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1.  Emotional learning and the development of differential moralities: Implications from research on psychopathy 
In this paper, we will argue that (1) four classes of norm can be distinguished from a neuro-cognitive perspective; (2) learning the prohibitive power of these norms relies on relatively independent emotional systems; (3) individuals with psychopathy show selective impairment for one of these emotional learning systems and two classes of norm: care-based and justice-based; and (4) while emotional learning systems are necessary for appropriate moral development/reasoning, they are not sufficient for moral development/reasoning.
PMCID: PMC4324693  PMID: 25684831
2.  Cognitive control of attention is differentially affected in trauma-exposed individuals with and without post-traumatic stress disorder 
Psychological medicine  2012;43(1):85-95.
This study aimed to determine whether patients with post-traumatic stress disorder (PTSD) show difficulty in recruitment of the regions of the frontal and parietal cortex implicated in top-down attentional control in the presence and absence of emotional distracters.
Unmedicated individuals with PTSD (n=14), and age-, IQ- and gender-matched individuals exposed to trauma (n=15) and healthy controls (n=19) were tested on the affective number Stroop task. In addition, blood oxygen level-dependent responses, as measured via functional magnetic resonance imaging, were recorded.
Patients with PTSD showed disrupted recruitment of lateral regions of the superior and inferior frontal cortex as well as the parietal cortex in the presence of negative distracters. Trauma-comparison individuals showed indications of a heightened ability to recruit fronto-parietal regions implicated in top-down attentional control across distracter conditions.
These results are consistent with suggestions that emotional responsiveness can interfere with the recruitment of regions implicated in top-down attentional control ; the heightened emotional responding of patients with PTSD may lead to the heightened interference in the recruitment of these regions.
PMCID: PMC4418499  PMID: 22571775
Anxiety; post-traumatic stress disorder; fMRI; fronto-parietal cortex; top down attention control
3.  Cortical and Subcortical Abnormalities in Youths with Conduct Disorder and Elevated Callous-Unemotional Traits 
Though there is growing evidence of brain abnormalities among individuals with Conduct Disorder (CD), the structural neuroimaging literature is mixed and frequently aggregates cortical volume rather than differentiating cortical thickness from surface area. The current study assesses CD-related differences in cortical thickness, surface area, and gyrification as well as volume differences in subcortical structures critical to neurodevelopmental models of CD (amygdala; striatum) in a carefully characterized sample. We also examined whether group structural differences were related to severity of callous-unemotional (CU) traits in the CD sample.
Participants were 49 community adolescents (aged 10-18 years); 22 with CD and 27 healthy comparison youth. Structural MRI was collected and the FreeSurfer image analysis suite was used to provide measures of cortical thickness, surface area and local gyrification as well as subcortical (amygdala and striatum) volumes.
Youths with CD showed reduced cortical thickness in superior temporal cortex. There were also indications of reduced gyrification in ventromedial frontal cortex particularly for youth with CD without comorbid Attention Deficit Hyperactivity Disorder. There were no group differences in cortical surface area. However, the youth with CD also showed reduced amygdala and striatum (putamen and pallidum) volumes. Right temporal cortical thickness was significantly inversely related to severity of CU traits.
Youths with CD show reduced cortical thickness within superior temporal regions, some indication of reduced gyrification within ventromedial frontal cortex and reduced amygdala and striatum (putamen and pallidum) volumes. These results are discussed with reference to neurobiological models of CD.
PMCID: PMC4281036  PMID: 24655655
conduct disorder; antisocial; amygdala; striatum; cortical thickness
4.  Looming animate and inanimate threats: The response of the amygdala and periaqueductal gray 
Social neuroscience  2013;8(6):621-630.
Looming stimuli are processed as threatening and activate basic neural defense systems. However, it is unclear how animacy information modulates this response. Participants (N = 25) viewed threatening or neutral images that were either animate (animals) or inanimate (objects) and which either approached (loomed) or receded from the participant. The amygdala was responsive to emotional, animacy and looming information (particularly to looming threats and looming animate stimuli). Periaqueductal gray was also sensitive to emotional information and particularly responsive to looming threats. The data are interpreted within category specific models of the amygdala and temporal cortex.
PMCID: PMC3919685  PMID: 24066700
Amygdala; Animacy; Threat; Looming; Periaqueductal Gray
5.  Reactive aggression and functional, not neural, specificity 
PMCID: PMC3941713  PMID: 20594400
6.  Dissociable Roles of Ventromedial Prefrontal Cortex (vmPFC) and Rostral Anterior Cingulate Cortex (rACC) in Value Representation and Optimistic Bias 
NeuroImage  2013;78:103-110.
Optimistic bias (OB) is seen when individuals underestimate their probability of experiencing negative life events and overestimate their probability of experiencing positive life events. A reduced OB has been linked with increased depression symptoms . However, given the relevance of this information to mood and anxiety disorders, little is currently known regarding the neurobiology of OB. In the current study, we examine the neural basis of OB in healthy individuals (n=33) during probability estimation of future positive and negative events occurring to themselves relative to other, comparable individuals. In line with previous work, subjects showed significant OB; they considered themselves significantly more likely to experience future positive and significantly less likely to experience future negative events relative to comparable others. Positive, relative to negative events, un-modulated by subjects’ probability estimates, were associated with significantly greater activity within ventromedial prefrontal cortex (vmPFC) and posterior cingulate cortex (PCC). Moreover, responses within both regions to positive events negatively related to the healthy subjects’ self reports of depression symptoms. However, there was no significant modulation of activity in either region by the subject’s OB, objectified as the level to which they thought the event was more likely [positive events] or less likely [negative events] to occur to them relative to comparable others. In contrast, activity within rostral anterior cingulate cortex (rACC) was positively modulated by OB for positive events and activity within anterior insula and dorsomedial prefrontal cortex (dmPFC) was negatively modulated by OB for negative events. However, there was no significant relationship between responsiveness within these regions and self reports of depression symptoms. The data are discussed with reference to current models of vmPFC, rACC and anterior insula functioning.
PMCID: PMC3686504  PMID: 23567883
fMRI; ventromedial prefrontal prefrontal cortex; rostral anterior cingulate cortex; optimistic bias
7.  Empathic responsiveness in amygdala and anterior cingulate cortex in youths with psychopathic traits 
Psychopathic traits are associated with increases in antisocial behaviors such as aggression and are characterized by reduced empathy for others’ distress. This suggests that psychopathic traits may also impair empathic pain sensitivity. However, whether psychopathic traits affect responses to the pain of others versus the self has not been previously assessed.
We used whole-brain functional magnetic resonance imaging (fMRI) scanning to measure neural activation in 14 adolescents with Oppositional Defiant Disorder or Conduct Disorder and psychopathic traits, as well as 21 healthy controls matched on age, sex, and intelligence. Activation in structures associated with empathic pain perception was assessed as adolescents viewed photographs of pain-inducing injuries. Adolescents imagined either that the body in each photograph was their own or that it belonged to another person. Behavioral and neuroimaging data were analyzed using random-effects analysis of variance.
Youths with psychopathic traits showed reduced activity within regions associated with empathic pain as the depicted pain increased. These regions included rostral anterior cingulate cortex, ventral striatum (putamen), and amygdala. Reductions in amygdala activity particularly occurred when the injury was perceived as occurring to another. Empathic pain responses within both amygdala and rostral anterior cingulate cortex were negatively correlated with the severity of psychopathic traits as indexed by PCL:YV scores.
Youths with psychopathic traits show less responsiveness in regions implicated in the affective response to another’s pain as the perceived intensity of this pain increases. Moreover, this reduced responsiveness appears to predict symptom severity.
PMCID: PMC3716835  PMID: 23488588
Psychopathy; adolescents; empathy; pain; amygdala; Conduct Disorder
8.  Dissociable Roles of Medial Orbitofrontal Cortex in Human Operant Extinction Learning 
NeuroImage  2008;43(4):748-755.
Operant extinction, which features modification of instrumental responses to stimuli following a change in associated reinforcement, is an important form of learning for organisms in dynamic environments. Animal studies have highlighted orbital and medial prefrontal cortex and amygdala as mediators of operant extinction. Yet little is known about the neural mediators of operant extinction learning in humans. Using a novel fMRI paradigm, we report dissociable functional responses in distinct regions of medial orbitofrontal cortex (mOFC) during successful appetitive and aversive based operant extinction. During successful operant extinction, increased activity was observed in frontopolar OFC, while decreased activity was observed in caudal mOFC and rostral anterior cingulate cortex (rACC) relative to both (i) successful control trials where the reinforcement associated with the stimulus does not change; and (ii) successful acquisition trials during initial learning of the stimulus- reinforcement associations. Functional connectivity analysis demonstrated inverse connectivity between frontopolar OFC and both rACC and the amygdala. These data support animal models suggesting the importance of mOFC - amygdala interaction during operant extinction and expand our knowledge of the neural systems in humans. These findings suggest that in humans, frontopolar OFC modulates activity in caudal mOFC, rACC and amygdala during successful operant extinction learning.
PMCID: PMC3941699  PMID: 18793731
frontopolar; amygdala; instrumental; anterior cingulate cortex
9.  Neuroimaging of Psychopathy and Antisocial Behavior: A Targeted Review 
Current psychiatry reports  2010;12(1):76-82.
The goal of this article is to provide a selective and targeted review of the neuroimaging literature on psychopathic tendencies and antisocial behavior and to explore the extent to which this literature supports recent cognitive neuroscientific models of psychopathy and antisocial behavior. The literature reveals that individuals who present with an increased risk for reactive, but not instrumental, aggression show increased amygdala responses to emotionally evocative stimuli. This is consistent with suggestions that such individuals are primed to respond strongly to an inappropriate extent to threatening or frustrating events. In contrast, individuals with psychopathic tendencies show decreased amygdala and orbitofrontal cortex responses to emotionally provocative stimuli or during emotional learning paradigms. This is consistent with suggestions that such individuals face difficulties with basic forms of emotional learning and decision making.
PMCID: PMC3941715  PMID: 20425314
Amygdala; Orbital frontal cortex; Psychopathy; Instrumental aggression; Reactive aggression
10.  Elevated amygdala responses to emotional faces in youths with chronic irritability or bipolar disorder 
NeuroImage : clinical  2013;2:637-645.
A major controversy in child psychiatry is whether bipolar disorder (BD) presents in children as severe, non-episodic irritability (operationalized here as severe mood dysregulation, SMD), rather than with manic episodes as in adults. Both classic, episodic BD and SMD are severe mood disorders characterized by deficits in processing emotional stimuli. Neuroimaging techniques can be used to test whether the pathophysiology mediating these deficits are similar across the two phenotypes. Amygdala dysfunction during face emotion processing is well-documented in BD, but little is known about amygdala dysfunction in chronically irritable youth. We compared neural activation in SMD (n=19), BD (n=19), and healthy volunteer (HV; n=15) youths during an implicit face-emotion processing task with angry, fearful and neutral expressions. In the right amygdala, both SMD and BD exhibited greater activity across all expressions than HV. However, SMD and BD differed from each other and HV in posterior cingulate cortex, posterior insula, and inferior parietal lobe. In these regions, only SMD showed deactivation in response to fearful expressions, whereas only BD showed deactivation in response to angry expressions. Thus, during implicit face emotion processing, youth with BD and those with SMD exhibit similar amygdala dysfunction but different abnormalities in regions involved in information monitoring and integration.
PMCID: PMC3746996  PMID: 23977455
severe mood dysregulation; bipolar disorder; amygdala; face; emotion
11.  The Influence of Oxytocin Administration on Responses to Infant Faces and Potential Moderation by OXTR Genotype 
Psychopharmacology  2012;224(4):469-476.
Oxytocin is a neuropeptide that is associated with increases in social affiliative behaviors, particularly toward infants. However, no previous study has investigated healthy adults’ responses to infant faces following oxytocin administration. In addition, given that preliminary evidence suggests that a single nucleotide polymorphism (SNP) of the oxytocin receptor (OXTR) gene, rs53576, may influence behaviors associated with parental sensitivity, we assessed whether such responses vary according to OXTR rs53576 genotype.
The present study assessed the effects of intranasally administered oxytocin and OXTR genotype on human adults’ preferences for infant faces.
A double-blind, between-groups design was used, with 57 genotyped volunteers randomly assigned to receive intranasally administered oxytocin or placebo. Fifty minutes following the administration of oxytocin or placebo, participants viewed infants’ and adults’ faces showing neutral expressions and assessed how appealing they found each face.
Infants’ faces were more strongly preferred following oxytocin inhalation relative to placebo. When participants were separated according to genotype, this effect was only observed for participants homozygous for the rs53576G allele. Parallel effects were not seen for adults’ faces.
The present results are consistent with the hypothesis that acute oxytocin administration increases sensitivity to reward-relevant features of infants and/or reduces sensitivity to their aversive properties. The results also are consistent with suggestions of more efficient oxytocinergic function in rs53576G homozygotes.
PMCID: PMC3500580  PMID: 22763666
Oxytocin; OXTR; parental; faces; affiliation
12.  Reduced dorsal anterior cingulate cortical activity during emotional regulation and top-down attentional control in Generalized Social Phobia (GSP), Generalized Anxiety Disorder (GAD) and comorbid GSP/GAD 
Biological psychiatry  2012;72(6):476-482.
Generalized Social Phobia (GSP) and Generalized Anxiety Disorder (GAD) are both associated with emotion dysregulation. In healthy subjects, research implicates dorsal anterior cingulate (dACC) in both explicit emotion regulation and top-down attentional control. While studies have examined these processes in GSP or GAD, no work compares findings across the two disorders. Moreover, no work examines functioning in cases comorbid for both disorders (GSP/GAD). Here we compare the neural correlates of explicit emotion regulation (EER) and top-down attentional control (TAC) in GSP, GAD, and GSP/GAD.
Medication-free adults with GSP (EER n=19; TAC n=18), GAD (EER n=17; TAC n =17), GSP/GAD (EER n=17; TAC=15), or no psychopathology (EER n=18; TAC n=18). During EER, individuals alternatively viewed, up-regulated, and down-regulated responses to emotional pictures. During TAC, they performed an emotional Stroop task.
For both tasks, significant group-by-condition interactions emerged in dACC and parietal cortices. Healthy adults showed significantly increased recruitment during emotion regulation, relative to emotion-picture viewing. GAD, GSP, and GSP/GAD subjects showed no such increases, with all three groups differing from healthy adults but not from each other. Evidence of emotion-related disorder-specificity emerged in medial prefrontal cortex (MPFC) and amygdala. This disorder-specific responding varied as a function of stimulus emotion content but not emotion-regulatory demands.
GSP and GAD both involve reduced capacity for engaging emotion-regulation brain networks, whether explicitly or via top-down attentional control. A reduced ability to recruit regions implicated in top-down attention might represent a general risk factor for anxiety disorders.
PMCID: PMC3424322  PMID: 22592057
imaging; social anxiety; generalized anxiety; emotion regulation; anterior cingulate cortex; top-down attentional control
13.  Differing Amygdala Responses to Facial Expressions in Children vs. Adults with Bipolar Disorder 
The American Journal of Psychiatry  2012;169(6):642-649.
Bipolar disorder (BD) is highly debilitating in both children and adults. Child and adult BD patients show both behavioral deficits in face emotion processing and abnormal amygdala activation. However, amygdala function in pediatric vs. adult BD patients has never been compared directly.
This study compared amygdala responses to emotional facial expressions in 74 subjects [pediatric (N=18) and adult (N=17) patients with BD, healthy volunteer (HV) children (N=15) and adults (N=22)]. Subjects performed a gender identification task while viewing fearful, angry, and neutral faces.
In response to fearful faces, patients with BD across age-groups showed right amygdala hyperactivity relative to healthy volunteers. However, when responses to all facial expressions were combined, pediatric patients exhibited greater right amygdala activation than either adults with BD or HV children.
Amygdala hyperactivity in response to fearful faces is present in both youths and adults with BD. However, compared to adults with BD or healthy youths, youths with BD show amygdala hyperactivity in response to a broad array of emotional faces. Thus, abnormal amygdala activation during face processing appears to be more pervasive in children than adults with BD.
PMCID: PMC3383787  PMID: 22535257
14.  Elevated amygdala responses to emotional faces in youths with chronic irritability or bipolar disorder☆ 
NeuroImage : Clinical  2013;2:637-645.
A major controversy in child psychiatry is whether bipolar disorder (BD) presents in children as severe, non-episodic irritability (operationalized here as severe mood dysregulation, SMD), rather than with manic episodes as in adults. Both classic, episodic BD and SMD are severe mood disorders characterized by deficits in processing emotional stimuli. Neuroimaging techniques can be used to test whether the pathophysiology mediating these deficits are similar across the two phenotypes. Amygdala dysfunction during face emotion processing is well-documented in BD, but little is known about amygdala dysfunction in chronically irritable youth. We compared neural activation in SMD (n = 19), BD (n = 19), and healthy volunteer (HV; n = 15) youths during an implicit face-emotion processing task with angry, fearful and neutral expressions. In the right amygdala, both SMD and BD exhibited greater activity across all expressions than HV. However, SMD and BD differed from each other and HV in posterior cingulate cortex, posterior insula, and inferior parietal lobe. In these regions, only SMD showed deactivation in response to fearful expressions, whereas only BD showed deactivation in response to angry expressions. Thus, during implicit face emotion processing, youth with BD and those with SMD exhibit similar amygdala dysfunction but different abnormalities in regions involved in information monitoring and integration.
•Youths with severe mood dysregulation (SMD), bipolar disorder (BD), controls.•Implicit face-emotion processing fMRI task with angry, fearful, neutral emotions.•In R amygdala, SMD and BD had greater activity across all expressions vs controls.•In whole brain analysis SMD had decreased, BD increased activity vs. other groups.
PMCID: PMC3746996  PMID: 23977455
Severe mood dysregulation; Bipolar disorder; Amygdala; Face; Emotion
15.  Considering anger from a cognitive neuroscience perspective 
The goal of this paper is to consider anger from a cognitive neuroscience perspective. Five main claims are made: First, reactive aggression is the ultimate behavioral expression of anger and thus we can begin to understand anger by understanding reactive aggression. Second, neural systems implicated in reactive aggression (amygdala, hypothalamus and periaqueductal gray; the basic threat system) are critically implicated in anger. Factors such as exposure to extreme threat that increase the responsiveness of these systems, should be (and are in the context of Post Traumatic Stress Disorder), associated with increased anger. Third, regions of frontal cortex implicated in regulating the basic threat system, when dysfunctional (e.g., in the context of lesions) should be associated with increased anger. Fourth, frustration occurs when an individual continues to do an action in the expectation of a reward but does not actually receive that reward, and is associated with anger. Individuals who show impairment in the ability to alter behavioral responding when actions no longer receive their expected rewards should be (and are in the context of psychopathy) associated with increased anger. Fifth, someone not doing what another person wants them to do (particularly if this thwarts the person’s goal) is frustrating and consequently anger inducing. The response to such a frustrating social event relies on the neural architecture implicated in changing behavioral responses in non-social frustrating situations.
PMCID: PMC3260787  PMID: 22267973
16.  Reduced amygdala-orbitofrontal connectivity during moral judgments in youths with disruptive behavior disorders and psychopathic traits 
Psychiatry research  2011;194(3):279-286.
We used functional magnetic resonance imaging (fMRI) to investigate dysfunction in the amygdala and orbitofrontal cortex in adolescents with disruptive behavior disorders and psychopathic traits during a moral judgment task. Fourteen adolescents with psychopathic traits and 14 healthy controls were assessed using fMRI while they categorized illegal and legal behaviors in a moral judgment implicit association task. fMRI data were then analyzed using random-effects analysis of variance and functional connectivity. Youths with psychopathic traits showed reduced amygdala activity when making judgments about legal actions and reduced functional connectivity between the amygdala and orbitofrontal cortex during task performance. These results suggest that psychopathic traits are associated with amygdala and orbitofrontal cortex dysfunction. This dysfunction may relate to previous findings of disrupted moral judgment in this population.
PMCID: PMC3225495  PMID: 22047730
Conduct disorder; psychopathy; moral reasoning; fMRI
17.  Adolescents with psychopathic traits report reductions in physiological responses to fear 
Psychopathy is characterized by profound affective deficits, including shallow affect and reduced empathy. Recent research suggests that these deficits may apply particularly to negative emotions, or to certain negative emotions such as fear. Despite increased focus on the cognitive and neural underpinnings of psychopathy, little is known about how psychopathy is associated with emotional deficits across a range of emotions. In addition, the relationship between psychopathy and the subjective experience of emotion has not yet been assessed.
Eighteen 10-17-year-olds with psychopathic traits and 24 comparison children and adolescents reported on their subjective experiences of emotion during five recent emotionally evocative life events, following a paradigm developed by Scherer and colleagues (Scherer & Wallbott, 1994). Group comparisons were then performed to assess variations in subjective experiences across emotions.
As predicted, psychopathy was associated with reductions in the subjective experience of fear relative to other emotions. Children and adolescents with psychopathic traits reported fewer symptoms associated with sympathetic nervous system arousal during fear-evoking experiences.
Rather than being related to uniformly impoverished emotional experience, psychopathic traits appear to be associated with greater deficits in subjective experiences of fear. This pattern of responding supports and extends previous observations that psychopathy engenders deficits in fear learning, physiological responses to threats, and the recognition of fear in others.
PMCID: PMC3116087  PMID: 21155775
Psychopathy; emotion; fear; antisocial behavior; autonomic
18.  Atypical modulation of medial prefrontal cortex to self-referential comments in generalized social phobia 
Psychiatry research  2011;193(1):38-45.
Generalized social phobia (GSP) involves the fear of being negatively evaluated. Previous work suggests that self-referentiality, mediated by medial prefrontal cortex (MFPC), plays an important role in the disorder. However, it is not clear whether this anomalous MPFC response to self-related information in patients with GSP concerns an increased representation of their own or others’ opinions. In this paper we examined whether GSP is associated with increased response to own (1st person) or other individuals’ (2nd person) opinions relative to healthy individuals. Unmedicated individuals with GSP (n=15) and age, IQ, and gender-matched comparison individuals (n=15) read 1st (e.g., I’m ugly), and 2nd (e.g., You’re ugly) person viewpoint comments during fMRI. We observed significant group-by-viewpoint interactions within ventral MPFC. Whereas the healthy comparison individuals showed significantly increased (or less decreased) BOLD responses to 1st relative to 2nd person viewpoints, the patients showed significantly increased responses to 2nd relative to 1st person viewpoints. The reduced BOLD responses to 1st person viewpoint comments shown by the patients correlated significantly with severity of social anxiety symptom severity. These results underscore the importance of dysfunctional self-referential processing and MPFC in GSP. We believe that these data reflect a reorganization of self-referential reasoning in the disorder with a self-concept perhaps atypically related to the view of others.
PMCID: PMC3105197  PMID: 21601433
Social Phobia; fMRI; self-referentiality; medial prefrontal cortex; amygdala
19.  Neural Response to Self- and Other Referential Praise and Criticism in Generalized Social Phobia 
Archives of general psychiatry  2008;65(10):1176-1184.
Generalized social phobia (GSP) is characterized by fear/avoidance of social situations. Previous studies have examined the neural responses in GSP to one class of social stimuli, facial expressions. However, studies have not examined the neural response in GSP to another equally important class of social stimuli, the communication of praise or criticism.
To examine the neural response to receipt of praise or criticism in GSP; specifically, to determine whether patients with GSP show an increased response to the receipt of both praise and criticism and whether self-relevance modulates this relationship.
Case-control study.
Government clinical research institute.
Unmedicated individuals with GSP (n=17) and age-, IQ-, and sex-matched healthy comparison individuals (n=17).
Main Outcome Measure
Blood oxygenation level–dependent signal, as measured via functional magnetic resonance imaging. During functional magnetic resonance imaging scans, individuals read positive (eg, You are beautiful), negative (eg, You are ugly), and neutral (eg, You are human) comments that could be either about the self or about somebody else (eg, He is beautiful).
Hypothesized significant group×valence×referent interactions were observed within regions of the medial prefrontal cortex and bilateral amygdala. In these regions, the patients with GSP showed significantly increased blood oxygenation level–dependent responses, relative to comparison individuals, to negative comments (criticism) referring to themselves. However, in contrast, there were no significant group differences with respect to negative comments referring to others or neutral or positive comments referring to self or others.
These results implicate the medial prefrontal cortex, involved in the representation of the self, together with the amygdala, in the pathophysiology of GSP. Further, findings demonstrate a meaningful effect of psychological context on neural-circuitry hyperactivity in GSP.
PMCID: PMC2785901  PMID: 18838634
20.  The neural substrates of action identification 
Mentalization is the process by which an observer views a target as possessing higher cognitive faculties such as goals, intentions and desires. Mentalization can be assessed using action identification paradigms, in which observers choose mentalistic (goals-focused) or mechanistic (action-focused) descriptions of targets’ actions. Neural structures that play key roles in inferring goals and intentions from others’ observed or imagined actions include temporo-parietal junction, ventral premotor cortex and extrastriate body area. We hypothesized that these regions play a role in action identification as well. Data collected using functional magnetic resonance imaging (fMRI) confirmed our predictions that activity in ventral premotor cortex and middle temporal gyrus near the extrastriate body area varies both as a function of the valence of the target and the extent to which actions are identified as goal-directed. In addition, the inferior parietal lobule is preferentially engaged when participants identify the actions of mentalized targets. Functional connectivity analyses suggest support from other regions, including the medial prefrontal cortex and amygdala, during mentalization. We found correlations between action identification and Autism Quotient scores, suggesting that understanding the neural correlates of action identification may enhance our understanding of the underpinnings of essential social cognitive processes.
PMCID: PMC2999755  PMID: 20150343
fMRI; action identification; mentalize; ventral premotor cortex; extrastriate body area; amygdale
21.  Serotonin Transporter Genotype (5-HTTLPR) Predicts Utilitarian Moral Judgments 
PLoS ONE  2011;6(10):e25148.
The psychological and neurobiological processes underlying moral judgment have been the focus of extensive recent research. Here we show that serotonin transporter (5-HTTLPR) genotype predicts responses to moral dilemmas featuring foreseen harm to an innocent.
Methodology/Principal Findings
Participants in this study judged the acceptability of actions that would unintentionally or intentionally harm an innocent victim in order to save others' lives. An analysis of variance revealed a genotype × scenario interaction, F(2, 63) = 4.52, p = .02. Results showed that, relative to long allele homozygotes (LL), carriers of the short (S) allele showed particular reluctance to endorse utilitarian actions resulting in foreseen harm to an innocent individual. LL genotype participants rated perpetrating unintentional harm as more acceptable (M = 4.98, SEM = 0.20) than did SL genotype participants (M = 4.65, SEM = 0.20) or SS genotype participants (M = 4.29, SEM = 0.30). No group differences in moral judgments were observed in response to scenarios featuring intentional harm.
The results indicate that inherited variants in a genetic polymorphism that influences serotonin neurotransmission influence utilitarian moral judgments as well. This finding is interpreted in light of evidence that the S allele is associated with elevated emotional responsiveness.
PMCID: PMC3187753  PMID: 21998637
22.  Social norm processing in adult Social Phobia: Atypically increased ventromedial frontal cortex responsiveness to unintentional (embarrassing) transgressions 
The American journal of psychiatry  2010;167(12):1526-1532.
Generalized social phobia (GSP) is defined by a persistent fear of social disapproval. However, the neural underpinnings of this increased fear and its mediating factors are unclear. Using event-related fMRI, we examined whether the intent of an event, which mediates the neural response to social disapproval in healthy individuals, differentially affects response in GSP. Specifically, would patients with GSP show particularly increased response to embarrassing, unintentional transgressions?
Sixteen patients with GSP and sixteen age, IQ, and gender matched healthy individuals read stories during fMRI scans that either involved neutral social events, unintentional social transgressions (e.g., choking on food at party, and coughing it up), or intentional social transgressions (e.g., disliking food at party, and spitting it out).
Significant group-by-transgression interactions were observed within ventral regions of medial prefrontal cortex (MPFC). Healthy individuals tended to show increased BOLD responses to intentional, relative to unintentional transgressions. The patients with GSP, however, showed significantly increased responses to the unintentional transgressions. In addition, they rated the unintentional transgressions as significantly more embarrassing than the comparison individuals. We also observed significant group main effects within the amygdala and bilateral insula, reflecting elevated GSP responses within these regions to all event types.
These results further implicate the MPFC in the pathophysiology of GSP, specifically through its involvement in distorted self-referential processing. In addition, the current results further underscore the extended role of the amygdala and insula in the processing of social stimuli more generally in GSP.
PMCID: PMC3175630  PMID: 20889651
23.  Stimulus-reinforcement Based Decision-making and Anxiety: Impairment in Generalized Anxiety Disorder (GAD), but not in Generalized Social Phobia (GSP) 
Psychological medicine  2008;39(7):1153-1161.
Generalized Social Phobia (GSP) involves the fear/avoidance of social situations while Generalized Anxiety Disorder (GAD) involves an intrusive worry about everyday life circumstances. It remains unclear whether these, highly comorbid, conditions represent distinct disorders or alternative presentations of a single underlying pathology. In this study, we examined stimulus-reinforcement based decision-making in GSP and GAD.
Twenty unmedicated patients with GSP, sixteen unmedicated patients with GAD and nineteen age, IQ, and gender matched healthy comparison individuals completed the Differential Reward/ Punishment Learning Task (DRPLT). In this task, the subject chooses between two objects associated with different levels of reward or punishment. Thus, response choice indexes not only reward/ punishment sensitivity but also sensitivity to reward/ punishment level according to between-object reinforcement distance.
We found that patients with GAD committed a significantly greater number of errors compared to both the patients with GSP and the healthy comparison individuals. In contrast, the patients with GSP and the healthy comparison individuals did not differ in performance on this task.
These results link GAD with an anomalous non-affective based decision-making. Further, they are indicative that GSP and GAD are associated with distinct pathophysiologies.
PMCID: PMC2999404  PMID: 19102795
24.  Response to Emotional Expressions in Generalized Social Phobia and Generalized Anxiety Disorder: Evidence for Separate Disorders 
The American journal of psychiatry  2008;165(9):1193-1202.
Generalized social phobia involves fear/avoidance, specifically of social situations, whereas generalized anxiety disorder involves intrusive worry about diverse circumstances. It remains unclear the degree to which these two, often comorbid, conditions represent distinct disorders or alternative presentations of a single, core underlying pathology. Functional magnetic resonance imaging assessed the neural response to facial expressions in generalized social phobia and generalized anxiety disorder.
Individuals matched on age, IQ, and gender with generalized social phobia without generalized anxiety disorder (N=17), generalized anxiety disorder (N= 17), or no psychopathology (N=17) viewed neutral, fearful, and angry expressions while ostensibly making a simple gender judgment.
The patients with generalized social phobia without generalized anxiety disorder showed increased activation to fearful relative to neutral expressions in several regions, including the amygdala, compared to healthy individuals. This increased amygdala response related to self-reported anxiety in patients with generalized social phobia without generalized anxiety disorder. In contrast, patients with generalized anxiety disorder showed significantly less activation to fearful relative to neutral faces compared to the healthy individuals. They did show significantly increased response to angry expressions relative to healthy individuals in a lateral region of the middle frontal gyrus. This increased lateral frontal response related to self-reported anxiety in patients with generalized anxiety disorder.
These results suggest that neural circuitry dysfunctions differ in generalized social phobia and generalized anxiety disorder.
PMCID: PMC2855133  PMID: 18483136
25.  Psychopathy, attention and emotion 
Psychological medicine  2008;39(4):543-555.
Psychopathy is a developmental disorder marked by emotional hypo-responsiveness and an increased risk for antisocial behavior. Influential attention based accounts of psychopathy have long been made; however, these accounts have made relatively little reference to general models of attention in healthy individuals. The current paper has three aims: (1) To briefly describe current cognitive neuroscience data on differing attentional systems; (2) To examine the functional integrity of these attentional systems in individuals with psychopathy; (3) To consider the implications of these data for attention and emotion dysfunction accounts of psychopathy.
PMCID: PMC2769016  PMID: 18700991

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