Spatial gradient in value representation along the medial prefrontal cortex reflects individual differences in prosociality.

Despite the importance of valuing another person's welfare for prosocial behavior, currently we have only a limited understanding of how these values are represented in the brain and, more importantly, how they give rise to individual variability in prosociality. In the present study, participants underwent functional magnetic resonance imaging while performing a prosocial learning task in which they could choose to benefit themselves and/or another person. Choice behavior indicated that participants valued the welfare of another person, although less so than they valued their own welfare. Neural data revealed a spatial gradient in activity within the medial prefrontal cortex (MPFC), such that ventral parts predominantly represented self-regarding values and dorsal parts predominantly represented other-regarding values. Importantly, compared with selfish individuals, prosocial individuals showed a more gradual transition from self-regarding to other-regarding value signals in the MPFC and stronger MPFC-striatum coupling when they made choices for another person rather than for themselves. The present study provides evidence of neural markers reflecting individual differences in human prosociality.

Structural covariance networks of the dorsal anterior insula predict females' individual differences in empathic responding.

Previous functional imaging studies have shown key roles of the dorsal anterior insula (dAI) and anterior midcingulate cortex (aMCC) in empathy for the suffering of others. The current study mapped structural covariance networks of these regions and assessed the relationship between networks and individual differences in empathic responding in 94 females. Individual differences in empathy were assessed through average state measures in response to a video task showing others' suffering, and through questionnaire-based trait measures of empathic concern. Overall, covariance patterns indicated that dAI and aMCC are principal hubs within prefrontal, temporolimbic, and midline structural covariance networks. Importantly, participants with high empathy state ratings showed increased covariance of dAI, but not aMCC, to prefrontal and limbic brain regions. This relationship was specific for empathy and could not be explained by individual differences in negative affect ratings. Regarding questionnaire-based empathic trait measures, we observed a similar, albeit weaker modulation of dAI covariance, confirming the robustness of our findings. Our analysis, thus, provides novel evidence for a specific contribution of frontolimbic structural covariance networks to individual differences in social emotions beyond negative affect.

Functional neural plasticity and associated changes in positive affect after compassion training.

The development of social emotions such as compassion is crucial for successful social interactions as well as for the maintenance of mental and physical health, especially when confronted with distressing life events. Yet, the neural mechanisms supporting the training of these emotions are poorly understood. To study affective plasticity in healthy adults, we measured functional neural and subjective responses to witnessing the distress of others in a newly developed task (Socio-affective Video Task). Participants' initial empathic responses to the task were accompanied by negative affect and activations in the anterior insula and anterior medial cingulate cortex--a core neural network underlying empathy for pain. Whereas participants reacted with negative affect before training, compassion training increased positive affective experiences, even in response to witnessing others in distress. On the neural level, we observed that, compared with a memory control group, compassion training elicited activity in a neural network including the medial orbitofrontal cortex, putamen, pallidum, and ventral tegmental area--brain regions previously associated with positive affect and affiliation. Taken together, these findings suggest that the deliberate cultivation of compassion offers a new coping strategy that fosters positive affect even when confronted with the distress of others.

Intentional social distance regulation alters affective responses towards victims of violence: an FMRI study.

We used functional magnetic resonance imaging (fMRI) to investigate brain processes underlying control of emotional responses towards a person in distress by cognitive social distance modulation. fMRI and peripheral physiological responses (startle response and electrodermal activity) were recorded from 24 women while they watched victim-offender scenes and modulated their social distance to the victim by cognitive reappraisal. We found that emotional responses, including startle eyeblink and amygdala responses, can effectively be modulated by social distance modulation. Furthermore, our data provide evidence that activity in the dorsomedial prefrontal cortex (dmPFC) and the anterior paracingulate cortex (aPCC), two brain regions that have previously been associated with brain processes related to distant and close others, is differentially modulated by intentional social distance modulation: activity in the dmPFC increased with increasing disengagement from the victim and activity in the aPCC increased with increasing engagement with the victim. We suggest that these two regions play opposing roles in cognitive modulation of social distance and affective responses towards persons in distress that enable the adaptive and flexible social behavior observed in humans.

Virtual faces as a tool to study emotion recognition deficits in schizophrenia.

Studies investigating emotion recognition in patients with schizophrenia predominantly presented photographs of facial expressions. Better control and higher flexibility of emotion displays could be afforded by virtual reality (VR). VR allows the manipulation of facial expression and can simulate social interactions in a controlled and yet more naturalistic environment. However, to our knowledge, there is no study that systematically investigated whether patients with schizophrenia show the same emotion recognition deficits when emotions are expressed by virtual as compared to natural faces. Twenty schizophrenia patients and 20 controls rated pictures of natural and virtual faces with respect to the basic emotion expressed (happiness, sadness, anger, fear, disgust, and neutrality). Consistent with our hypothesis, the results revealed that emotion recognition impairments also emerged for emotions expressed by virtual characters. As virtual in contrast to natural expressions only contain major emotional features, schizophrenia patients already seem to be impaired in the recognition of basic emotional features. This finding has practical implication as it supports the use of virtual emotional expressions for psychiatric research: the ease of changing facial features, animating avatar faces, and creating therapeutic simulations makes validated artificial expressions perfectly suited to study and treat emotion recognition deficits in schizophrenia.

The temporal dynamics of insula activity to disgust and happy facial expressions: a magnetoencephalography study.

The insula has consistently been shown to be involved in processing stimuli that evoke the emotional response of disgust. Recently, its specificity for processing disgust has been challenged and a broader role of the insula in the representation of interoceptive information has been suggested. Studying the temporal dynamics of insula activation during emotional processing can contribute valuable information pertaining to this issue. Few studies have addressed the insula's putative specificity to disgust and the dynamics of its underlying neural processes. In the present study, neuromagnetic responses of 13 subjects performing an emotional continuous performance task (CPT) to faces with disgust, happy, and neutral expressions were obtained. Magnetic field tomography extracted the time course of bilateral insula activities. Right insula activation was stronger to disgust and happy than neutral facial expressions at about 200 ms after stimulus onset. Later only at about 350 ms after stimulus onset the right insula was activated stronger to disgust than happy facial expressions. Thus, the early right insula response reflects activation to emotionally arousing stimuli regardless of valence, and the later right insula response differentiates disgust from happy facial expressions. Behavioral performance but not the insula activity differed between 100 ms and 1000 ms presentation conditions. Present findings support the notion that the insula is involved in the representation of interoceptive information.

Towards a human self-regulation system: Common and distinct neural signatures of emotional and behavioural control.

Self-regulation refers to controlling our emotions and actions in the pursuit of higher-order goals. Although research suggests commonalities in the cognitive control of emotion and action, evidence for a shared neural substrate is scant and largely circumstantial. Here we report on two large-scale meta-analyses of human neuroimaging studies on emotion or action control, yielding two fronto-parieto-insular networks. The networks' overlap, however, was restricted to four brain regions: posteromedial prefrontal cortex, bilateral anterior insula, and right temporo-parietal junction. Conversely, meta-analytic contrasts revealed major between-network differences, which were independently corroborated by clustering domain-specific regions based on their intrinsic functional connectivity, as well as by functionally characterizing network sub-clusters using the BrainMap database for quantitative forward and reverse inference. Collectively, our analyses identified a core system for implementing self-control across emotion and action, beyond which, however, either regulation facet appears to rely on broadly similar yet distinct subnetworks. These insights into the neurocircuitry subserving affective and executive facets of self-control suggest both processing commonalities and differences between the two aspects of human self-regulation.

Prefrontal gamma-band activity distinguishes between sound durations.

The present study used magnetoencephalography to assess the cortical representation of brief sound durations during a short-term memory task. Twelve subjects were instructed to memorize sounds S1 with durations of either 100 or 200 ms during an 800-ms delay phase. Subsequently, they had to judge whether the duration of a probe sound S2 matched the memorized stimulus. Statistical probability mapping of oscillatory signals revealed several components of gamma-band activity (GBA) over prefrontal cortex. A first component with a center frequency of 40 Hz responded more strongly to longer than shorter sounds during the encoding of S1. During the subsequent delay phase, shorter and longer durations were associated with topographically and spectrally distinct GBA enhancements at 71 and 80 Hz, respectively. S2 was again associated with stronger oscillatory activation for longer than shorter sounds at approximately 72 Hz. Non matching compared with matching S1-S2 pairs elicited an additional approximately 66 Hz GBA component peaking at approximately 200 ms after the offset of S2. The analysis of magnetoencephalographic GBA thus served to identify prefrontal network components underlying the representation of different sound durations during the various phases of a delayed matching-to-sample task.

The multiple facets of empathy: a survey of theory and evidence.

Empathy is the ability to perceive and understand other people's emotions and to react appropriately. This ability is a necessary prerequisite for successful interpersonal interaction. Empathy is a multifaceted construct including low-level mechanisms like emotional contagion as well as high-level processes like perspective-taking. The ability to empathize varies between individuals and is considered a stable personality trait: some people are generally more successful in empathizing than others. In this chapter we will first present different conceptualizations of the construct of empathy, and refer to empathy-regulating processes as well as to the relationship between empathy and social behavior. Then, we will review peripheral physiological and brain imaging studies pertaining to low- and high-level empathic processes, empathy-modulating processes, and the link between empathy and social behavior. Further, we will present evidence regarding interindividual differences in these processes as an important source of information for solving the conundrum of how the comprehension of others' emotions is achieved by our brains.

Effects of memory load on cortical oscillatory activity during auditory pattern working memory.

The present magnetoencephalography study investigated memory load-dependent changes in cortical oscillatory activity during a modified auditory version of the Sternberg paradigm. Twelve subjects were presented with test stimulus sets of 1-3 syllables spoken in a natural female voice. After an 800-ms delay period, a probe syllable was presented and subjects had to judge whether the probe had been included in the preceding test set. Compared to a non-memory-control task, memory trials elicited an increase of beta activity over right temporal regions and an increase of alpha activity over right middle prefrontal cortex at the end of the delay phase. Monotonic increases in spectral amplitude as a function of memory load were revealed for the beta band over right temporal sensors and the alpha band over right frontal sensors during the delay period. The results demonstrate the relevance of both beta and alpha oscillations for the memorization of multiple stimuli. The former may be associated with the representation of task-relevant stimulus features, while the latter may reflect the top-down control of these representations.

The brain's functional network architecture reveals human motives.

Goal-directed human behaviors are driven by motives. Motives are, however, purely mental constructs that are not directly observable. Here, we show that the brain's functional network architecture captures information that predicts different motives behind the same altruistic act with high accuracy. In contrast, mere activity in these regions contains no information about motives. Empathy-based altruism is primarily characterized by a positive connectivity from the anterior cingulate cortex (ACC) to the anterior insula (AI), whereas reciprocity-based altruism additionally invokes strong positive connectivity from the AI to the ACC and even stronger positive connectivity from the AI to the ventral striatum. Moreover, predominantly selfish individuals show distinct functional architectures compared to altruists, and they only increase altruistic behavior in response to empathy inductions, but not reciprocity inductions.

The Obsessive-Compulsive Inventory: development and validation of a short version.

This article reports on the development of a revised version of the Obsessive-Compulsive Inventory (OCI; E. B. Foa, M. J. Kozak, P. Salkovskis, M. E. Coles, & N. Amir, 1998), a psychometrically sound, theoretically driven, self-report measure. The revised OCI (OCI-R) improves on the parent version in 3 ways: It eliminates the redundant frequency scale, simplifies the scoring of the subscales, and reduces overlap across subscales. The reliability and validity of the OCI-R were examined in 215 patients with obsessive-compulsive disorder (OCD), 243 patients with other anxiety disorders, and 677 nonanxious individuals. The OCI-R, which contains 18 items and 6 subscales, has retained excellent psychometric properties. The OCI-R and its subscales differentiated well between individuals with and without OCD. Receiver operating characteristic (ROC) analyses demonstrated the usefulness of the OCI-R as a diagnostic tool for screening patients with OCD, utilizing empirically derived cutscores.

Differential pattern of functional brain plasticity after compassion and empathy training.

Although empathy is crucial for successful social interactions, excessive sharing of others' negative emotions may be maladaptive and constitute a source of burnout. To investigate functional neural plasticity underlying the augmentation of empathy and to test the counteracting potential of compassion, one group of participants was first trained in empathic resonance and subsequently in compassion. In response to videos depicting human suffering, empathy training, but not memory training (control group), increased negative affect and brain activations in anterior insula and anterior midcingulate cortex-brain regions previously associated with empathy for pain. In contrast, subsequent compassion training could reverse the increase in negative effect and, in contrast, augment self-reports of positive affect. In addition, compassion training increased activations in a non-overlapping brain network spanning ventral striatum, pregenual anterior cingulate cortex and medial orbitofrontal cortex. We conclude that training compassion may reflect a new coping strategy to overcome empathic distress and strengthen resilience.

Harder than expected: increased conflict in clearly disadvantageous delayed choices in a computer game.

When choosing between immediate and temporally delayed goods, people sometimes decide disadvantageously. Here, we aim to provide process-level insight into differences between individually determined advantageous and disadvantageous choices. Participants played a computer game, deciding between two different rewards of varying size and distance by moving an agent towards the chosen reward. We calculated individual models of advantageous choices and characterized the decision process by analyzing mouse movements. The larger amount of participants' choices was classified as advantageous and the disadvantageous choices were biased towards choosing sooner/smaller rewards. The deflection of mouse movements indicated more conflict in disadvantageous choices compared with advantageous choices when the utilities of the options differed clearly. Further process oriented analysis revealed that disadvantageous choices were biased by a tendency for choice-repetition and an undervaluation of the value information in favour of the delay information, making rather simple choices harder than could be expected from the properties of the decision situation.

Short-term compassion training increases prosocial behavior in a newly developed prosocial game.

Compassion has been suggested to be a strong motivator for prosocial behavior. While research has demonstrated that compassion training has positive effects on mood and health, we do not know whether it also leads to increases in prosocial behavior. We addressed this question in two experiments. In Experiment 1, we introduce a new prosocial game, the Zurich Prosocial Game (ZPG), which allows for repeated, ecologically valid assessment of prosocial behavior and is sensitive to the influence of reciprocity, helping cost, and distress cues on helping behavior. Experiment 2 shows that helping behavior in the ZPG increased in participants who had received short-term compassion training, but not in participants who had received short-term memory training. Interindividual differences in practice duration were specifically related to changes in the amount of helping under no-reciprocity conditions. Our results provide first evidence for the positive impact of short-term compassion training on prosocial behavior towards strangers in a training-unrelated task.

Recognition profile of emotions in natural and virtual faces.

BACKGROUND: Computer-generated virtual faces become increasingly realistic including the simulation of emotional expressions. These faces can be used as well-controlled, realistic and dynamic stimuli in emotion research. However, the validity of virtual facial expressions in comparison to natural emotion displays still needs to be shown for the different emotions and different age groups. METHODOLOGY/PRINCIPAL FINDINGS: Thirty-two healthy volunteers between the age of 20 and 60 rated pictures of natural human faces and faces of virtual characters (avatars) with respect to the expressed emotions: happiness, sadness, anger, fear, disgust, and neutral. Results indicate that virtual emotions were recognized comparable to natural ones. Recognition differences in virtual and natural faces depended on specific emotions: whereas disgust was difficult to convey with the current avatar technology, virtual sadness and fear achieved better recognition results than natural faces. Furthermore, emotion recognition rates decreased for virtual but not natural faces in participants over the age of 40. This specific age effect suggests that media exposure has an influence on emotion recognition. CONCLUSIONS/SIGNIFICANCE: Virtual and natural facial displays of emotion may be equally effective. Improved technology (e.g. better modelling of the naso-labial area) may lead to even better results as compared to trained actors. Due to the ease with which virtual human faces can be animated and manipulated, validated artificial emotional expressions will be of major relevance in future research and therapeutic applications.

The involvement of ipsilateral temporoparietal cortex in tactile pattern working memory as reflected in beta event-related desynchronization.

Cortical oscillatory activity in various frequency bands has been shown to reflect working memory processes operating on visual and auditory stimulus information. Here we use magnetoencephalography to investigate cortical oscillatory activity related to working memory for tactile patterns. Right-handed subjects made same-different judgements on two dot patterns sequentially applied with a 3-s delay to the right middle fingertip. Spectral analysis revealed beta desynchronization (17+/-2.5 Hz) at contralateral postcentral and ipsilateral temporoparietal regions preceding and during the presentation of both tactile stimuli as well as during the early and late delay periods. Whereas contralateral beta desynchronization preceding tactile stimulation may reflect anticipation of incoming stimuli, ipsilateral beta desynchronization may underlie working memory maintenance of tactile patterns. The later hypothesis is supported by a significant positive correlation between subjects' performance and the amplitude of ipsilateral beta desynchronization 800 ms to 500 ms before the onset of the second pattern stimulus. Thus, our results suggest that ipsilateral temporoparietal cortex contributes to the maintenance of tactile pattern information in working memory.

BOLD adaptation in vibrotactile stimulation: neuronal networks involved in frequency discrimination.

The present functional magnetic resonance imaging (fMRI) study investigated human brain regions subserving the discrimination of vibrotactile frequency. An event-related adaptation paradigm was used in which blood-oxygen-level-dependent (BOLD) responses are lower to same compared with different pairs of stimuli (BOLD adaptation). This adaptation effect serves as an indicator for feature-specific responding of neuronal subpopulations. Subjects had to discriminate two vibrotactile stimuli sequentially applied with a delay of 600 ms to their left middle fingertip. The stimulus frequency was in the flutter range of 18-26 Hz. In half of the trials, the two stimuli possessed identical frequency (same), whereas in the other half, a frequency difference of +/-2 Hz was used (diff). As a result, BOLD adaptation was observed in the contralateral primary somatosensory cortex (S1), precentral gyrus, superior temporal gyrus (STG); ipsilateral insula as well as bilateral secondary somatosensory cortex and supplementary motor area. When statistically comparing the BOLD time courses between same and diff trials in these cortical areas, it was found that the vibrotactile BOLD adaptation is initiated in the contralateral S1 and STG simultaneously. These findings suggest that the cortical areas responsive to the frequency difference between two serially presented stimuli sequentially process the frequency of a vibrotactile stimulus and constitute a putative neuronal network underlying human vibrotactile frequency discrimination.