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1.  Valence-specific conflict moderation in the dorso-medial PFC and the caudate head in emotional speech 
Emotional speech comprises of complex multimodal verbal and non-verbal information that allows deducting others’ emotional states or thoughts in social interactions. While the neural correlates of verbal and non-verbal aspects and their interaction in emotional speech have been identified, there is very little evidence on how we perceive and resolve incongruity in emotional speech, and whether such incongruity extends to current concepts of task-specific prediction errors as a consequence of unexpected action outcomes (‘negative surprise’). Here, we explored this possibility while participants listened to congruent and incongruent angry, happy or neutral utterances and categorized the expressed emotions by their verbal (semantic) content. Results reveal valence-specific incongruity effects: negative verbal content expressed in a happy tone of voice increased activation in the dorso-medial prefrontal cortex (dmPFC) extending its role from conflict moderation to appraisal of valence-specific conflict in emotional speech. Conversely, the caudate head bilaterally responded selectively to positive verbal content expressed in an angry tone of voice broadening previous accounts of the caudate head in linguistic control to moderating valence-specific control in emotional speech. Together, these results suggest that control structures of the human brain (dmPFC and subcompartments of the basal ganglia) impact emotional speech differentially when conflict arises.
doi:10.1093/scan/nsu021
PMCID: PMC4321614  PMID: 24526187
fMRI; conflict; prediction error; dmPFC; caudate head; emotional speech
2.  Empathy for social exclusion involves the sensory-discriminative component of pain: a within-subject fMRI study 
Recent research has shown that experiencing events that represent a significant threat to social bonds activates a network of brain areas associated with the sensory-discriminative aspects of pain. In the present study, we investigated whether the same brain areas are involved when witnessing social exclusion threats experienced by others. Using a within-subject design, we show that an ecologically valid experience of social exclusion recruits areas coding the somatosensory components of physical pain (posterior insular cortex and secondary somatosensory cortex). Furthermore, we show that this pattern of activation not only holds for directly experienced social pain, but also during empathy for social pain. Finally, we report that subgenual cingulate cortex is the only brain area conjointly active during empathy for physical and social pain. This supports recent theories that affective processing and homeostatic regulation are at the core of empathic responses.
doi:10.1093/scan/nsu038
PMCID: PMC4321615  PMID: 24563529
social exclusion; physical pain; empathy; fMRI; somatosensory cortex
3.  Neural correlates of ‘distracting’ from emotion during autobiographical recollection 
Remembering emotional autobiographical memories (AMs) is important for emotional well-being, and investigation of the role of emotion regulation (ER) during AM recollection has relevance for understanding mental health issues. Although significant progress has been made in understanding the brain mechanisms underlying ER and AM, less is known about the role of ER during AM recollection. The present study investigated how focusing away (or ‘distracting’) from the emotional content during AM recollection influences the subjective re-experiencing of emotions and the associated neural correlates, by manipulating the retrieval focus of participants who remembered emotional AMs while fMRI data were recorded. First, focusing away from emotion led to decreased self-reported emotional responses, along with increased engagement of ER-related regions (ventro-medial prefrontal cortex, vmPFC), and reduced activity in emotion-related regions (amygdala, AMY). Second, increased vmPFC activity was linked to reduced emotional ratings, during the non-emotional focus. Third, mediation analysis identified vmPFC as a functional hub integrating affective signals from AMY and mediating their impact on the subjective re-experiencing of emotion, according to the current retrieval focus. Collectively, these findings shed light on the neural mechanisms underlying the ability to effectively switch attentional focus away from emotions during AM recollections and have direct relevance for understanding, preventing and treating affective disorders, characterised by reduced ability to regulate emotions.
doi:10.1093/scan/nsu039
PMCID: PMC4321616  PMID: 24603021
personal memories; emotion control; ventro-medial PFC; emotional disorders
4.  Brain structure links trait creativity to openness to experience 
Creativity is crucial to the progression of human civilization and has led to important scientific discoveries. Especially, individuals are more likely to have scientific discoveries if they possess certain personality traits of creativity (trait creativity), including imagination, curiosity, challenge and risk-taking. This study used voxel-based morphometry to identify the brain regions underlying individual differences in trait creativity, as measured by the Williams creativity aptitude test, in a large sample (n = 246). We found that creative individuals had higher gray matter volume in the right posterior middle temporal gyrus (pMTG), which might be related to semantic processing during novelty seeking (e.g. novel association, conceptual integration and metaphor understanding). More importantly, although basic personality factors such as openness to experience, extroversion, conscientiousness and agreeableness (as measured by the NEO Personality Inventory) all contributed to trait creativity, only openness to experience mediated the association between the right pMTG volume and trait creativity. Taken together, our results suggest that the basic personality trait of openness might play an important role in shaping an individual’s trait creativity.
doi:10.1093/scan/nsu041
PMCID: PMC4321617  PMID: 24603022
openness to experience; trait creativity; creativity assessment packet; voxel-based morphometry
5.  Bad and worse: neural systems underlying reappraisal of high- and low-intensity negative emotions 
One of the most effective strategies for regulating emotional responses is cognitive reappraisal. While prior work has made great strides in characterizing reappraisal’s neural mechanisms and behavioral outcomes, the key issue of how regulation varies as a function of emotional intensity remains unaddressed. We compared the behavioral and neural correlates of reappraisal of high- and low-intensity emotional responses using functional magnetic resonance imaging (fMRI). We found that successful reappraisal of both high- and low-intensity emotions depends upon recruitment of dorsomedial (dmPFC) as well as left dorsolateral (dlPFC) and ventrolateral (vlPFC) prefrontal cortex. However, reappraisal of high-intensity emotions more strongly activated left dlPFC, and in addition, activated right lateral and dorsomedial PFC regions not recruited by low-intensity reappraisal. No brain regions were more strongly recruited during reappraisal of low when compared with high-intensity emotions. Taken together, these results suggest that reappraisal of high-intensity emotion requires greater cognitive resources as evidenced by quantitative and qualitative differences in prefrontal recruitment. These data have implications for understanding how and when specific PFC systems are needed to regulate different types of emotional responses.
doi:10.1093/scan/nsu043
PMCID: PMC4321618  PMID: 24603024
reappraisal; emotional intensity
6.  Cumulative activation during positive and negative events and state anxiety predicts subsequent inertia of amygdala reactivity 
Inertia, together with intensity and valence, is an important component of emotion. We tested whether positive and negative events generate lingering changes in subsequent brain responses to unrelated threat stimuli and investigated the impact of individual anxiety. We acquired fMRI data while participants watched positive or negative movie-clips and subsequently performed an unrelated task with fearful and neutral faces. We quantified changes in amygdala reactivity to fearful faces as a function of the valence of preceding movies and cumulative neural activity evoked during them. We demonstrate that amygdala responses to emotional movies spill over to subsequent processing of threat information in a valence-specific manner: negative movies enhance later amygdala activation whereas positive movies attenuate it. Critically, the magnitude of such changes is predicted by a measure of cumulative amygdala responses to the preceding positive or negative movies. These effects appear independent of overt attention, are regionally limited to amygdala, with no changes in functional connectivity. Finally, individuals with higher state anxiety displayed stronger modulation of amygdala reactivity by positive movies. These results suggest that intensity and valence of emotional events as well as anxiety levels promote local changes in amygdala sensitivity to threat, highlighting the importance of past experience in shaping future affective reactivity.
doi:10.1093/scan/nsu044
PMCID: PMC4321619  PMID: 24603023
emotional states; emotional inertia; positive emotions; mood; sensitization; desensitization; state anxiety; amygdala; fMRI
7.  Acting on social exclusion: neural correlates of punishment and forgiveness of excluders 
This functional magnetic resonance imaging study examined the neural correlates of punishment and forgiveness of initiators of social exclusion (i.e. ‘excluders’). Participants divided money in a modified Dictator Game between themselves and people who previously either included or excluded them during a virtual ball-tossing game (Cyberball). Participants selectively punished the excluders by decreasing their outcomes; even when this required participants to give up monetary rewards. Punishment of excluders was associated with increased activation in the pre-supplementary motor area (pre-SMA) and bilateral anterior insula. Costly punishment was accompanied by higher activity in the pre-SMA compared with punishment that resulted in gains or was non-costly. Refraining from punishment (i.e. forgiveness) was associated with self-reported perspective-taking and increased activation in the bilateral temporoparietal junction, dorsomedial prefrontal cortex, dorsal anterior cingulate cortex, and ventrolateral and dorsolateral prefrontal cortex. These findings show that social exclusion can result in punishment as well as forgiveness of excluders and that separable neural networks implicated in social cognition and cognitive control are recruited when people choose either to punish or to forgive those who excluded them.
doi:10.1093/scan/nsu045
PMCID: PMC4321620  PMID: 24652858
Dictator Game; fairness; fMRI; ostracism; perspective-taking
8.  Age differences in the default network at rest and the relation to self-referential processing 
Older adults show a ‘positivity bias’ in tasks involving emotion and self-referential processing. A critical network that is involved in self-referencing and shows age-related decline is the default network (DN). The purpose of the current study was to investigate age differences in pre- and post-task DN functional connectivity (FC) and signal variability, and to examine whether they are predictive of the positivity bias in self-referencing. We measured FC and within-subject variability of the DN in resting-state scans preceding and following tasks involving personality judgements on the self and a close other. Older adults endorsed more positive traits than younger adults on both tasks. FC was weaker post-task in older vs younger adults, and younger adults had greater variability than older adults in DN nodes. Younger adults with higher post-task DN variability had more negative self-ratings. For both age groups, greater FC in the DN was associated with more negative self-ratings. Neither FC nor variability was related to other ratings, despite the potential for self-processing when making other judgements. Our findings suggest that ageing leads to reduced FC and variability in the DN, which is most apparent after task, and may be one mechanism underlying the positive bias with age.
doi:10.1093/scan/nsu046
PMCID: PMC4321621  PMID: 24652859
ageing; default network; resting state; self-referential processing; positivity effect
9.  Eye’m talking to you: speakers’ gaze direction modulates co-speech gesture processing in the right MTG 
Recipients process information from speech and co-speech gestures, but it is currently unknown how this processing is influenced by the presence of other important social cues, especially gaze direction, a marker of communicative intent. Such cues may modulate neural activity in regions associated either with the processing of ostensive cues, such as eye gaze, or with the processing of semantic information, provided by speech and gesture. Participants were scanned (fMRI) while taking part in triadic communication involving two recipients and a speaker. The speaker uttered sentences that were and were not accompanied by complementary iconic gestures. Crucially, the speaker alternated her gaze direction, thus creating two recipient roles: addressed (direct gaze) vs unaddressed (averted gaze) recipient. The comprehension of Speech&Gesture relative to SpeechOnly utterances recruited middle occipital, middle temporal and inferior frontal gyri, bilaterally. The calcarine sulcus and posterior cingulate cortex were sensitive to differences between direct and averted gaze. Most importantly, Speech&Gesture utterances, but not SpeechOnly utterances, produced additional activity in the right middle temporal gyrus when participants were addressed. Marking communicative intent with gaze direction modulates the processing of speech–gesture utterances in cerebral areas typically associated with the semantic processing of multi-modal communicative acts.
doi:10.1093/scan/nsu047
PMCID: PMC4321622  PMID: 24652857
co-speech gestures; speech–gesture integration; eye gaze; communicative intent; middle temporal gyrus
10.  Potentiation of the early visual response to learned danger signals in adults and adolescents 
The reinforcing effects of aversive outcomes on avoidance behaviour are well established. However, their influence on perceptual processes is less well explored, especially during the transition from adolescence to adulthood. Using electroencephalography, we examined whether learning to actively or passively avoid harm can modulate early visual responses in adolescents and adults. The task included two avoidance conditions, active and passive, where two different warning stimuli predicted the imminent, but avoidable, presentation of an aversive tone. To avoid the aversive outcome, participants had to learn to emit an action (active avoidance) for one of the warning stimuli and omit an action for the other (passive avoidance). Both adults and adolescents performed the task with a high degree of accuracy. For both adolescents and adults, increased N170 event-related potential amplitudes were found for both the active and the passive warning stimuli compared with control conditions. Moreover, the potentiation of the N170 to the warning stimuli was stable and long lasting. Developmental differences were also observed; adolescents showed greater potentiation of the N170 component to danger signals. These findings demonstrate, for the first time, that learned danger signals in an instrumental avoidance task can influence early visual sensory processes in both adults and adolescents.
doi:10.1093/scan/nsu048
PMCID: PMC4321623  PMID: 24652856
N170 event-related potential; danger signals; avoidance behaviour; adolescence; learning-dependent plasticity
11.  Reduced long-range functional connectivity in young children with autism spectrum disorder 
Autism spectrum disorder (ASD) is often described as a disorder of aberrant neural connectivity. Although it is important to study the pathophysiology of ASD in the developing cortex, the functional connectivity in the brains of young children with ASD has not been well studied. In this study, brain activity was measured non-invasively during consciousness in 50 young human children with ASD and 50 age- and gender-matched typically developing human (TD) children. We employed a custom child-sized magnetoencephalography (MEG) system in which sensors were located as close to the brain as possible for optimal recording in young children. We focused on theta band oscillations because they are thought to be involved in long-range networks associated with higher cognitive processes. The ASD group showed significantly reduced connectivity between the left-anterior and the right-posterior areas, exhibiting a decrease in the coherence of theta band (6 Hz) oscillations compared with the TD group. This reduction in coherence was significantly correlated with clinical severity in right-handed children with ASD. This is the first study to demonstrate reduced long-range functional connectivity in conscious young children with ASD using a novel MEG approach.
doi:10.1093/scan/nsu049
PMCID: PMC4321624  PMID: 24652855
autism spectrum disorder (ASD); coherence; long-range connectivity; magnetoencephalography (MEG); theta oscillation; young children
12.  Developmental trajectories of cortical–subcortical interactions underlying the evaluation of trust in adolescence 
Social decision making is guided by the ability to intuitively judge personal attributes, including analysis of facial features to infer the trustworthiness of others. Although the neural basis for trustworthiness evaluation is well characterized in adults, less is known about its development during adolescence. We used event-related functional magnetic resonance imaging to examine age-related changes in neural activation and functional connectivity during the evaluation of trust in faces in a sample of adolescent females. During scanning, participants viewed masked presentations of faces and rated their trustworthiness. Parametric modeling of trust ratings revealed enhanced activation in amygdala and insula to untrustworthy faces, effects which peaked during mid-adolescence. Analysis of amygdala functional connectivity demonstrated enhanced amygdala–insula coupling during the evaluation of untrustworthy faces. This boost in connectivity was attenuated during mid-adolescence, suggesting a functional transition within face-processing circuits. Together, these findings underscore adolescence as a period of reorganization in neural circuits underlying socioemotional behavior.
doi:10.1093/scan/nsu050
PMCID: PMC4321625  PMID: 24682131
trust; adolescence; amygdala; insula; face processing
13.  Activation of frontoparietal attention networks by non-predictive gaze and arrow cues 
Gaze and arrow cues automatically orient visual attention, even when they have no predictive value, but the neural circuitry by which they direct attention is not clear. Recent evidence has indicated that the ventral frontoparietal attention network is primarily engaged by breaches of a viewer’s cue-related expectations. Accordingly, we hypothesized that to the extent that non-predictive gaze and arrow cues automatically engender expectations with regard to cue location, they should activate the ventral attention network when they cue attention invalidly. Using event-related fMRI, we found that invalid gaze but not arrow cues activated the ventral attention network, specifically in the area of the right temporal parietal junction (TPJ), as well as nodes along the dorsal attention network associated with a redirection of attention to the correct target location. In additional whole-brain analyses, facilitation of behavioral response time by valid gaze cues was linearly associated with the degree of activation in the right TPJ. We conclude from our findings that gaze direction elicits potent expectations in humans with regard to an actor’s intention that engage attention networks if not differently from, at least more robustly than, arrow cues.
doi:10.1093/scan/nsu054
PMCID: PMC4321626  PMID: 24748545
gaze; arrow; visual attention; spatial orienting; intention
14.  Are you gonna leave me? Separation anxiety is associated with increased amygdala responsiveness and volume 
The core feature of separation anxiety is excessive distress when faced with actual or perceived separation from people to whom the individual has a strong emotional attachment. So far little is known about the neurobiological underpinnings of separation anxiety. Therefore, we investigated functional (amygdala responsiveness and functional connectivity during threat-related emotion processing) and structural (grey matter volume) imaging markers associated with separation anxiety as measured with the Relationship Scale Questionnaire in a large sample of healthy adults from the Münster Neuroimaging Cohort (N = 320). We used a robust emotional face-matching task and acquired high-resolution structural images for morphometric analyses using voxel-based morphometry. The main results were positive associations of separation anxiety scores with amygdala reactivity to emotional faces as well as increased amygdala grey matter volumes. A functional connectivity analysis revealed positive associations between separation anxiety and functional coupling of the amygdala with areas involved in visual processes and attention, including several occipital and somatosensory areas. Taken together, the results suggest a higher emotional involvement in subjects with separation anxiety while watching negative facial expressions, and potentially secondary neuro-structural adaptive processes. These results could help to understand and treat (adult) separation anxiety.
doi:10.1093/scan/nsu055
PMCID: PMC4321627  PMID: 24752071
adult separation anxiety; fMRI; voxel-based morphometry; amygdala
15.  Alexithymia influences brain activation during emotion perception but not regulation 
Alexithymia is a psychological construct that can be divided into a cognitive and affective dimension. The cognitive dimension is characterized by difficulties in identifying, verbalizing and analysing feelings. The affective dimension comprises reduced levels of emotional experience and imagination. Alexithymia is widely regarded to arise from an impairment of emotion regulation. This is the first functional magnetic resonance imaging (fMRI) study to critically evaluate this by investigating the neural correlates of emotion regulation as a function of alexithymia levels. The aim of the current study was to investigate the neural correlates underlying the two alexithymia dimensions during emotion perception and emotion regulation. Using fMRI, we scanned 51 healthy subjects while viewing, reappraising or suppressing negative emotional pictures. The results support the idea that cognitive alexithymia, but not affective alexithymia, is associated with lower activation in emotional attention and recognition networks during emotion perception. However, in contrast with several theories, no alexithymia-related differences were found during emotion regulation (neither reappraisal nor suppression). These findings suggest that alexithymia may result from an early emotion processing deficit rather than compromised frontal circuits subserving higher-order emotion regulation processes.
doi:10.1093/scan/nsu056
PMCID: PMC4321628  PMID: 24760016
alexithymia; emotion processing; emotion regulation; amygdala; neuroimaging
16.  Age-related differences in function and structure of rSMG and reduced functional connectivity with DLPFC explains heightened emotional egocentricity bias in childhood 
Humans often judge others egocentrically, assuming that they feel or think similarly to themselves. Emotional egocentricity bias (EEB) occurs in situations when others feel differently to oneself. Using a novel paradigm, we investigated the neurocognitive mechanisms underlying the developmental capacity to overcome such EEB in children compared with adults. We showed that children display a stronger EEB than adults and that this correlates with reduced activation in right supramarginal gyrus (rSMG) as well as reduced coupling between rSMG and left dorsolateral prefrontal cortex (lDLPFC) in children compared with adults. Crucially, functional recruitment of rSMG was associated with age-related differences in cortical thickness of this region. Although in adults the mere presence of emotional conflict occurs between self and other recruited rSMG, rSMG-lDLPFC coupling was only observed when implementing empathic judgements. Finally, resting state analyses comparing connectivity patterns of rSMG with that of right temporoparietal junction suggested a unique role of rSMG for self-other distinction in the emotional domain for adults as well as for children. Thus, children’s difficulties in overcoming EEB may be due to late maturation of regions distinguishing between conflicting socio-affective information and relaying this information to regions necessary for implementing accurate judgments.
doi:10.1093/scan/nsu057
PMCID: PMC4321629  PMID: 24771281
functional and structural brain development; socio-affective development; emotional egocentricity bias; self–other distinction; supramarginal gyrus; dorsolateral prefrontal cortex
17.  Altered emotion regulation capacity in social phobia as a function of comorbidity 
Social phobia (SP) has been associated with amygdala hyperreactivity to fear-relevant stimuli. However, little is known about the neural basis of SP individuals’ capacity to downregulate their responses to such stimuli and how such regulation varies as a function of comorbid depression and anxiety. We completed an functional magnetic resonance imaging (fMRI) study wherein SP participants without comorbidity (n = 30), with comorbid depression (n = 18) and with comorbid anxiety (n = 19) and healthy controls (n = 15) were scanned while completing an affect labeling emotion regulation task. Individuals with SP as a whole exhibited a reversal of the pattern observed in healthy controls in that they showed upregulation of amygdala activity during affect labeling. However, subsequent analyses revealed a more complex picture based on comorbidity type. Although none of the SP subgroups showed the normative pattern of amygdala downregulation, it was those with comorbid depression specifically who showed significant upregulation. Effects could not be attributed to differences in task performance, amygdala reactivity or right ventral lateral prefrontal cortex (RVLPFC) engagement, but may stem from dysfunctional communication between amygdala and RVLPFC. Furthermore, the particularly altered emotion regulation seen in those with comorbid depression could not be fully explained by symptom severity or state anxiety. Results reveal altered emotion regulation in SP, especially when comorbid with depression.
doi:10.1093/scan/nsu058
PMCID: PMC4321630  PMID: 24813437
social anxiety; depression; affect labeling; functional magnetic resonance imaging
18.  ‘Why should I care?’ Challenging free will attenuates neural reaction to errors 
Whether human beings have free will has been a philosophical question for centuries. The debate about free will has recently entered the public arena through mass media and newspaper articles commenting on scientific findings that leave little to no room for free will. Previous research has shown that encouraging such a deterministic perspective influences behavior, namely by promoting cursory and antisocial behavior. Here we propose that such behavioral changes may, at least partly, stem from a more basic neurocognitive process related to response monitoring, namely a reduced error detection mechanism. Our results show that the error-related negativity, a neural marker of error detection, was reduced in individuals led to disbelieve in free will. This finding shows that reducing the belief in free will has a specific impact on error detection mechanisms. More generally, it suggests that abstract beliefs about intentional control can influence basic and automatic processes related to action control.
doi:10.1093/scan/nsu068
PMCID: PMC4321631  PMID: 24795441
free will; belief; response monitoring; error detection; error-related negativity
19.  Evidence for a role of the oxytocin system, indexed by genetic variation in CD38, in the social bonding effects of expressed gratitude 
Oxytocin is thought to play a central role in promoting close social bonds via influence on social interactions. The current investigation targeted interactions involving expressed gratitude between members of romantic relationships because recent evidence suggests gratitude and its expression provides behavioral and psychological ‘glue’ to bind individuals closer together. Specifically, we took a genetic approach to test the hypothesis that social interactions involving expressed gratitude would be associated with variation in a gene, CD38, which has been shown to affect oxytocin secretion. A polymorphism (rs6449182) that affects CD38 expression was significantly associated with global relationship satisfaction, perceived partner responsiveness and positive emotions (particularly love) after lab-based interactions, observed behavioral expression of gratitude toward a romantic partner in the lab, and frequency of expressed gratitude in daily life. A separate polymorphism in CD38 (rs3796863) previously associated with plasma oxytocin levels and social engagement was also associated with perceived responsiveness in the benefactor after an expression of gratitude. The combined influence of the two polymorphisms was associated with a broad range of gratitude-related behaviors and feelings. The consistent pattern of findings suggests that the oxytocin system is associated with solidifying the glue that binds adults into meaningful and important relationships.
doi:10.1093/scan/nst182
PMCID: PMC4249462  PMID: 24396004
close relationships; gratitude; emotion expression; CD38; oxytocin
20.  Encoding-related EEG oscillations during memory formation are modulated by mood state 
Mood states have a strong impact on how we process incoming information. It has been proposed that positive mood facilitates elaborative, relational encoding, whereas negative mood promotes a more careful, stimulus-driven encoding style. Previous electrophysiological studies have linked successful information encoding to power increases in slow (<8 Hz) delta/theta and fast (>30 Hz) gamma oscillations, as well as to power decreases in midrange (8–30 Hz) alpha/beta oscillations. Whether different mood states modulate encoding-related oscillations has not been investigated yet. In order to address this question, we used an experimental mood induction procedure and recorded electroencephalograms from 20 healthy participants while they performed a free recall memory task after positive and negative mood induction. We found distinct oscillatory patterns in positive and negative mood. Successful encoding in positive mood was accompanied by widespread power increases in the delta band, whereas encoding success in negative mood was specifically accompanied by frontal power decreases in the beta band. On the behavioral level, memory performance was enhanced in positive mood. Our findings show that mood differentially modulates the neural correlates of successful information encoding and thus contribute to an understanding of how mood shapes different processing styles.
doi:10.1093/scan/nst184
PMCID: PMC4249463  PMID: 24464848
subsequent memory paradigm; mood induction; EEG; delta ERS; beta ERD
21.  The brain network reflecting bodily self-consciousness: a functional connectivity study 
Several brain regions are important for processing self-location and first-person perspective, two important aspects of bodily self-consciousness. However, the interplay between these regions has not been clarified. In addition, while self-location and first-person perspective in healthy subjects are associated with bilateral activity in temporoparietal junction (TPJ), disturbed self-location and first-person perspective result from damage of only the right TPJ. Identifying the involved brain network and understanding the role of hemispheric specializations in encoding self-location and first-person perspective, will provide important information on system-level interactions neurally mediating bodily self-consciousness. Here, we used functional connectivity and showed that right and left TPJ are bilaterally connected to supplementary motor area, ventral premotor cortex, insula, intraparietal sulcus and occipitotemporal cortex. Furthermore, the functional connectivity between right TPJ and right insula had the highest selectivity for changes in self-location and first-person perspective. Finally, functional connectivity revealed hemispheric differences showing that self-location and first-person perspective modulated the connectivity between right TPJ, right posterior insula, and right supplementary motor area, and between left TPJ and right anterior insula. The present data extend previous evidence on healthy populations and clinical observations in neurological deficits, supporting a bilateral, but right-hemispheric dominant, network for bodily self-consciousness.
doi:10.1093/scan/nst185
PMCID: PMC4249464  PMID: 24396007
self-location; first-person perspective; temporoparietal junction; insula; multisensory integration
22.  Neural disruption to theory of mind predicts daily social functioning in individuals at familial high-risk for schizophrenia 
Theory-of-mind (ToM) ability is foundational for successful social relationships, and dependent on a neurocognitive system, which includes temporoparietal junction and medial prefrontal cortex. Schizophrenia is associated with ToM impairments, and initial studies demonstrate similar, though more subtle deficits, in unaffected first-degree relatives, indicating that ToM deficits are a potential biomarker for the disorder. Importantly, the social consequences of ToM deficits could create an additional vulnerability factor for individuals at familial high risk (FHR). However, behavioral studies of ToM are inconsistent and virtually nothing is known about the neural basis of ToM in FHR or the relationship between ToM and social functioning. Here, FHR and non-FHR control participants underwent functional MRI scanning while reasoning about a story character’s thoughts, emotions or physical appearance. Afterwards, participants completed a 28-day online ‘daily-diary’ questionnaire in which they reported daily social interactions and degree of ToM reasoning. FHR participants demonstrated less neural activity in bilateral temporoparietal junction when reasoning about thoughts and emotions. Moreover, across all participants, the degree of neural activity during ToM reasoning predicted several aspects of daily social behavior. Results suggest that vulnerability for schizophrenia is associated with neurocognitive deficits in ToM and the degree of deficit is related to day-to-day social functioning.
doi:10.1093/scan/nst186
PMCID: PMC4249465  PMID: 24396009
theory of mind; social functioning; schizophrenia; familial high risk; fMRI
23.  Three stages of emotional word processing: an ERP study with rapid serial visual presentation 
Rapid responses to emotional words play a crucial role in social communication. This study employed event-related potentials to examine the time course of neural dynamics involved in emotional word processing. Participants performed a dual-target task in which positive, negative and neutral adjectives were rapidly presented. The early occipital P1 was found larger when elicited by negative words, indicating that the first stage of emotional word processing mainly differentiates between non-threatening and potentially threatening information. The N170 and the early posterior negativity were larger for positive and negative words, reflecting the emotional/non-emotional discrimination stage of word processing. The late positive component not only distinguished emotional words from neutral words, but also differentiated between positive and negative words. This represents the third stage of emotional word processing, the emotion separation. Present results indicated that, similar with the three-stage model of facial expression processing; the neural processing of emotional words can also be divided into three stages. These findings prompt us to believe that the nature of emotion can be analyzed by the brain independent of stimulus type, and that the three-stage scheme may be a common model for emotional information processing in the context of limited attentional resources.
doi:10.1093/scan/nst188
PMCID: PMC4249467  PMID: 24526185
emotional word; three stages; rapid serial visual presentation; event-related potential
24.  Individual differences in anxiety predict neural measures of visual working memory for untrustworthy faces 
When facing strangers, one of the first evaluations people perform is to implicitly assess their trustworthiness. However, the underlying processes supporting trustworthiness appraisal are poorly understood. We hypothesized that visual working memory (VWM) maintains online face representations that are sensitive to physical cues of trustworthiness, and that differences among individuals in representing untrustworthy faces are associated with individual differences in anxiety. Participants performed a change detection task that required encoding and maintaining for a short interval the identity of one face parametrically manipulated to be either trustworthy or untrustworthy. The sustained posterior contralateral negativity (SPCN), an event-related component (ERP) time-locked to the onset of the face, was used to index the resolution of face representations in VWM. Results revealed greater SPCN amplitudes for trustworthy faces when compared with untrustworthy faces, indicating that VWM is sensitive to physical cues of trustworthiness, even in the absence of explicit trustworthiness appraisal. In addition, differences in SPCN amplitude between trustworthy and untrustworthy faces correlated with participants’ anxiety, indicating that healthy college students with sub-clinical high anxiety levels represented untrustworthy faces in greater detail compared with students with sub-clinical low anxiety levels. This pattern of findings is discussed in terms of the high flexibility of aversive/avoidance and appetitive/approach motivational systems.
doi:10.1093/scan/nst189
PMCID: PMC4249468  PMID: 24493843
trustworthiness; visual working memory; anxiety; event-related potentials; faces
25.  A comparison of neural correlates underlying social cognition in Klinefelter syndrome and autism 
Klinefelter syndrome (KS) is a genetic syndrome characterized by the presence of an extra X chromosome that appears to increase the risk of psychopathology, such as autism symptoms. This study used functional magnetic resonance imaging to determine underlying mechanisms related to this risk, with the aim of gaining insight into neural mechanisms behind social-cognitive dysfunction in KS and autism, and understanding similarities and differences in social information processing deficits. Fourteen boys with KS, seventeen boys with autism spectrum disorders (ASD) and nineteen non-clinical male controls aged 10–18 years were scanned while matching and labeling facial expressions (i.e. face processing and affect labeling, respectively). No group differences in neural activation were found during face processing. However, during affect labeling, the ASD group showed increased activation in the amygdala compared with controls, while the KS group showed increased activation in frontal areas compared with both controls and the ASD group. No group differences in task performance were found. Although behavioral symptoms of social dysfunction appear similar both in boys with KS and ASD, this is the first study to demonstrate different underlying etiologies. These results may aid in identifying different pathways to autism symptoms, which may help understanding variability within the ASD spectrum.
doi:10.1093/scan/nst190
PMCID: PMC4249469  PMID: 24396006
Klinefelter; autism; social cognition; fMRI; facial affect processing

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