming input with the world knowledge learned over the lifetime (Proverbio & Riva, 2009;Proverbio, Riva, & Zani, 2010;Proverbio, Crotti, Manfredi, Adorni, & Zani, 2012;Sitnikova, Holcomb, Kiyonaga, & Kuperberg, 2008;Giglio, Minati, & Boggio, 2013). Also, it has been showed that the N400 elicited by pictures has a more anterior scalp topography than the N400 observed in most language studies (McPherson & Holcomb, 1999). ...
... Using voxelbased morphometry analysis, concurrent with the dispositional empathy measures, researchers also found that young adult females had significantly larger gray matter volume locatged in the mirror-neuron system, which is highly related in empathy ability [28]. Females showed earlier and stronger brain processing of the action's purpose in the females brain compared with males in an ERP study [29]. Females were also more accurate when their feelings of the target [30]. ...
Article
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Previous studies have demonstrated that the brain responds differentially to others' gains and losses relative to one's own, moderated by social context factors such as competition and interpersonal relationships. In the current study, we tested the hypothesis that the neural response to others' outcomes could be modulated by a short-term induced affective preference. We engaged 17 men and 18 women in a social-exchange game, in which two confederates played fairly or unfairly. Both men and women rated the fair player as likable and the unfair players as unlikable. Afterwards, ERPs were recorded while participants observed each confederates playing a gambling game individually. This study examines feedback related negativity (FRN), an ERP component sensitive to negative feedback. ANOVA showed a significant interaction in which females but not males displayed stronger FRNs when observing likable players' outcomes compared to unlikable ones'. However, males did not respond differently under either circumstance. These findings suggest that, at least in females, the neural response is influenced by a short-term induced affective preference.
... The N400 component, a large negative deflection peaking at about 400 ms, is usually largest at central and parietal channels and is sensitive to semantic integration in the verbal domain (Brown & Hagoort, 1993) as well as to lexical decision processes (Federmeier, 2007;Kutas & Federmeier, 2000;Kutas & Hillyard, 1980;Lau, Phillips, & Poeppel, 2008). Recent works have indicated that the N400 does not specifically reflect a linguistic processing, but rather contextual comprehension and the integration process of incoming input with the world knowledge learned over the lifetime (Proverbio & Riva, 2009;Proverbio, Riva, & Zani, 2010;Proverbio, Crotti, Manfredi, Adorni, & Zani, 2012;Sitnikova, Holcomb, Kiyonaga, & Kuperberg, 2008;Giglio, Minati, & Boggio, 2013). Also, it has been showed that the N400 elicited by pictures has a more anterior scalp topography than the N400 observed in most language studies (McPherson & Holcomb, 1999). ...
Article
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The goal of the present study was to shed some light on a particular kind of humour, called slapstick, by measuring brain bioelectrical activity during the perception of funny vs. non funny pictures involving misfortunate circumstances. According to our hypothesis, the element mostly providing a comic feature in a misfortunate situation is the facial expression of the victims: the observer's reaction will usually be laughing only if the victims will show a funny bewilderment face and not a painful or anger expression. Several coloured photos depicting people involved in misfortunate situations were presented to 30 Italian healthy volunteers, while their EEG was recorded. Three different situations were considered: people showing a painful or an angry expression (affective); people showing a bewilderment expression and, so, a comic look (Comic); people engaged in similar misfortunate situations but with no face visible (No Face). Results showed that the mean amplitude of both the posterior N170 and anterior N220 components were much larger in amplitude to comic pictures, than the other stimuli. This early response could be considered the first identification of a comic element and the evidence of the compelling and automatic response that usually characterizes people amused reaction during a misfortune. In addition, we observed a larger P300 amplitude in response to comic than affective pictures, probably reflecting a more conscious processing of the comic element. Finally, no face pictures elicited an anteriorly distributed N400, which might reflect the effort to comprehend the nature of the situation displayed without any affective facial information, and a late positivity, possibly indexing a re-analysis processing of the unintelligible misfortunate situation (comic or unhappy) depicted in the No Face stimuli. These data support the hypothesis that the facial expression of the victims acts as a specific trigger for the amused feeling that observers usually experience when someone falls down. Overall, the data indicate the existence of a neural circuit that is capable to recognize and appreciate the comic element of a misfortunate situation in a group of young adults.
... Several late ERP components have been related to the processing of visual actions. Modulations of the N400 component by the congruency between an action and the context in which it takes place (e.g., squeezing a lemon in the bathroom instead of the kitchen) have been repeatedly reported [24][25][26][27][28][29][30] . Such modulations have been interpreted as a marker of the integration between different pieces of action information 30 . ...
Article
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Recent behavioural evidence suggests that when processing others’ actions, motor acts and goal-related information both contribute to action recognition. Yet the neuronal mechanisms underlying the dynamic integration of the two action dimensions remain unclear. This study aims to elucidate the ERP components underlying the processing and integration of grip and goal-related information. The electrophysiological activity of 28 adults was recorded during the processing of object-directed action photographs (e.g., writing with pencil) containing either grip violations (e.g. upright pencil grasped with atypical-grip), goal violations (e.g., upside-down pencil grasped with typical-grip), both grip and goal violations (e.g., upside-down pencil grasped with atypical-grip), or no violations. Participants judged whether actions were overall typical or not according to object typical use. Brain activity was sensitive to the congruency between grip and goal information on the N400, reflecting the semantic integration between the two dimensions. On earlier components, brain activity was affected by grip and goal typicality independently. Critically, goal typicality but not grip typicality affected brain activity on the N300, supporting an earlier role of goal-related representations in action recognition. Findings provide new insights on the neural temporal dynamics of the integration of motor acts and goal-related information during the processing of others’ actions.
... For example, the stimuli in the communicative condition appear more humorously than those of the private condition. Second of all, the gender difference (Proverbio et al., 2010) in processing the intention cannot be considered adequately here due to the gender proportion of participants. Further studies need to be done to address these questions to better understand the modulation of communicative vs. private intention, as a function of gender differences and arousal or emotional content. ...
Article
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Although a growing body of research provides critical information about the spatio-temporal dynamic of the brain network mediating the understanding of causality between an action and its outcome at the individual level, little remains known about this cognitive process when the action outcome has a social connotation. To address this question, we recorded electrical brain activity from 16 healthy adults while they performed an intention understanding task including actions with three different types of causality; 1) private intentions by two agents acting independently from one another; 2) communicative intentions by two agents acting in an interactive way with one another; 3) physical causality among objects. Electrophysiological results showed differential electrical activity for private compared to communicative intentions within 400 ms post-stimulus. Brain source localization of the difference waves between communicative and private actions showed a generator located in the vicinity of middle cingulate cortex, which reinforces the role of this brain area in predicting social intentions.
... In particular, the frontal N400 is implicated in the evaluation of the appropriateness of different human actions, such as the meaning of hand postures (26), appropriateness of tool use (27), and semantic anticipation of action sequences (28). Further the frontal N400 has been source-localized in regions encompassing a mentalizing neural network (i.e., inferior frontal cortex, superior frontal gyrus, superior temporal gyrus) (28,29). One conjecture might be that, whereas both Americans and Chinese are equally likely to detect discrepancies between an observed behavior and the behavior normatively expected (as revealed in the centro-parietal N400), only Chinese go beyond the detected norm violation to infer the mental state of the person violating the norms. ...
Article
Humans are unique among all species in their ability to develop and enforce social norms, but there is wide variation in the strength of social norms across human societies. Despite this fundamental aspect of human nature, there has been surprisingly little research on how social norm violations are detected at the neurobiological level. Building on the emerging field of cultural neuroscience, we combine noninvasive electroencephalography (EEG) with a new social norm violation paradigm to examine the neural mechanisms underlying the detection of norm violations and how they vary across cultures. EEG recordings from Chinese and US participants (n = 50) showed consistent negative deflection of event-related potential around 400 ms (N400) over the central and parietal regions that served as a culture-general neural marker of detecting norm violations. The N400 at the frontal and temporal regions, however, was only observed among Chinese but not US participants, illustrating culture-specific neural substrates of the detection of norm violations. Further, the frontal N400 predicted a variety of behavioral and attitudinal measurements related to the strength of social norms that have been found at the national and state levels, including higher culture superiority and self-control but lower creativity. There were no cultural differences in the N400 induced by semantic violation, suggesting a unique cultural influence on social norm violation detection. In all, these findings provided the first evidence, to our knowledge, for the neurobiological foundations of social norm violation detection and its variation across cultures.
... Therefore, N400 activity can be considered as an electrophysiological indicator of incongruency detection and modulated by expertise, reflecting the construction of action meaning from previous experiences and current contextual information (Amoruso et al., 2013). The neural sources of action-N400 activity involve widely distributed semantic and motor regions (Proverbio et al., 2010(Proverbio et al., , 2012van Elk et al., 2010;Amoruso et al., 2014;Orlandi et al., 2017). In particular, the activation of the superior temporal gyrus and middle temporal gyrus is modulated by an individual's expertise, reflecting action processing in the ventral pathway (Amoruso et al., 2014). ...
Article
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Congruency monitoring of action occurs in individuals with relevant motor experience while observing actions. However, it remains unclear whether congruency monitoring can occur at the motor level and the extent to which expertise contributes. Here, we examined behavioral performance and electrophysiological brain activity of individuals with and without domain-specific expertise when judging the action congruency of occluded video clips of a snowboard halfpipe trick and normal walking. For the halfpipe trick, experts exhibited better task performance and greater midline theta oscillations before possible incongruency compared with controls. Source reconstruction for the theta oscillation revealed stronger activation in the middle and superior frontal gyrus for experts in response to incongruency compared with controls. Incongruent halfpipe actions elicited higher N400 amplitude in experts compared with congruent actions, while no such differences were observed in controls. Source reconstruction revealed the activation in the board frontal regions and middle temporal gyrus for experts. These findings suggest that congruency monitoring can occur at the motor level during action observations, and is modulated by individual expertise. The modulation of expertise reflects in the special N400 effect and midline theta oscillation.
... This area could support semantic anomaly representation for an object-related action inserted into sequential frames. This increased N400 effect for incongruent conditions was found in previous studies (Amoruso et al. 2013;Balconi and Caldiroli 2011;Proverbio et al. 2010;Sitnikova et al. 2008). In another study, the DLPFC was found to be responsive to action semantic anomalies tested by brain oscillations (Balconi et al. 2014). ...
Conference Paper
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In this experimental we investigated the cortical neuromodulation during a tDCS (transcranial direct current stimulation) section to induce a temporary inhibition of the frontal area. The induced effect of brain modulation was tested on the EEG (electroencephalography) and ERPs (event-related potentials) profile when subjects performed a task in which they had to respond if the object represented in the sequence was correctly or incorrectly used. It was shown that an increased negative peak deflection (N400) is observable in case of semantic anomalies. We attended a significant reduction of this ERPs deflections when tDCS was applied to frontal area. During the detection task, participants were asked to evaluate the semantic correctness of some motor sequences that manipulated simple objects. EEG were registered during the tDCS or no tDCS stimulation. Significant differences between the two conditions and a reduction of the peak amplitude were observed in case of tDCS stimulation.
... For instance, in our previous study, we found the larger N1 for moral pictures than non-moral pictures, which may reflect the early moral intuition processing without the emotional impact. For mental states attribution, Proverbio and Riva, 2009;Proverbio et al., 2010) reported an enhanced posterior negative component that peaked around 250 ms (N250), which reflected the early recognition of comprehensible behaviors and the early processing of action's purpose. Another study further suggested that the N250 may reflect the early cognitive processing to understand private intention (Wang et al., 2012). ...
Article
The ability to integrate the moral intention information with the outcome of an action plays a crucial role in mature moral judgment. Functional magnetic resonance imaging (fMRI) studies implicated that both prefrontal and temporo-parietal cortices are involved in moral intention and outcome processing. Here, we used the event-related potentials (ERPs) technique to investigate the temporal dynamics of the processing of the integration between intention and outcome information in harmful and helpful moral judgment. In two experiments, participants were asked to make moral judgments for agents who produced either negative/neutral outcomes with harmful/neutral intentions (harmful judgment) or positive/neutral outcomes with helpful/neutral intentions (helpful judgment). Significant ERP differences between attempted and successful actions over prefrontal and bilateral temporo-parietal regions were found in both harmful and helpful moral judgment, which suggest a possible time course of the integration processing in the brain, starting from the right temporo-parietal area (N180) to the left temporo-parietal area (N250), then the prefrontal area (FSW) and the right temporo-parietal area (TP450 and TPSW) again. These results highlighted the fast moral intuition reaction and the late integration processing over the right temporo-parietal area.
... In fact, sex differences have been shown in action understanding tasks. Women have been found to be better at understanding the action purpose, as indexed by earlier and larger discriminative ERP responses to incongruent and purposeless behavior (Proverbio et al., 2010c), compared with men. Consistently, the combined fMRI and ERP study by Canessa et al. (2012) and Proverbio et al. (2011c) found differences across male and female participants involving a stronger activation of the action understanding system, the STS, and the ventral premotor cortex (associated with the mirror resonance of others' actions) during the observation of cooperative (vs. ...
Article
Several studies have demonstrated that women show a greater interest for social information and empathic attitude than men. This article reviews studies on sex differences in the brain, with particular reference to how males and females process faces and facial expressions, social interactions, pain of others, infant faces, faces in things (pareidolia phenomenon), opposite-sex faces, humans vs. landscapes, incongruent behavior, motor actions, biological motion, erotic pictures, and emotional information. Sex differences in oxytocin-based attachment response and emotional memory are also mentioned. In addition, we investigated how 400 different human faces were evaluated for arousal and valence dimensions by a group of healthy male and female University students. Stimuli were carefully balanced for sensory and perceptual characteristics, age, facial expression, and sex. As a whole, women judged all human faces as more positive and more arousing than men. Furthermore, they showed a preference for the faces of children and the elderly in the arousal evaluation. Regardless of face aesthetics, age, or facial expression, women rated human faces higher than men. The preference for opposite- vs. same-sex faces strongly interacted with facial age. Overall, both women and men exhibited differences in facial processing that could be interpreted in the light of evolutionary psychobiology.
... Consistent with this idea, available evidence on the neural mechanisms of action observation points to the existence of a distinct network of brain regions subserving these processes. Previous studies of action-related N400/N450 using source reconstruction techniques have identified the potential origins of these ERP components in regions including the superior temporal (BA 20/BA 21), lateral parietal (BA 39), cingulate, and premotor areas (BA 6), among others (Proverbio, Riva, & Zani, 2010;van Elk, van Schie, & Bekkering, 2010). Notably, these regions partially overlap with those identified as the possible generators of the LP effect linked to observation of dynamic actions, as discussed above . ...
Article
Despite evidence identifying the role of group membership in social cognition, the neural mechanisms associated with the perception and evaluation of nonverbal behaviors displayed by in-group versus out-group members remain unclear. Here, 42 White participants underwent electroencephalographic recording while observing social encounters involving dynamic displays of nonverbal behaviors by racial in-group and out-group avatar characters. Dynamic behaviors included approach and avoidance poses and expressions, followed by the participants' ratings of the avatars displaying them. Behaviorally, participants showed longer RTs when evaluating in-group approach behavior compared with other behaviors, possibly suggesting increased interest and attention devoted to processing positive social encounters with their in-group members. Analyses of ERPs revealed differential sensitivity of the N450 and late positivity components to social cues, with the former showing initial sensitivity to the presence of a humanoid avatar character at the beginning of social encounters and the latter showing sensitivity to dynamic nonverbal behaviors displayed by the avatars. Moreover, time–frequency analysis of electroencephalography data also identified suppression of beta-range power linked to the observation of dynamic nonverbal behaviors. Notably, the magnitude of these responses was modulated by the degree of behavioral racial in-group bias. This suggests that differential neural sensitivity to nonverbal cues while observing social encounters is associated with subsequent in-group bias manifested in the evaluation of such encounters. Collectively, these findings shed light on the mechanisms of racial in-group bias in social cognition and have implications for understanding factors related to successful interactions with individuals from diverse racial backgrounds.
... However, other investigations have revealed that the right STG is not only sensitive to facial expression processing but may also be involved in the processing of emotions from other sources, such as vocal emotion (Lavan and Lima 2014), body language (Proverbio et al. 2014), action comprehension (Proverbio et al. 2010(Proverbio et al. , 2011(Proverbio et al. , 2012, and emotional audiovisual information (Jeong et al. 2011). In addition, a recent investigation by Wyk et al. (2016) analyzed activity in the STS region as participants viewed actions that were congruent or incongruent with the intentions established by a previous actress's facial expression. ...
Article
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The superior temporal gyrus (STG) has been found to play a crucial role in the recognition of actions and facial expressions and may, therefore, be critical for the processing of humorous information. Here we investigated whether tDCS application to the STG would modulate the ability to recognize and appreciate the comic element in serious and comedic situations of misfortune. To this aim, the effects of different types of tDCS stimulation on the STG were analyzed during a task in which the participants were instructed to categorize various misfortunate situations as “comic” or “not comic”. Participants underwent three different tDCS conditions: Anodal-right/Cathodal-left; Cathodal-right/Anodal-left; Sham. Images depicting people involved in accidents were grouped into three categories based on the facial expression of the victim: angry or painful (Affective); bewildered and funny (Comic); and images that did not contain the victim’s face (No Face). An improvement in mean reaction times in response to both the Comic and No Face stimuli was observed following Anodal-left/Cathodal-right stimulation when compared to sham stimulation. This suggests that this stimulation type reduced the reaction times to socio-emotional complex scenes, regardless of facial expression. The Anodal-right/Cathodal-left stimulation reduced the mean reaction times for Comic stimuli only, suggesting that specifically the right STG may be involved in facial expression recognition and in the appreciation of the comic element in misfortunate situations. These results suggest a functional hemispheric asymmetry in STG response to social stimuli: the left STG might have a role in a general comprehension of social complex situations, while the right STG may be involved in the ability to recognize and integrate specific emotional aspects in a complex scene.
... However, other investigations have revealed that the right STG is not only sensitive to facial expression processing but may also be involved in the processing of emotions from other sources, such as vocal emotion (Lavan and Lima 2014), body language (Proverbio et al. 2014), action comprehension (Proverbio et al. 2010(Proverbio et al. , 2011(Proverbio et al. , 2012, and emotional audiovisual information (Jeong et al. 2011). In addition, a recent investigation by Wyk et al. (2016) analyzed activity in the STS region as participants viewed actions that were congruent or incongruent with the intentions established by a previous actress's facial expression. ...
... The N400, of particular interest for the present study, is a negative component peaking around 400 ms after the onset of a meaningful stimulus. It is sensitive not only to semantic and linguistic processing [16][17][18] , but also to communicative gestures 19 , to the perception of images that are incongruent with context and expectations, or to violations of world-knowledge of the observer [20][21][22][23][24] . ...
Article
To date, most investigations in the field of affective neuroscience mainly focused on the processing of facial expressions, overlooking the exploration of emotional body language (EBL), its capability to express our emotions notwithstanding. Few electrophysiological studies investigated the time course and the neural correlates of EBL and the integration of face and body emotion-related information. The aim of the present study was to investigate both the time course and the neural correlates underlying the integration of affective information conveyed by faces and bodies. We analysed EEG activities evoked during an expression matching task, requiring the judgment of emotional congruence between sequentially presented pairs of stimuli belonging to the same category (face-face or body-body), and between stimuli belonging to different categories (face-body or body-face). We focused on N400 time window and results showed that incongruent stimuli elicited a modulation of the N400 in all comparisons except for body-face condition. This modulation was mainly detected in the Middle Temporal Gyrus and within regions related to the mirror mechanism. More specifically, while the perception of incongruent facial expressions activates somatosensory-related representations, incongruent emotional body postures also require the activation of motor and premotor representations, suggesting a strict link between emotion and action.
... Studies using event-related potentials also support the involvement of semantic network in action understanding. Although the N400 was initially described following the onset of incongruent verbal stimuli, it has recently been detected also for incongruent non-verbal stimuli such as actions (Balconi and Caldiroli, 2011;Lee et al., 2018;Proverbio et al., 2010). Incongruent actions, i.e. movements that mismatch to the preceding context, evoke the classic N400 effect (Amoruso et al., 2014;Reid and Striano, 2008;Sitnikova et al., 2003). ...
... Les primates, et donc les humains, sont des créatures hautement sociales. Connaître les intentions des autres (empathie) a de tout temps été crucial pour notre survie(Proverbio et al. 2010). ...
Thesis
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We make decisions every waking day of our life. Facing our options, we tend to pick the most likely to get our expected outcome. Taking into account our past experiences and their outcome is mandatory to identify the best option. This cognitive process is called reinforcement learning. To date, the underlying neural mechanisms are debated. Despite a consensus on the role of dopaminergic neurons in reward processing, several hypotheses on the neural bases of reinforcement learning coexist: either two distinct opposite systems covering cortical and subcortical areas, or a segregation of neurons within brain regions to process reward-based and punishment-avoidance learning.This PhD work aimed to identify the brain dynamics of human reinforcement learning. To unravel the neural mechanisms involved, we used intracerebral recordings in refractory epileptic patients during a probabilistic learning task. In the first study, we used a computational model to tackle the brain dynamics of reinforcement signal encoding, especially the encoding of reward and punishment prediction errors. Local field potentials exhibited the central role of high frequency gamma activity (50-150Hz) in these encodings. We report a role of the ventromedial prefrontal cortex in reward prediction error encoding while the anterior insula and the dorsolateral prefrontal cortex encoded punishment prediction errors. In addition, the magnitude of the neural response in the insula predicted behavioral learning and trial-to-trial behavioral adaptations. These results are consistent with the existence of two distinct opposite cortical systems processing reward and punishments during reinforcement learning. In a second study, we recorded the neural activity of the anterior and dorsomedial nuclei of the thalamus during the same cognitive task. Local field potentials recordings highlighted the role of low frequency theta activity in punishment processing, supporting an implication of these nuclei during punishment-avoidance learning. In a third behavioral study, we investigated the influence of risk on reinforcement learning. We observed a risk-aversion during punishment-avoidance, affecting the performance, as well as a risk-seeking behavior during reward-seeking, revealed by an increased reaction time towards appetitive risky choices. Taken together, these results suggest we are risk-seeking when we have something to gain and risk-averse when we have something to lose, in contrast to the prediction of the prospect theory.Improving our common knowledge of the brain dynamics of human reinforcement learning could improve the understanding of cognitive deficits of neurological patients, but also the decision bias all human beings can exhibit.
... In this vein, a larger N400 and a smaller positivity would index a difficulty in integrating semantic information and in making a decision about gesture/caption congruence. Overall, by manipulating the congruence of pictures and captions, we expected to find an enhanced anterior N400 in response to incorrect associations, which would match previous findings (Sitnikova et al., 2003;Gunter and Bach, 2004;Coulson, 2005, 2007;Reid and Striano, 2008;Proverbio et al., 2010aProverbio et al., , 2012aProverbio et al., ,b, 2015. According to Proverbio and Riva (2009), the N400 would reflect the activation of brain mechanisms that automatically process biological actions as meaningful units by means of visuomotor mirror neurons (Rizzolatti et al., 2001;Fogassi et al., 2005). ...
Article
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The role of facial coding in body language comprehension was investigated by ERP recordings in 31 participants viewing 800 photographs of gestures (iconic, deictic and emblematic), which could be congruent or incongruent with their caption. Facial information was obscured by blurring in half of the stimuli. The task consisted of evaluating picture/caption congruence. Quicker response times were observed in women than in men to congruent stimuli, and a cost for incongruent vs. congruent stimuli was found only in men. Face obscuration did not affect accuracy in women as reflected by omission percentages, nor reduced their cognitive potentials, thus suggesting a better comprehension of face deprived pantomimes. N170 response (modulated by congruity and face presence) peaked later in men than in women. Late Positivity was much larger for congruent stimuli in the female brain, regardless of face blurring. Face presence specifically activated the right superior temporal and fusiform gyri, cingulate cortex and insula, according to source reconstruction. These regions have been reported to be insufficiently activated in face-avoiding individuals with social deficits. Overall, the results corroborate the hypothesis that females might be more resistant to the lack of facial information or better at understanding body language in face-deprived social information.
Article
Objective To design and evaluate the effectiveness of a stimulus material in eliciting the N400 event related potential (ERP). Design A set of 700 semantically congruent and incongruent sentences was developed in accordance with current linguistic norms, and validated with an electroencephalography (EEG) study, in which the influence of age and gender on the N400 ERP magnitude was analysed. Study sample Forty-five normal-hearing subjects (19–57 years, 21 females) participated in the EEG study. Results The stimulus material used in the EEG study elicited a robust N400 ERP, with a morphology consistent with the literature. Results also showed no statistically significant effect of age or gender on the N400 magnitude. Conclusions The material presented in this paper constitutes the largest complete stimulus set suitable for both auditory and text-based N400 experiments. This material may help facilitate the efficient implementation of future N400 ERP studies, as well as promote standardisation and consistency across studies.
Article
Many studies have reported sex differences in empathy and social skills. In this review, several lines of empirical evidences about sex differences in functions and anatomy of social brain are discussed. The most relevant differences involve face processing, facial expression recognition, response to baby schema, the ability to see faces in things, the processing of social interactions, the response to the others’ pain, interest in social information, processing of gestures and actions, biological motion, erotic, and affective stimuli. Sex differences in oxytocin‐based parental response are also reported. In conclusion, the female and male brains show several neuro‐functional differences in various aspects of social cognition, and especially in emotional coding, face processing, and response to baby schema. An interpretation of this sexual dimorphism is provided in the view of evolutionary psychobiology.
Article
Integrative semantic processing and mirror-neuron system activation in response to incongruence detection during action observation have so far been investigated only through paradigms that involve unfeasible action endings, introduction of extraneous objects or synthetic pictures. The ecological validity of the reported effects, particularly modulation of the N400 event-related potential (ERP), therefore remains questionable. To address this shortcoming, we investigated the ERPs evoked by physically feasible and naturalistic but unexpected action endings. We hypothesized that in comparison to existing work our more realistic scenarios would elicit slower processing, due to engagement of deeper analysis as opposed to fast detection based on automatic processes. ERPs were recorded in 14 healthy participants in response to 60 sequences, each containing three pictures, the last of which could be an expected or unexpected action ending. The N400 was larger for the first image compared to the others, and the LPP was enhanced for unexpected endings. Source localization implicated anterior fronto-temporal and temporo-parietal junction regions in generation of these potentials. Our findings challenge the view that the N400 universally indices verbal and action-perception incongruence detection, and point to situation-contingent effects which, in the case of subtle violations, are reflected in later activity components, speculatively reflecting recourse to the "mentalizing" system.
Article
In this study, we explored the representation of an incongruent action (instrumentally incorrect use of an object) in comparison with sentences ending with an incongruent action word, taking into account the role of the activation of the left dorsolateral pFC (DLPFC). This activity was appositely modulated by transcranial direct current stimulation (tDCS). The effect of tDCS when participants processed congruent/incongruent object-related actions (Experiment 1) or sentences (Experiment 2) was verified by measuring changes in the ERP N400, error rates (ERs), and RTs. In Experiment 1, 30 participants performed the detection task within a dynamic context (video tapes representing a sequence of four action frames). In Experiment 2, 28 participants read sentences that represented object-related actions. The stimulation effect (a cathode applied to the DLPFC and an anode to the right supraorbital region) was analyzed by comparing the ER, RT, and ERP profiles before and after stimulation (or sham treatment). A significant reduction of the N400 was observed for incongruent stimuli in the case of cathodal (inhibitory) stimulation of the DLPFC in comparison with prestimulation conditions for Experiment 1, but not Experiment 2. Moreover, ERs were increased, and RTs were reduced in response to incongruent conditions after tDCS, but not after sham stimulation in Experiment 1. It is suggested that perturbation of the DLPFC may limit the ability to analyze a semantically anomalous action sequence as a reduced N400 ERP effect and increased random responses was observed. Finally, the contribution of the frontal area to the semantic processing of actions is discussed.
Article
Background The correct outcome of transitive actions depends on an object’s use, considering both the instrumental and functional features. Indeed, the representation of transitive action includes the correct/incorrect instrumental use and the congruous/incongruous functional use of the object. Objectives. The contribution of frontal (experiment 1) and posterior parietal (experiment 2) areas to process semantic violations in instrumental vs. functional action representation was explored. Methods These cortical areas were appositely modulated by anodal tDCS (transcranial direct current stimulation) or sham tDCS when 43 healthy subjects processed semantic violations (congruous/incongruous pairs) and they had to perform an error detection task (measuring ERs – error rates; and RTs – response times). Results Decreased RTs were observed for incongruous pairs in the case of prefrontal stimulation compared to the sham condition. This effect was related to both the instrumental and functional conditions, suggesting that the activation of the prefrontal area may increase the ability to analyze the semantic incongruence independently from the anomaly type, which would limit the cognitive costs. In contrast, posterior parietal stimulation reduced RTs only in response to functional action in the case of semantic incongruence. Conclusions Thus, different cortical contributions by anterior vs. posterior areas occurred in response to semantic incongruences respectively for generic expectancy violations (more frontally distributed) and specific functional violations (more parietally distributed).
Article
The strength relationship between action and language was largely discussed. In the present research, we explored this link by considering the cortical response (N400 event-related potential effect) to the semantic incongruence induced by a final anomalous object-related action within an actions' sequence. Seventeen participants performed an explicit task to distinguish congruous from incongruous final target action. Event-related potentials analysis showed a significant N400-like effect more frontally (dorsolateral prefrontal cortex) and temporoparietal (mainly left supramarginal gyrus) distribution in response to incongruous condition. It can be argued that the N400-like effect is similar in nature to the N400, which is generally evoked by linguistic stimuli. Nevertheless, the cortical source analysis (low-resolution electromagnetic tomography) showed significant differences for the cortical generators induced by an action processing. This fact may be explained by assuming that object-related action representation activates a specific cortical network, more directly related to congruous/incongruous object use comprehension in relationship with a specific context.
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Given the complexity of ERP recording and processing pipeline, the resulting variability of methodological options, and the potential for these decisions to influence study outcomes, it is important to understand how ERP studies are conducted in practice and to what extent researchers are transparent about their data collection and analysis procedures. The review gives an overview of methodology reporting in a sample of 132 ERP papers, published between January 1980 – June 2018 in journals included in two large databases: Web of Science and PubMed. Because ERP methodology partly depends on the study design, we focused on a well-established component (the N400) in the most commonly assessed population (healthy neurotypical adults), in one of its most common modalities (visual images). The review provides insights into 73 properties of study design, data pre-processing, measurement, statistics, visualization of results, and references to supplemental information across studies within the same subfield. For each of the examined methodological decisions, the degree of consistency, clarity of reporting and deviations from the guidelines for best practice were examined. Overall, the results show that each study had a unique approach to ERP data recording, processing and analysis, and that at least some details were missing from all papers. In the review, we highlight the most common reporting omissions and deviations from established recommendations, as well as areas in which there was the least consistency. Additionally, we provide guidance for a priori selection of the N400 measurement window and electrode locations based on the results of previous studies.
Research
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The sex difference between nature and nurture has been a debate for a long period. As body language play a critical part of our interpersonal communication, understanding it is critical to reveal all the hidden meanings in any message. Therefore, the aim of this research is to find if sex affects body language decoding accuracy. And if true, who is the better decoder? This research finds that women are better body language decoder than men, due to many natural and psychological differences. The natural difference tested by measuring the brain response of both men and women. The results showed a higher response and a larger brain area when subjected to face stimuli. The second argument based on the gender role in the evolutionary psychology theory. The third method argument stands on the sexual selection theory. Some researchers countered these arguments by studying this difference between homosexual and heterosexual people difference, and on the different responses showed in different experimental environments. Refuting this argument was by shedding light on the atypical development of homosexual people, and explaining the competitive environment complexity.
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Abstract While the mechanism of sign language comprehension in deaf people has been widely investigated, little is known about the neural underpinnings of spontaneous gesture comprehension in healthy speakers. Bioelectrical responses to 800 pictures of actors showing common Italian gestures (e.g., emblems, deictic or iconic gestures) were recorded in 14 persons. Stimuli were selected from a wider corpus of 1122 gestures. Half of the pictures were preceded by an incongruent description. ERPs were recorded from 128 sites while participants decided whether the stimulus was congruent. Congruent pictures elicited a posterior P300 followed by late positivity, while incongruent gestures elicited an anterior N400 response. N400 generators were investigated with swLORETA reconstruction. Processing of congruent gestures activated face- and body-related visual areas (e.g., BA19, BA37, BA22), the left angular gyrus, mirror fronto/parietal areas. The incongruent–congruent contrast particularly stimulated linguistic and semantic brain areas, such as the left medial and the superior temporal lobe. ERPs; Action processing; Language; Semantic violation; N400; Body language; Mirror neurons
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The present research used the Iowa Gambling Task to test the effect of the reward-sensitivity Behavioral Activation System-Reward (BAS-Reward) construct on the ability to distinguish between high- and low-risk decisions. To elucidate the individual differences that influence the decisional processes, making the strategies more or less advantageous, we considered the impact of the BAS motivational system and the frontal left and right cortical activity on subjects' decisions. More specifically, the lateralization effect, which is related to the increased activation of the left (BAS-Reward-related) hemisphere, was explored by using frequency band analysis. Specifically, behavioral responses (gain/loss options), metacognition, and delta, theta, alpha, and beta band modulation (asymmetry index) were considered. Thirty subjects were divided into high-BAS and low-BAS groups. In comparison with low-BAS, the high-BAS group showed an increased tendency to opt in favor of the immediate reward (losing strategy) instead of the long-term option (winning strategy), and members of this group were more impaired in metacognitive monitoring of their strategies and showed an increased left hemisphere activation when they responded to losing choices. A "reward bias" effect was hypothesized to act for high BAS, based on a left-hemisphere hyperactivation.
Article
In the present study, we explored the contribution of different cortical areas in processing different semantic violations in action representation-that is, instrumental or functional violations. The cortical contribution in object-related action comprehension was verified by measuring changes in event-related potential (N400 effect), error rates (ERs), and response times (RTs), by applying an inhibitory transcranial direct current stimulation (tDCS) on the dorsolateral prefrontal cortex (DLPFC). Thirty-three subjects performed the detection task (action frames ending with a congruous vs. incongruous action). The tDCS effect was analyzed by comparing the N400, ERs, and RTs before and after stimulation. A significant reduction of the N400 and increased RTs were observed for incongruous stimuli in the case of inhibitory stimulation of the DLPFC. These results highlighted that DLPFC inhibition may limit the ability to analyze a semantically incongruous action, with a reduced N400 ERP effect and increased "cognitive costs" (higher RTs). Moreover, functional violation showed also the contribution of the temporoparietal areas to modulate the N400 amplitude. Therefore the existence of different cortical generators was supposed for the instrumental (more frontal) and the functional (more frontal and temporoparietal) semantic anomaly processing.
Article
Coherent representation of action sequences implies that the logical temporal order of each action can be correctly represented. Violation of this logical order may induce a sort of expectancies disruption of the temporal structure. Thus the present study explored the event-related potential (ERP) effect related to the cortical response to this violation. Action sequence composed by four frames with final congruous or incongruous endings was submitted to 28 subjects. Two distinct ERP effects, feedback-related negativity (FRN), and P300, were found in response to incongruous endings, with also significant increased RTs. The functional significance of these two ERP deflections was related respectively to the perception of an erroneous action outcome as the ending of an illogical sequence (FRN) and to the necessity to updating the relationship action-context by changing the cognitive model which supports the cognitive expectancies (P300). The significant correlation between the RTs and the ERP measures, especially in case of FRN effect, supported this interpretation. Indeed increased cognitive costs are supposed in case of expectancies violations which require further processes of reanalysis of the coherence between the action and the background (the temporal background) where the action was produced. Two different cortical localizations were found for FRN and P300, respectively a more fronto-central (dorsolateral prefrontal cortex) and posterior (superior temporal gyrus) site. The significance of these results for the temporal order effect for action comprehension was discussed.
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Sex differences in empathy for pain have been repeatedly observed. However, it is unclear whether this is due to sex differences in “bottom-up” somatomotor representations of others’ pain (self-other resonance) or to “top-down” prefrontal control of such responses. Here, we provide data from 70 subjects suggesting that sex differences in empathy for pain lie primarily in pre-reflective, bottom-up resonance mechanisms. Subjects viewed a right hand pierced by a needle during fMRI. They also filled out a self-report measure of trait empathy, the Interpersonal Reactivity Index. A permutation-based analysis (FSL’s Randomise) found that females showed greater signal in a cluster in primary somatomotor cortex that includes the motor hand area. No significant differences were observed in other task-implicated areas. An examination of condition-specific parameter estimates found that this difference was due to reduced signal in this cluster in males. No significant differences in resting connectivity or within-task (generalized psychophysiological interaction analysis or gPPI) dynamic connectivity of this region with prefrontal areas were observed. While female subjects scored higher on affective subscales of the IRI, there were no sex differences in Perspective-Taking, the primary index of cognitive, top-down empathy processes. These findings suggest that localized internal somatomotor representations of others’ pain, a functional index of bottom-up resonance processes, are stronger in female subjects.
Chapter
I had just received my badge, the conference program, and the complimentary laptop sleeve; I turned around and stood face to face with John, who was also waiting in line to register for the conference. We exchanged a short word of greeting and we shook hands wholeheartedly. When this encounter was over, I bumped into Mary; we both expressed our surprise and then hugged each other. Right after this hug, I nodded politely to Paul passing behind Mary, noticing that he wanted to engage in a conversation. Had Mary been accompanied by her spouse, perhaps the hug would have been toned down to a warm handshake. Had I not hugged Mary, I might have had more trouble avoiding Paul, whom I suspected to be reviewer #3 of my recently rejected paper. This type of familiar scene is usually fluid and many of these social interactions feel automatic. However, if we analyze further the simple action of the greeting, the initial social contact with others, we learn that incalculable cognitive operations subserved by a number of neurofunctional systems are at play. But at the source of this cascade of cerebral processing often lie the actions of the others.
Book
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Múltiples aproximaciones y modelos han tratado de explicar el funcionamiento de la memoria humana. En esta obra se habla tanto de lo cognitivo como de lo neurocientífico, y en cada capítulo los autores van contando diferentes aproximaciones a múltiples problemas relacionados con los procesos de memoria, aproximaciones tanto descriptivas como funcionales. Cuentan desde la topografía de las distintas áreas cerebrales o zonas que se estimulan al realizar tareas conductuales de memoria, hasta la explicación que tratan de darnos los investigadores que se han acercado a diseccionar estos problemas desde la psicología experimental o clínica. Es una obra excepcionalmente completa sobre la memoria. Su extensión permite abordar los distintos ámbitos desde los que considerar y estudiar la memoria, con una exposición atractiva, aunque no exenta de complejidad y conocimientos técnicos.
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O artigo apresenta estudo experimental sobre tomada de decisao tendo como objetivo geral: Identificar mecanismos de aprendizagem implicita e sua relacao com congruencia da informacao e influencia social em decisoes sobre metas de investimento. Procedimentos metodologicos: Estudo experimental junto a 22 sujeitos (alunos de graduacao da area de negocios) tendo como tarefa um jogo de decisao especialmente desenvolvido para o estudo, durante o qual os sujeitos estavam conectados a um EEG (Eletroencefalograma) que captava as ondas eletricas do cerebro no momento da decisao. A decisao envolvia escolher o nivel de investimento para 200 filiais de uma mesma empresa baseando-se em informacoes contabeis apresentadas em formato de graficos. Havia tres condicoes experimentais: congruencia das informacoes fornecidas, influencia de propositores que davam dicas sobre o nivel de investimento e correcao/incorrecao dessas dicas tendo em vista as informacoes contabeis. Principais resultados: Nas 4200 decisoes analisadas, todos os 21 sujeitos identificaram que havia uma regra para ganho/perda de pontos associada a decisao tomada, embora nao acertassem qual era a regra; nao levavam em consideracao a incongruencia semântica das informacoes; decidiam conforme a regra aprendiada, mesmo quando esta os levava a uma decisao tecnicamente erronea, contra a empresa. A analise do EEG mostrou que a incongruencia era captada (amplitude do componente N400, associado a violacoes de informacoes semânticas). Os resultados (comportamental e do EEG) comprovam em tempo real a presenca do conflito de interesses (auto-beneficio) e a violacao sistematica da analise da informacao para decisao.
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The goal of this study was to investigate whether the semantic processing of the audiovisual combination of communicative gestures with speech differs between men and women. We recorded event-related brain potentials in women and men during the presentation of communicative gestures that were either congruent or incongruent with the speech. Our results showed that incongruent gestures elicited an N400 effect over frontal sites compared to congruent ones in both groups. Moreover, the females showed an earlier N2 response to incongruent stimuli than congruent ones, while larger sustained negativity and late positivity in response to incongruent stimuli was observed only in males. These results suggest that women rapidly recognise and process audiovisual combinations of communicative gestures and speech (as early as 300 ms) whereas men analyse them at the later stages of the process.
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Previous studies did not directly compare the time course of neural correlates underlying the different kinds of social motor intentions, specifically the communicative intentions produced by two agents’ whole body movements. In order to characterize the time course of the communicative intention inference (vs. non-interactive intention) in the brain, we employed the ERP (event-related potential) technology to record 16 healthy participants’ electrical brain activity while they were watching photos depicting two actors’ actions, and performing an intention inference task (IIT). There were three different types of action intentions: kindness, hostility and non-interactiveness. Electrophysiological results revealed the peak amplitudes of N2 over the frontal sites for kindness intentions were significantly larger than those for hostility and non-interactive intentions. On the other hand, there were no significant differences between hostility and non-interactive intention at N2. At a later stage (i.e., during the 270–500 ms epoch), the peak amplitude of the P3 over the parietal sites and the right hemisphere was significantly larger for hostility intentions compared to the kindness and non-interactive intentions, while there were no significant differences between kindness and non-interactive intentions. The present study provides preliminary evidence of the temporal dynamics sustaining the dissociation between the understanding of different kinds of social motor intentions versus non-interactive motor intention.
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Actions are complex, goal-directed, movements, and, despite being hidden in the actor’s mind, observers successfully identify and anticipate actor’s goal. In this thesis, we identified two main approaches to explain how observers recognise others’ actions. Sensorimotor approaches consider action recognition as bottom-up propagation from the perception of visual kinematics to the recognition of action goals. Visual kinematics are viewed here as the primary source of visual information from which goal-related information is extracted. In contrast, predictive approaches assume that observers cannot make sense of visual kinematics without a prediction about the actor’s goal. Observers would extract goal-related information from non-motor sources of information to guide the processing of the visual kinematics. Information about the temporal dynamics of activation of visual kinematics and goal-related information during action visual processing is critical to disentangle the two approaches and to provide a better understanding of the mechanisms underlying action recognition, but empirical data in this direction are clearly lacking. In order to fill this gap, we investigated the relative priority given to visual kinematics versus non-motor goal-related information during the recognition of others’ actions. The contribution of visual kinematics and non-motor goal-related information was independently evaluated by introducing violations of grip and/or visual goal in photographs of object-directed actions. Using behavioural methods (priming and visual-search paradigms), we demonstrated that non-motor goal-related information was prioritised over visual kinematics during the first steps of visual action processing, whereas visual kinematics were prioritised over goal-related information later during visual action processing. Using neurophysiological methods (event-related potential and transcranial magnetic stimulation priming paradigms), we found that both visual kinematics and non-motor goal-related information are already processed during the perceptual stages of action processing, but that action semantic processing is guided by goal-related information rather than visual kinematics. We further provide evidence supporting the critical involvement of the frontoparietal network in the later integration of visual kinematics and non-motor goal-related information. We finally showed that the priority given to non-motor goal-related information over visual kinematics during action visual processing depends on individual social characteristics. Together, the findings reported are consistent with predictive approaches of action recognition. Results are discussed in the light of converging evidence suggesting that visual kinematics are used to update goal predictions that have been previously derived from non-motor goal-related information. Yet findings further orient towards a pluralist view of action understanding, in which the strategies used to process others’ actions may vary depending on situations and individuals.
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Event-related brain potentials (ERPs) were used to investigate the visual processing of actions belonging to the typical human repertoire. Two hundred and sixty coloured pictures representing persons differing in number, age and gender, engaged in simple actions, were presented to 23 right-handed students. Perception of meaningful actions (e.g., young woman trying shoes in shop) was contrasted with perception of actions lacking an understandable goal (e.g., businesswoman balancing on one foot in desert). The results indicated early recognition of comprehensible behaviour in the form of an enhanced posterior "recognition potential" (RP) (N250), which was followed by a larger negativity (N400) in response to incongruent actions. The results suggest that incoming visual information regarding human gestures is processed similarly to linguistic inputs from a conceptual point of view, thus eliciting a posterior RP when the action code is visually recognized and comprehended, and a later N400 when the action is not recognized or is difficult to integrate with previous knowledge.
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Misuse of tools and objects by patients with left brain damage is generally recognized as a manifestation of apraxia, caused by parietal lobe damage. The use of tools and objects can, however, be subdivided in several components. The purpose of our study was to find out which of these are dependent on parietal lobe function. Thirty-eight patients with left brain damage and aphasia were examined using tests to assess the retrieval of functional knowledge from semantic memory (Functional Associations), mechanical problem solving (Novel Tools) and use of everyday tools and objects (Common Tools). Voxel-wise analysis of magnetic resonance images revealed two regions where lesions had a significant impact on the test results. One extended rostrally from the central region and ventrally through the middle frontal cortex to the dorsal margin of the inferior frontal gyrus. The other reached dorsally and caudally from the supramarginal gyrus, through the inferior, to superior parietal lobe. Whereas the frontal lesions had an adverse influence on all experimental tests as well as on the subtests of the Aachen Aphasia test, parietal lesions impaired Novel and Common Tools, but did not have an adverse effect on the Functional Associates. An association between Functional Associations and temporal lesions became apparent when patients with only a selective deficit in the test were considered, but did not show up in the whole group analysis. The parietal influence was as strong for the selection as for the use of either novel or common tools, although choice of appropriate manual configuration and movements was more important for use than for selection. We conclude that the contribution of the parietal lobe to tool use concerns general principles of tool use rather than knowledge about the prototypical use of common tools and objects, and the comprehension of mechanical interactions of the tool with other tools, recipients or material rather than the selection of grip formation and manual movements.
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Our recent magnetoencephalography study demonstrated that the mu rhythm can reliably indicate sensorimotor resonance during the perception of pain in others (Cheng, Y., Yang, C.Y., Lin, C.P., Lee, P.L., Decety, J., 2008b. The perception of pain in others suppresses somatosensory oscillations: a magnetoencephalography study. NeuroImage 40, 1833-1840). The current study further investigated the neurophysiological mechanism underpinning empathy for pain in relation with gender through the measurements of the electroencephalographic mu suppression in healthy female (N=16) and male (N=16) adults during the observation of body parts in painful or no-painful situations. The results demonstrate that both genders exhibited sensorimotor activation related to pain empathy. However, females showed stronger mu suppressions than males when watching the painful as well as the non-painful situations. Further, the mu suppression for pain empathy was positively correlated with the scoring on the personal distress subscale of the interpersonal reactivity index only in the female participants. The present findings suggest the existence of a gender difference in pain empathy in relation with the sensorimotor cortex resonance. The mu rhythm can be a potential biomarker of empathic mimicry.
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The understanding of actions of tool use depends on the motor act that is performed and on the function of the objects involved in the action. We used event-related potentials (ERPs) to investigate the processes that derive both kinds of information in a task in which inserting actions had to be judged. The actions were presented as two consecutive frames, one showing an effector/instrument and the other showing a potential target object of the action. Two mismatches were possible. An orientation mismatch occurred when the spatial object properties were not consistent with a motor act of insertion being performed (i.e., different orientations of insert and slot). A functional mismatch happened when the instrument (e.g., screwdriver) would usually not be applied to the target object (e.g., keyhole). The order in which instrument and target object were presented was also varied. The two kinds of mismatch gave rise to similar but not identical negativities in the latency range of the N400 followed by a positive modulation. The results indicate that the motor act and the function of the objects are derived by two at least partially different subprocesses and become integrated into a common representation of the observed action.
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Females frequently perform better in empathy, interpersonal sensitivity, and emotional recognition than do males. The mirror-neuron system has been proposed to play an important role in social cognition. It remains to be clarified, however, whether the neuroanatomy underlying the human mirror neuron system exhibits sex differences. With the use of voxel-based morphometry analysis, a whole-brain unbiased technique to characterize regional cerebral volume differences in structural magnetic resonance images, concurrent with the dispositional empathy measures, we demonstrate that young adult females (n=25) had significantly larger gray matter volume in the pars opercularis and inferior parietal lobule than matched males (n=25) participants. Moreover, higher self-report scores in the emotional empathic disposition was tightly coupled with larger gray matter volume of the pars opercularis across all female and male participants (P=0.002). These results indicate that the existence of neuroanatomical sex differences in the human mirror-neuron system. They also suggest that the network of the human mirror-neuron system is strongly linked to empathy competence.
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The discovery of the mirror neuron system (MNS) has led researchers to speculate that this system evolved from an embodied visual recognition apparatus in monkey to a system critical for social skills in humans. It is accepted that the MNS is specialized for processing animate stimuli, although the degree to which social interaction modulates the firing of mirror neurons has not been investigated. In the current study, EEG mu wave suppression was used as an index of MNS activity. Data were collected while subjects viewed four videos: (1) Visual White Noise: baseline, (2) Non-interacting: three individuals tossed a ball up in the air to themselves, (3) Social Action, Spectator: three individuals tossed a ball to each other and (4) Social Action, Interactive: similar to video 3 except occasionally the ball would be thrown off the screen toward the viewer. The mu wave was modulated by the degree of social interaction, with the Non-interacting condition showing the least suppression, followed by the Social Action, Spectator condition and the Social Action, Interactive condition showing the most suppression. These data suggest that the human MNS is specialized not only for processing animate stimuli, but specifically stimuli with social relevance.
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Hand preference for the original items proposed by Oldfield (1971) and information concerning age, sex, familial sinistrality were obtained from a population of 1694 subjects. An item analysis was performed which resulted in the elimination of some of the items. Handedness distribution derived on the basis of the selected items was compared with the distribution obtained on the basis of the Oldfield's selection. Results show that handedness distributions depend on item selection, familial sinistrality and age, while no effect of sex is found.
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Face perception, perhaps the most highly developed visual skill in humans, is mediated by a distributed neural system in humans that is comprised of multiple, bilateral regions. We propose a model for the organization of this system that emphasizes a distinction between the representation of invariant and changeable aspects of faces. The representation of invariant aspects of faces underlies the recognition of individuals, whereas the representation of changeable aspects of faces, such as eye gaze, expression, and lip movement, underlies the perception of information that facilitates social communication. The model is also hierarchical insofar as it is divided into a core system and an extended system. The core system is comprised of occipitotemporal regions in extrastriate visual cortex that mediate the visual analysis of faces. In the core system, the representation of invariant aspects is mediated more by the face-responsive region in the fusiform gyrus, whereas the representation of changeable aspects is mediated more by the face-responsive region in the superior temporal sulcus. The extended system is comprised of regions from neural systems for other cognitive functions that can be recruited to act in concert with the regions in the core system to extract meaning from faces.
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Although the sentences that we hear or read have meaning, this does not necessarily mean that they are also true. Relatively little is known about the critical brain structures for, and the relative time course of, establishing the meaning and truth of linguistic expressions. We present electroencephalogram data that show the rapid parallel integration of both semantic and world knowledge during the interpretation of a sentence. Data from functional magnetic resonance imaging revealed that the left inferior prefrontal cortex is involved in the integration of both meaning and world knowledge. Finally, oscillatory brain responses indicate that the brain keeps a record of what makes a sentence hard to interpret.
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Results obtained with functional magnetic resonance imaging show that both feeling a moderately painful pinprick stimulus to the fingertips and witnessing another person's hand undergo similar stimulation are associated with common activity in a pain-related area in the right dorsal anterior cingulate cortex (ACC). Common activity in response to noxious tactile and visual stimulation was restricted to the right inferior Brodmann's area 24b. These results suggest a shared neural substrate for felt and seen pain for aversive ecological events happening to strangers and in the absence of overt symbolic cues. In contrast to ACC 24b, the primary somatosensory cortex showed significant activations in response to both noxious and innocuous tactile, but not visual, stimuli. The different response patterns in the two areas are consistent with the ACC's role in coding the motivational-affective dimension of pain, which is associated with the preparation of behavioral responses to aversive events.
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Understanding the intentions of others while watching their actions is a fundamental building block of social behavior. The neural and functional mechanisms underlying this ability are still poorly understood. To investigate these mechanisms we used functional magnetic resonance imaging. Twenty-three subjects watched three kinds of stimuli: grasping hand actions without a context, context only (scenes containing objects), and grasping hand actions performed in two different contexts. In the latter condition the context suggested the intention associated with the grasping action (either drinking or cleaning). Actions embedded in contexts, compared with the other two conditions, yielded a significant signal increase in the posterior part of the inferior frontal gyrus and the adjacent sector of the ventral premotor cortex where hand actions are represented. Thus, premotor mirror neuron areas-areas active during the execution and the observation of an action-previously thought to be involved only in action recognition are actually also involved in understanding the intentions of others. To ascribe an intention is to infer a forthcoming new goal, and this is an operation that the motor system does automatically.
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Empathizing is the capacity to predict and to respond to the behavior of agents (usually people) by inferring their mental states and responding to these with an appropriate emotion. Systemizing is the capacity to predict and to respond to the behavior of nonagentive deterministic systems by analyzing input-operation-output relations and inferring the rules that govern such systems. At a population level, females are stronger empathizers and males are stronger systemizers. The “extreme male brain” theory posits that autism represents an extreme of the male pattern (impaired empathizing and enhanced systemizing). Here we suggest that specific aspects of autistic neuroanatomy may also be extremes of typical male neuroanatomy.
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The neural processes underlying empathy are a subject of intense interest within the social neurosciences. However, very little is known about how brain empathic responses are modulated by the affective link between individuals. We show here that empathic responses are modulated by learned preferences, a result consistent with economic models of social preferences. We engaged male and female volunteers in an economic game, in which two confederates played fairly or unfairly, and then measured brain activity with functional magnetic resonance imaging while these same volunteers observed the confederates receiving pain. Both sexes exhibited empathy-related activation in pain-related brain areas (fronto-insular and anterior cingulate cortices) towards fair players. However, these empathy-related responses were significantly reduced in males when observing an unfair person receiving pain. This effect was accompanied by increased activation in reward-related areas, correlated with an expressed desire for revenge. We conclude that in men (at least) empathic responses are shaped by valuation of other people's social behaviour, such that they empathize with fair opponents while favouring the physical punishment of unfair opponents, a finding that echoes recent evidence for altruistic punishment.
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Current cognitive neuroscience models predict a right-hemispheric dominance for face processing in humans. However, neuroimaging and electromagnetic data in the literature provide conflicting evidence of a right-sided brain asymmetry for decoding the structural properties of faces. The purpose of this study was to investigate whether this inconsistency might be due to gender differences in hemispheric asymmetry. In this study, event-related brain potentials (ERPs) were recorded in 40 healthy, strictly right-handed individuals (20 women and 20 men) while they observed infants' faces expressing a variety of emotions. Early face-sensitive P1 and N1 responses to neutral vs. affective expressions were measured over the occipital/temporal cortices, and the responses were analyzed according to viewer gender. Along with a strong right hemispheric dominance for men, the results showed a lack of asymmetry for face processing in the amplitude of the occipito-temporal N1 response in women to both neutral and affective faces. Men showed an asymmetric functioning of visual cortex while decoding faces and expressions, whereas women showed a more bilateral functioning. These results indicate the importance of gender effects in the lateralization of the occipito-temporal response during the processing of face identity, structure, familiarity, or affective content.
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Self-other discrimination is fundamental to social interaction, however, little is known about the neural systems underlying this ability. In a previous functional magnetic resonance imaging study, we demonstrated that a right fronto-parietal network is activated during viewing of self-faces as compared with the faces of familiar others. Here we used image-guided repetitive transcranial magnetic stimulation (rTMS) to create a 'virtual lesion' over the parietal component of this network to test whether this region is necessary for discriminating self-faces from other familiar faces. The current results indeed show that 1 Hz rTMS to the right inferior parietal lobule (IPL) selectively disrupts performance on a self-other discrimination task. Applying 1 Hz rTMS to the left IPL had no effect. It appears that activity in the right IPL is essential to the task, thus providing for the first time evidence for a causal relation between a human brain area and this high-level cognitive capacity.
Article
Sophisticated analysis methods for EEG and MEG play a key role in the better understanding of brain functions as measured by high-density EEG and MEG. Being commercially available since 1996, the ASA software (ANT Software BV, Enschede, Netherlands) has been gaining growing popularity among clinical and cognitive researchers. With the following article, we present an overview on the currently available functionality of the software and provide examples of its application.
Article
The long-term consequences of early prefrontal cortex lesions occurring before 16 months were investigated in two adults. As is the case when such damage occurs in adulthood, the two early-onset patients had severely impaired social behavior despite normal basic cognitive abilities, and showed insensitivity to future consequences of decisions, defective autonomic responses to punishment contingencies and failure to respond to behavioral interventions. Unlike adult-onset patients, however, the two patients had defective social and moral reasoning, suggesting that the acquisition of complex social conventions and moral rules had been impaired. Thus early-onset prefrontal damage resulted in a syndrome resembling psychopathy.
Article
My initial scope will be limited: starting from a neurobiological standpoint, I will analyse how actions are possibly represented and understood. The main aim of my arguments will be to show that, far from being exclusively dependent upon mentalistic/linguistic abilities, the capacity for understanding others as intentional agents is deeply grounded in the relational nature of action. Action is relational, and the relation holds both between the agent and the object target of the action (see Gallese, 2000b), as between the agent of the action and his/her observer (see below). Agency constitutes a key issue for the understanding of intersubjectivity and for explaining how individuals can interpret their social world. This account of intersubjectivity, founded on the empirical findings of neuroscientific investigation, will be discussed and put in relation with a classical tenet of phenomenology: empathy. I will provide an 'enlarged' account of empathy that will be defined by means of a new conceptual tool: the shared manifold of intersubjectivity.
Article
Aim: Neuroimaging studies on biological motion have established the view that the posterior superior temporal sulcus (pSTS) is involved in detecting intention of others. Those studies have consistently reported other regions such as body-selective extrastriate body area (EBA) and motion-sensitive middle temporal, in close proximity to pSTS. Whether EBA responds only to static body parts or has a more extended role as part of a system for inferring intention of others has remained an elusive issue. The aim of the present study was to investigate the role of EBA in processing goal-directed actions. Methods: Twelve healthy volunteers participated in the present study. Using sports-related motions as visual stimuli, brain activations were examined during observation of goal-directed actions and non-goal-directed actions on functional magnetic resonance imaging. Results: Compared to non-goal-directed actions, goal-directed actions produced greater activations in EBA along with the mirror neuron system. Conclusions: EBA might contribute to understanding others' actions by representing the dynamic aspects of human motions.
Article
Numerous cortical regions respond to aspects of the human form and its actions. What is the contribution of the extrastriate body area (EBA) to this network? In particular, is the EBA involved in constructing a dynamic representation of observed actions? We scanned 16 participants with fMRI while they viewed two kinds of stimulus sequences. In the coherent condition, static frames from a movie of a single, intransitive whole-body action were presented in the correct order. In the incoherent condition, a series of frames from multiple actions (involving one actor) were presented. ROI analyses showed that the EBA, unlike area MT + and the posterior superior temporal sulcus, responded more to the incoherent than to the coherent conditions. Whole brain analyses revealed increased activation to the coherent sequences in parietal and frontal regions that have been implicated in the observation and control of movement. We suggest that the EBA response adapts when succeeding images depict relatively similar postures (coherent condition) compared to relatively different postures (incoherent condition). We propose that the EBA plays a unique role in the perception of action, by representing the static structure, rather than dynamic aspects, of the human form.
Article
Event-related potentials (ERPs) were recorded during the evaluation of the appropriateness of cooperative actions between two people. We used pictures of pass-and-receive actions as one type of cooperative actions, in which one person passed an object and another reached out to receive it with a preshaped hand. Eleven participants judged whether the receiving actions were appropriate or inappropriate in relation to the passing ones. The inappropriate actions elicited a widely distributed and parietal maximum N400 as compared with the appropriate ones. These results suggest that the N400 is evoked in the incongruous context in which two people perform inappropriate cooperative actions and is related to the semantic processing that involves the prediction of interpersonal action sequences.
Article
Tager-Flusberg and Sullivan [Tager-Flusberg, H., Sullivan, K., 2000. A componential view of theory of mind: evidence from Williams syndrome. Cognition 76, 59-90] have argued for a distinction between the social-perceptive component of theory of mind (ToM), involving judgment of mental state from facial and body expressions, and the social-cognitive component, which is representation-based and linked to language and theory-building. This is analogous to the distinction made by others [Gallese, V., Keysers, C., Rizzolatti, G., 2004. A unifying view of the basis of social cognition. Trends in Cognitive Science 8, 396-403] between representing the mental state of another as if it was one's own (simulation theory), which requires involvement of the mirror neuron system, and explicit or declarative reasoning about mental states (theory theory), which does not. This componential view of ToM was tested by examining mirroring, as indexed by EEG mu rhythm suppression, in subjects performing tasks assumed to tap both dimensions. Mu suppression was positively correlated with accuracy on the social-perceptual task but not in the social-cognitive task. In a ToM control task requiring judgments about person-object interactions accuracy was correlated with mu suppression. This implies that mirroring is involved in making judgments about emotions and person-object interactions. However, mirroring is insensitive to the distinction between correct and incorrect inferences in the social-cognitive task suggesting that additional mechanisms are needed to make mental attributions of beliefs and intentions. These results are consistent with a refined componential view of ToM.
Article
Recent findings have demonstrated that women might be more reactive than men to viewing painful stimuli (vicarious response to pain), and therefore more empathic [Han, S., Fan, Y., & Mao, L. (2008). Gender difference in empathy for pain: An electrophysiological investigation. Brain Research, 1196, 85-93]. We investigated whether the two sexes differed in their cerebral responses to affective pictures portraying humans in different positive or negative contexts compared to natural or urban scenarios. 440 IAPS slides were presented to 24 Italian students (12 women and 12 men). Half the pictures displayed humans while the remaining scenes lacked visible persons. ERPs were recorded from 128 electrodes and swLORETA (standardized weighted Low-Resolution Electromagnetic Tomography) source reconstruction was performed. Occipital P115 was greater in response to persons than to scenes and was affected by the emotional valence of the human pictures. This suggests that processing of biologically relevant stimuli is prioritized. Orbitofrontal N2 was greater in response to positive than negative human pictures in women but not in men, and not to scenes. A late positivity (LP) to suffering humans far exceeded the response to negative scenes in women but not in men. In both sexes, the contrast suffering-minus-happy humans revealed a difference in the activation of the occipito/temporal, right occipital (BA19), bilateral parahippocampal, left dorsal prefrontal cortex (DPFC) and left amygdala. However, increased right amygdala and right frontal area activities were observed only in women. The humans-minus-scenes contrast revealed a difference in the activation of the middle occipital gyrus (MOG) in men, and of the left inferior parietal (BA40), left superior temporal gyrus (STG, BA38) and right cingulate (BA31) in women (270-290 ms). These data indicate a sex-related difference in the brain response to humans, possibly supporting human empathy.
Article
Event-related brain potentials (ERPs) were recorded while subjects silently read several prose passages, presented one word at a time. Semantic anomalies and various grammatical errors had been inserted unpredictably at different serial positions within some of the sentences. The semantically inappropriate words elicited a large N400 component in the ERP, whereas the grammatical errors were associated with smaller and less consistent components that had scalp distributions different from that of the N400. This result adds to the evidence that the N400 wave is more closely related to semantic than to grammatical processing. Additional analyses revealed that different ERP configurations were elicited by open-class (“content”) and closed-class (“function”) words in these prose passages.
Article
This paper presents a new method for localizing the electric activity in the brain based on multichannel surface EEG recordings. In contrast to the models presented up to now the new method does not assume a limited number of dipolar point sources nor a distribution on a given known surface, but directly computes a current distribution throughout the full brain volume. In order to find a unique solution for the 3-dimensional distribution among the infinite set of different possible solutions, the method assumes that neighboring neurons are simultaneously and synchronously activated. The basic assumption rests on evidence from single cell recordings in the brain that demonstrates strong synchronization of adjacent neurons. In view of this physiological consideration the computational task is to select the smoothest of all possible 3-dimensional current distributions, a task that is a common procedure in generalized signal processing. The result is a true 3-dimensional tomography with the characteristic that localization is preserved with a certain amount of dispersion, i.e., it has a relatively low spatial resolution. The new method, which we call Low Resolution Electromagnetic Tomography (LORETA) is illustrated with two different sets of evoked potential data, the first showing the tomography of the P100 component to checkerboard stimulation of the left, right, upper and lower hemiretina, and the second showing the results for the auditory N100 component and the two cognitive components CNV and P300. A direct comparison of the tomography results with those obtained from fitting one and two dipoles illustrates that the new method provides physiologically meaningful results while dipolar solutions fail in many situations. In the case of the cognitive components, the method offers new hypotheses on the location of higher cognitive functions in the brain.
To characterize the effects of normal aging on the amplitude, latency and scalp distribution of the N400 congruity effect. Event-related brain potentials (ERPs) were recorded from 72 adults (half of them men) between the ages of 20 and 80 years (12/decade) as they performed a semantic categorization task. Participants listened to spoken phrases (e.g. 'a type of fruit' or 'the opposite of black') followed about 1 s later by a visually-presented word that either did or did not fit with the sense of the preceding phrase; they reported the word read and whether or not it was appropriate. ERP measurements (mean amplitudes, peak amplitudes, peak latencies) were subjected to analysis of variance and linear regression analyses. All participants, regardless of age, produced larger N400s to words that did not fit than to those that did. The N400 congruity effect (no-fit ERPs - fit ERPs) showed a reliable linear decrease in the amplitude (0.05-0.09 microV per year, r = 0.40) and a reliable linear increase peak latency (1.5-2.1 ms/year, r = 0.60) with age. In sum, the N400 semantic congruity effect at the scalp gets smaller, slower and more variable with age, consistent with a quantitative rather than qualitative change in semantic processing (integration) with normal aging.
Article
Although it is widely accepted that the cortex participates in pain perception, there is no direct evidence for the existence of cortical neurons that respond to noxious or painful stimuli in humans. Anatomical and neurophysiological studies in animals as well as brain imaging and evoked potential studies in humans suggest that the anterior cingulate cortex (ACC) is an important area for processing sensory information related to pain1, 2, 3, 4, 5, 6, 7. We have now identified single neurons in ACC that respond selectively to painful thermal and mechanical stimuli, supporting a role for the ACC in pain perception.
Article
The long-term consequences of early prefrontal cortex lesions occurring before 16 months were investigated in two adults. As is the case when such damage occurs in adulthood, the two early-onset patients had severely impaired social behavior despite normal basic cognitive abilities, and showed insensitivity to future consequences of decisions, defective autonomic responses to punishment contingencies and failure to respond to behavioral interventions. Unlike adult-onset patients, however, the two patients had defective social and moral reasoning, suggesting that the acquisition of complex social conventions and moral rules had been impaired. Thus early-onset prefrontal damage resulted in a syndrome resembling psychopathy.
Article
In two ERP experiments we investigated how and when the language comprehension system relates an incoming word to semantic representations of an unfolding local sentence and a wider discourse. In Experiment 1, subjects were presented with short stories. The last sentence of these stories occasionally contained a critical word that, although acceptable in the local sentence context, was semantically anomalous with respect to the wider discourse (e.g., Jane told the brother that he was exceptionally slow in a discourse context where he had in fact been very quick). Relative to coherent control words (e.g., quick), these discourse-dependent semantic anomalies elicited a large N400 effect that began at about 200 to 250 msec after word onset. In Experiment 2, the same sentences were presented without their original story context. Although the words that had previously been anomalous in discourse still elicited a slightly larger average N400 than the coherent words, the resulting N400 effect was much reduced, showing that the large effect observed in stories depended on the wider discourse. In the same experiment, single sentences that contained a clear local semantic anomaly elicited a standard sentence-dependent N400 effect (e.g., Kutas & Hillyard, 1980). The N400 effects elicited in discourse and in single sentences had the same time course, overall morphology, and scalp distribution. We argue that these findings are most compatible with models of language processing in which there is no fundamental distinction between the integration of a word in its local (sentence-level) and its global (discourse-level) semantic context.