Load-related brain activation predicts spatial working memory performance in youth aged 9–12 and is associated with executive function at earlier ages

Spatial working memory is a central cognitive process that matures through adolescence in conjunction with major changes in brain function and anatomy. Here we focused on late childhood and early adolescence to more closely examine the neural correlates of performance variability during this important transition period. Using a modified spatial 1-back task with two memory load conditions in an fMRI study, we examined the relationship between load-dependent neural responses and task performance in a sample of 39 youth aged 9–12 years. Our data revealed that between-subject differences in task performance was predicted by load-dependent deactivation in default network regions, including the ventral anterior cingulate cortex (vACC) and posterior cingulate cortex (PCC). Although load-dependent increases in activation in prefrontal and posterior parietal regions were only weakly correlated with performance, increased prefrontal–parietal coupling was associated with better performance. Furthermore, behavioral measures of executive function from as early as age 3 predicted current load-dependent deactivation in vACC and PCC. These findings suggest that both task positive and task negative brain activation during spatial working memory contributed to successful task performance in late childhood/early adolescence. This may serve as a good model for studying executive control deficits in developmental disorders.     

Neural representation of maternal face processing: a functional magnetic resonance imaging study.

OBJECTIVE: The mother-child relationship may have important implications for emotional development and adult psychopathology. The objective of this study was to examine brain responses to processing maternal faces in healthy adult women. METHODS: Ten healthy adult female volunteers with adequate early-life maternal care and a normal relationship with their living mothers participated in the study. Functional magnetic resonance imaging was used to examine brain responses to pictures of the subject's mother, a close female friend, and 2 age-matched female strangers during passive viewing, valence (emotional), and salience (self-relevance) evaluations. RESULTS: The main contrast of mother, compared with all others (that is, friend and older and younger strangers), demonstrated the following: first, significant activation in the left posterior cingulate cortex-precuneus (PCC-Pcu), collapsed across all tasks; second, right ventromedial prefrontal cortex-anterior cingulate cortex (VMPFC-ACC) activation during the valence condition; and third, left dorsolateral prefrontal cortex (DLPFC) activation during the salience condition. In the region-of-interest analyses, the VMPFC-ACC and DLPFC showed significant activations in response to mothers' faces and deactivation in response to control faces. Among the 3 regions, only VMPFC-ACC activity distinguished the unique processing of one's own mother's face from that of a close friend. PCC-Pcu activations demonstrate a graded response (mother > friend > strangers) and, further, demonstrated differential response with respect to mothering style. CONCLUSIONS: The activation in prefrontal and cingulate cortices related to maternal face processing is consistent with their implicated roles in mother-infant interactions, personal familiarity, and emotional and self-relevant processing. These findings suggest a neural basis for maternal attachment and propose a focus for future studies aimed at investigating the impact of disrupted maternal attachment on emotional development.     

fMRI reveals alteration of spatial working memory networks across adolescence.

Recent studies have described neuromaturation and cognitive development across the lifespan, yet few neuroimaging studies have investigated task-related alterations in brain activity during adolescence. We used functional magnetic resonance imaging (fMRI) to examine brain response to a spatial working memory (SWM) task in 49 typically developing adolescents (25 females and 24 males; ages 12-17). No gender or age differences were found for task performance during SWM. However, age was positively associated with SWM brain response in left prefrontal and bilateral inferior posterior parietal regions. Age was negatively associated with SWM activation in bilateral superior parietal cortex. Gender was significantly associated with SWM response; females demonstrated diminished anterior cingulate activation and males demonstrated greater response in frontopolar cortex than females. Our findings indicate that the frontal and parietal neural networks involved in spatial working memory change over the adolescent age range and are further influenced by gender. These changes may represent evolving mnemonic strategies subserved by ongoing adolescent brain development.     

Altered resting state networks in mild cognitive impairment and mild Alzheimer's disease: an fMRI study

Activity and reactivity of the default mode network in the brain was studied using functional magnetic resonance imaging (fMRI) in 28 nondemented individuals with mild cognitive impairment (MCI), 18 patients with mild Alzheimer's disease (AD), and 41 healthy elderly controls (HC). The default mode network was interrogated by means of decreases in brain activity, termed deactivations, during a visual encoding task and during a nonspatial working memory task. Deactivation was found in the default mode network involving the anterior frontal, precuneus, and posterior cingulate cortex. MCI patients showed less deactivation than HC, but more than AD. The most pronounced differences between MCI, HC, and AD occurred in the very early phase of deactivation, reflecting the reactivity and adaptation of the network. The default mode network response in the anterior frontal cortex significantly distinguished MCI from both HC (in the medial frontal) and AD (in the anterior cingulate cortex). The response in the precuneus could only distinguish between patients and HC, not between MCI and AD. These findings may be consistent with the notion that MCI is a transitional state between healthy aging and dementia and with the proposed early changes in MCI in the posterior cingulate cortex and precuneus. These findings suggest that altered activity in the default mode network may act as an early marker for AD pathology.     

Direct replication of task‐dependent neural activation patterns during sadness introspection in two independent adolescent samples

Functional neuroimaging results need to replicate to inform sound models of human social cognition and its neural correlates. Introspection, the capacity to reflect on one's thoughts and feelings, is one process required for normative social cognition and emotional functioning. Engaging in introspection draws on a network of brain regions including medial prefrontal cortex (mPFC), posterior cingulate cortex (PCC), middle temporal gyri (MTG), and temporoparietal junction (TPJ). Maturation of these regions during adolescence mirrors the behavioral advances seen in adolescent social cognition, but the neural correlates of introspection in adolescence need to replicate to confirm their generalizability and role as a possible mechanism. The current study investigated whether reflecting upon one's own feelings of sadness would activate and replicate similar brain regions in two independent samples of adolescents. Participants included 156 adolescents (50% female) from the California Families Project and 119 adolescent girls from the Pittsburgh Girls Study of Emotion. All participants completed the Emotion Regulation Questionnaire (ERQ) and underwent a functional magnetic resonance imaging scan while completing the same facial emotion‐processing task at age 16–17 years. Both samples showed similar whole‐brain activation patterns when engaged in sadness introspection and when judging a nonemotional facial feature. Whole‐brain activation was unrelated to ERQ scores in both samples. Neural responsivity to task manipulations replicated in regions recruited for socio‐emotional (mPFC, PCC, MTG, TPJ) and attention (dorsolateral PFC, precentral gyri, superior occipital gyrus, superior parietal lobule) processing. These findings demonstrate robust replication of neural engagement during sadness introspection in two independent adolescent samples.     

Serotonin transporter genotype modulates cognitive reappraisal of negative emotions: a functional magnetic resonance imaging study

A functional polymorphism within the serotonin transporter gene (5-HTTLPR) has been reported to modulate emotionality and risk for affective disorders. The short (S) allele has less functional efficacy than the long (L) allele and has been associated with enhanced emotional reactivity. One possible contributing factor to the high emotionality in S carriers may be inefficient use of cognitive strategies such as reappraisal to regulate emotional responses. The aim of the present study was to test whether the 5-HTTLPR genotype modulates the neural correlates of emotion regulation. To determine neural differences between S and L allele carriers during reappraisal of negative emotions, 15 homozygous S (S′/S′) and 15 homozygous L (L′/L′) carriers underwent functional magnetic resonance imaging (fMRI), while performing an instructed emotion regulation task including downregulation, upregulation and passive viewing of negative emotional pictures. Compared to L′/L′ allele carriers, subjects who carry the S′/S′ allele responded with lower posterior insula and prefrontal brain activation during passive perception of negative emotional information but showed greater prefrontal activation and anterior insula activation during down- and upregulation of negative emotional responses. The current results support and extend previous findings of enhanced emotionality in S carriers by providing additional evidence of 5-HTTLPR modulation of volitional emotion regulation.     

The influence of emotional associations on the neural correlates of semantic priming

Emotions influence our everyday life in several ways. With the present study, we wanted to examine the impact of emotional information on neural correlates of semantic priming, a well-established technique to investigate semantic processing. Stimuli were presented with a short SOA of 200 ms as subjects performed a lexical decision task during fMRI measurement. Seven experimental conditions were compared: positive/negative/neutral related, positive/negative/neutral unrelated, nonwords (all words were nouns). Behavioral data revealed a valence specific semantic priming effect (i.e., unrelated > related) only for neutral and positive related word pairs. On a neural level, the comparison of emotional over neutral relations showed activation in left anterior medial frontal cortex, superior frontal gyrus, and posterior cingulate. Interactions for the different relations were located in left anterior part of the medial frontal cortex, cingulate regions, and right hippocampus (positive > neutral + negative) and left posterior part of medial frontal cortex (negative > neutral + positive). The results showed that emotional information have an influence on semantic association processes. While positive and neutral information seem to share a semantic network, negative relations might induce compensatory mechanisms that inhibit the spread of activation between related concepts. The neural correlates highlighted a distributed neural network, primarily involving attention, memory and emotion related processing areas in medial fronto-parietal cortices. The differentiation between anterior (positive) and posterior part (negative) of the medial frontal cortex was linked to the type of affective manipulation with more cognitive demands being involved in the automatic processing of negative information.     

Neural correlates of adolescents’ viewing of parents’ and peers’ emotions: Associations with risk-taking behavior and risky peer affiliations

Social reorientation from parents to same-age peers is normative in adolescence, but the neural correlates of youths’ socioemotional processing of parents and peers have not been explored. In the current study, 22 adolescents (average age 16.98) underwent neuroimaging (functional magnetic resonance imaging) while viewing and rating emotions shown in brief video clips featuring themselves, their parents, or an unfamiliar peer. Viewing self vs. other and parents vs. the peer activated regions in the medial prefrontal cortex, replicating prior findings that this area responds to self-relevant stimuli, including familiar and not just similar others. Viewing the peer compared with parents elicited activation in posterior ‘mentalizing’ structures, the precuneus, posterior cingulate cortex (PCC), bilateral posterior superior temporal sulcus and right temporoparietal junction, as well as the ventral striatum and bilateral amygdala and hippocampus. Relative activations in the PCC and precuneus to the peer vs. the parent were related both to reported risk-taking behavior and to affiliations with more risk-taking peers. The results suggest neural correlates of the adolescent social reorientation toward peers and away from parents that may be associated with adolescents’ real-life risk-taking behaviors and social relationships.     

Local glutamate in cingulate cortex subregions differentially correlates with affective network activations during face perception

The cingulate cortex is involved in emotion recognition/perception and regulation. Rostral and caudal subregions belong to different brain networks with distinct roles in affective perception. Despite recent accounts of the relevance of cingulate cortex glutamate (Glu) on blood‐oxygen‐level‐dependent (BOLD) responses, the specificity of the subregional Glu levels during emotional tasks remains unclear. Seventy‐two healthy participants (age = 27.33 ± 6.67, 32 women) performed an affective face‐matching task and underwent magnetic resonance spectroscopy (MRS) at 7 Tesla. Correlations between the BOLD response during emotion perception and Glu concentration in the pregenual anterior cingulate cortex (pgACC) and anterior midcingulate cortex (aMCC) were compared on a whole‐brain level. Post hoc specificity of the association with an affect was assessed. Lower Glu in the pgACC correlated with stronger activation differences between negative and positive faces in the left inferior and superior frontal gyrus (L IFG and L SFG). In contrast, lower Glu in the aMCC correlated with BOLD contrasts in the posterior cingulate cortex (PCC). Furthermore, negative face detection was associated with prolonged response time (RT). Our results demonstrate a subregion‐specific involvement of cingulate cortex Glu in interindividual differences during viewing of affective facial expressions. Glu levels in the pgACC were correlated with frontal area brain activations, whereas Glu in the salience network component aMCC modulated responses in the PCC–precuneus. We show that region‐specific metabolite mapping enables specific activation of different BOLD signals in the brain underlying emotional perception.     

Default-mode activity during a passive sensory task: uncoupled from deactivation but impacting activation

Deactivation refers to increased neural activity during low-demand tasks or rest compared with high-demand tasks. Several groups have reported that a particular set of brain regions, including the posterior cingulate cortex and the medial prefrontal cortex, among others, is consistently deactivated. Taken together, these typically deactivated brain regions appear to constitute a default-mode network of brain activity that predominates in the absence of a demanding external task. Examining a passive, block-design sensory task with a standard deactivation analysis (rest epochs vs. stimulus epochs), we demonstrate that the default-mode network is undetectable in one run and only partially detectable in a second run. Using independent component analysis, however, we were able to detect the full default-mode network in both runs and to demonstrate that, in the majority of subjects, it persisted across both rest and stimulus epochs, uncoupled from the task waveform, and so mostly undetectable as deactivation. We also replicate an earlier finding that the default-mode network includes the hippocampus suggesting that episodic memory is incorporated in default-mode cognitive processing. Furthermore, we show that the more a subject's default-mode activity was correlated with the rest epochs (and "deactivated" during stimulus epochs), the greater that subject's activation to the visual and auditory stimuli. We conclude that activity in the default-mode network may persist through both experimental and rest epochs if the experiment is not sufficiently challenging. Time-series analysis of default-mode activity provides a measure of the degree to which a task engages a subject and whether it is sufficient to interrupt the processes--presumably cognitive, internally generated, and involving episodic memory--mediated by the default-mode network.     

Ketamine alters neural processing of facial emotion recognition in healthy men: an fMRI study

Disruption of facial emotion perception occurs in neuropsychiatric disorders where the expression of emotion is dulled or blunted, for example depersonalization disorder and schizophrenia. It has been suggested that, in the clinical context of emotional blunting, there is a shift in the relative contribution of brain regions subserving cognitive and emotional processing. The non-competitive glutamate receptor antagonist ketamine produces such emotional blunting in healthy subjects. Therefore, we hypothesised that in healthy subjects ketamine would elicit neural responses to emotional stimuli which mimicked those reported in depersonalization disorder and schizophrenia. Thus, we predicted that ketamine would produce reduced activity in limbic and visual brain regions involved in emotion processing, and increased activity in dorsal regions of the prefrontal cortex and cingulate gyrus, both associated with cognitive processing and, putatively, with emotion regulation. Measuring BOLD signal change in fMRI, we examined the neural correlates of ketamine-induced emotional blunting in eight young right-handed healthy men receiving an infusion of ketamine or saline placebo while viewing alternating 30 s blocks of faces displaying fear versus neutral expressions. The normal pattern of neural response occurred in limbic and visual cortex to fearful faces during the placebo infusion. Ketamine abolished this: significant BOLD signal change was demonstrated only in left visual cortex. However, with ketamine, neural responses were demonstrated to neutral expressions in visual cortex, cerebellum and left posterior cingulate gyrus. Emotional blunting may be associated with reduced limbic responses to emotional stimuli and a relative increase in the visual cortical response to neutral stimuli.     

FMRI activations of amygdala, cingulate cortex, and auditory cortex by infant laughing and crying

One of the functions of emotional vocalizations is the regulation of social relationships like those between adults and children. Listening to infant vocalizations is known to engage amygdala as well as anterior and posterior cingulate cortices. But, the functional relationships between these structures still need further clarification. Here, nonparental women and men listened to laughing and crying of preverbal infants and to vocalization-derived control stimuli, while performing a pure tone detection task during low-noise functional magnetic resonance imaging. Infant vocalizations elicited stronger activation in amygdala and anterior cingulate cortex (ACC) of women, whereas the alienated control stimuli elicited stronger activation in men. Independent of listeners' gender, auditory cortex (AC) and posterior cingulate cortex (PCC) were more strongly activated by the control stimuli than by infant laughing or crying. The gender-dependent correlates of neural activity in amygdala and ACC may reflect neural predispositions in women for responses to preverbal infant vocalizations, whereas the gender-independent similarity of activation patterns in PCC and AC may reflect more sensory-based and cognitive levels of neural processing. In comparison to our previous work on adult laughing and crying, the infant vocalizations elicited manifold higher amygdala activation.     

Neural correlates of orthographic and phonological consistency effects in children

The objective of this study was to examine the neural correlates of phonological inconsistency (relationship of spelling to sound) and orthographic inconsistency (relationship of sound to spelling) in visual word processing using functional magnetic resonance imaging (fMRI). Children (9- to 15-year-old) performed a rhyming and spelling task in which two words were presented sequentially in the visual modality. Consistent with previous studies in adults, higher phonological inconsistency was associated with greater activation in several regions including left inferior frontal gyrus and medial frontal gyrus/anterior cingulate cortex. We additionally demonstrated an effect of orthographic inconsistency in these same areas, suggesting that these regions are involved in the integration of orthographic and phonological information and, with respect to the medial frontal/anterior cingulate, greater demands on executive function. Higher phonological and orthographic consistency was associated with greater activation in precuneus/posterior cingulate cortex, the putative steady state system active during resting, suggesting lower demands on cognitive resources for consistent items. Both consistency effects were larger for the rhyming compared with the spelling task suggesting greater demands of integrating spelling and sound in the former task. Finally, accuracy on the rhyming task was negatively correlated with the consistency effect in left fusiform gyrus. In particular, this region showed insensitivity to consistency in low performers, sensitivity to inconsistency (higher activity) in moderate performers, and sensitivity to inconsistency (high activation) and to consistency (deactivation). In general, these results show that the influence of spelling-sound (and sound-spelling) correspondences on processing in fusiform gyrus develops as a function of skill.     

The default mode network integrity in patients with Parkinson’s disease is levodopa equivalent dose-dependent

Disturbances in the default mode network (DMN) have been described in many neurological and psychiatric disorders including Parkinson’s disease (PD). The DMN is characterized by basal activity that increases during rest or passive visual fixation and decreases (“deactivates”) during cognitive tasks. The network is believed to be involved in cognitive processes. We examined the DMN in PD patients on dopaminergic medication with normal cognitive performance compared to age- and gender-matched healthy controls (HC) using fMRI and three methodological procedures: independent component analysis of resting-state data, analysis of deactivation during a complex visual scene-encoding task, and seed-based functional connectivity analysis. In the PD group, we also studied the effect of dopaminergic medication on the DMN integrity. We did not find any difference between the PD and HC groups in the DMN, but using the daily levodopa equivalent dose as a covariate, we observed an enhanced functional connectivity of the DMN in the posterior cingulate cortex and decreased activation in the left parahippocampal gyrus during the cognitive task. We conclude that dopaminergic therapy has a specific effect on both the DMN integrity and task-related brain activations in cognitively unimpaired PD patients, and these effects seem to be dose-dependent.     

Fronto-striatal overactivation in euthymic bipolar patients during an emotional go/nogo task

OBJECTIVE: Although euthymic bipolar patients show minimal manic and depressive symptoms, they continue to show impaired emotional regulation and cognitive functioning. Few studies have directly examined the interference of emotional information with cognitive processes and its underlying cerebral mechanisms in euthymic bipolar patients. The authors examined the emotional modulation of cognitive processes and its underlying neural mechanisms in euthymic bipolar patients. METHOD: Seventeen euthymic bipolar patients and 17 healthy comparison subjects underwent functional magnetic resonance imaging (MRI) while performing an emotional and nonemotional go/nogo task. Neural responses associated with the overall task performance, as well as with the impact of emotional information on the task performance, were assessed. RESULTS: Bipolar disorder patients exhibited increased activity in the temporal cortex, specifically to emotional go/nogo conditions, as well as in the orbitofrontal cortex, the insula, the caudate nuclei, and the dorsal anterior and posterior cingulate cortices when inhibiting emotional stimuli compared with neutral stimuli. Conversely, no global attentional deficits were observed on either a behavioral or neural response level, indicated by similar task performances for all task conditions and similar brain activation patterns when comparing all the go/nogo conditions with the resting state. CONCLUSIONS: The present study provides evidence of an altered emotional modulation of cognitive processing in euthymic bipolar patients, indicated by an overactivation in ventral-limbic, temporal, and dorsal brain structures during emotional go/nogo conditions in patients relative to comparison subjects.     

Abnormal neural responses to emotional visual stimuli in adolescents with conduct disorder.

BACKGROUND: It is widely held that aggression and antisocial behavior arise as a consequence of a deficiency in responding to emotional cues in the social environment. We asked whether neural responses evoked by affect-laden pictures would be abnormal in adolescents with conduct disorder (CD). METHODS: Functional magnetic resonance imaging during passive viewing of pictures with neutral or strong negative affective valence was performed in 13 male adolescents with severe CD aged 9 to 15 years and in 14 healthy age-matched control subjects. RESULTS: Main effects for negative-neutral affective valence included activations in the amygdala and hippocampus, ventral extrastriate visual cortex, and intraparietal sulcus bilaterally. There was a significant group-by-condition interaction in the right dorsal anterior cingulate cortex that was due to a pronounced deactivation in the patient group during viewing of negative pictures. When correcting for anxiety and depressive symptoms, we additionally found a reduced responsiveness of the left amygdala to negative pictures in patients compared with control subjects. CONCLUSIONS: We suggest that these findings reflect an impairment of both the recognition of emotional stimuli and the cognitive control of emotional behavior in patients with CD, resulting in a propensity for aggressive behavior.     

Differential dorsal and ventral medial prefrontal representations of the implicit self modulated by individualism and collectivism: An fMRI study

Individualism and collectivism, or self-construal style, refer to cultural values that influence how people think about themselves and their relation to the social and physical environment. Recent neuroimaging evidence suggests that cultural values of individualism and collectivism dynamically modulate neural response within cortical midline structures, such as the medial prefrontal cortex (MPFC) and posterior cingulate cortex (PCC), during explicit self-evaluation. However, it remains unknown whether cultural priming modulates neural response during self-evaluation due to explicit task demands. Here we investigated how cultural priming of self-construal style affects neural activity within cortical midline structures during implicit self-evaluation in bicultural individuals. Results indicate that ventral MPFC showed relatively less deactivation during implicit evaluation of both self- and father-relevant information as compared to control condition (e.g., information of an unfamiliar person), irrespective of cultural priming. By contrast, dorsal MPFC showed relatively less deactivation during implicit evaluation of father-relevant information, but not self-relevant information, as compared to control condition, only when they were primed with individualism. Furthermore, dorsal MPFC showed relatively less deactivation during implicit evaluation of father-relevant information as compared to self-relevant condition only when they were primed with individualism. Hence, our results indicate that cultural priming modulates neural response within dorsal, but not ventral, portions of MPFC in a stimulus-driven rather than task-driven manner. More broadly, these findings suggest that cultural values dynamically shape neural representations during the evaluation, rather than the detection, of self-relevant information.     

Mood alters amygdala activation to sad distractors during an attentional task.

BACKGROUND: A behavioral hallmark of mood disorders is biased perception and memory for sad events. The amygdala is poised to mediate internal mood and external event processing because of its connections with both the internal milieu and the sensory world. There is little evidence showing that the amygdala's response to sad sensory stimuli is functionally modulated by mood state, however. METHODS: We investigated the impact of mood on amygdala activation evoked by sad and neutral pictures presented as distractors during an attentional oddball task. Healthy adults underwent functional magnetic resonance imaging during task runs that were preceded by sad or happy movie clips. Happy and sad mood induction was conducted within-subjects on consecutive days in counterbalanced order. RESULTS: Amygdala activation to sad distractors was enhanced after viewing sad movies relative to happy ones and was correlated with reaction time costs to detect attentional targets. The activation was higher in female subjects in the right hemisphere. The anterior cingulate, ventromedial and orbital prefrontal cortex, insula, and other posterior regions also showed enhanced responses to sad distractors during sad mood. CONCLUSIONS: These findings reveal brain mechanisms that integrate emotional input and current mood state, with implications for understanding cognitive distractibility in depression.     

Affective modulation of anterior cingulate cortex in young people at increased familial risk of depression

BACKGROUND: We previously found that children of parents with depression showed impaired performance on a task of emotional categorisation. AIMS: To test the hypothesis that children of parents with depression would show abnormal neural responses in the anterior cingulate cortex, a brain region involved in the integration of emotional and cognitive information. METHOD: Eighteen young people (mean age 19.8 years) with no personal history of depression but with a biological parent with a history of major depression (FH+ participants) and 16 controls (mean age 19.9 years) underwent functional magnetic resonance imaging while completing an emotional counting Stroop task. RESULTS: Controls showed significant activation in the pregenual anterior cingulate cortex to both positive and negative words during the emotional Stroop task. This activation was absent in FH+ participants. CONCLUSIONS: Our findings show that people at increased familial risk of depression demonstrate impaired modulation of the anterior cingulate cortex in response to emotionally valenced stimuli.     

An fMRI study of simple ethical decision-making

Recent functional neuroimaging studies suggest that ventromedial prefrontal cortex (vmPFC), left posterior superior temporal sulcus (pSTS) and posterior cingulate cortex are engaged during moral decision-making on complex dilemmatic or salient emotional stimuli. In this fMRI study we investigated which of these brain regions are activated during simple ethical decision-making about unambiguous scenarios not containing direct bodily harm or violence. Simple moral decisions compared to semantic decisions resulted in activation of left pSTS and middle temporal gyrus, bilateral temporal poles, left lateral PFC and bilateral vmPFC. These results suggest that pSTS and vmPFC are a common neuronal substrate of decision-making about complex ethical dilemmas, processing material evocative of moral emotions and simple ethical decision-making about scenarios devoid of violence and direct bodily harm.