Functional Changes in Temporal Lobe Activity During Transient Global Amnesia

Neurology. Feb, 2002  |  Pubmed ID: 11865146

The integrity of temporal lobe activity during and after recovery from transient global amnesia (TGA) was assessed in a case study using functional MRI. TGA was associated with scene-encoding deficits in a temporolimbic circuit that recovered over time. Frontoparietal areas recruited during the amnesic state may signify a compensatory reliance on visuospatial or working memory strategies. Reduction of extrastriate cortex responses over repeated testing sessions possibly indicates intact visual priming in TGA.

Dissociable Prefrontal Brain Systems for Attention and Emotion

Proceedings of the National Academy of Sciences of the United States of America. Aug, 2002  |  Pubmed ID: 12177452

The prefrontal cortex has been implicated in a variety of attentional, executive, and mnemonic mental operations, yet its functional organization is still highly debated. The present study used functional MRI to determine whether attentional and emotional functions are segregated into dissociable prefrontal networks in the human brain. Subjects discriminated infrequent and irregularly presented attentional targets (circles) from frequent standards (squares) while novel distracting scenes, parametrically varied for emotional arousal, were intermittently presented. Targets differentially activated middle frontal gyrus, posterior parietal cortex, and posterior cingulate gyrus. Novel distracters activated inferior frontal gyrus, amygdala, and fusiform gyrus, with significantly stronger activation evoked by the emotional scenes. The anterior cingulate gyrus was the only brain region with equivalent responses to attentional and emotional stimuli. These results show that attentional and emotional functions are segregated into parallel dorsal and ventral streams that extend into prefrontal cortex and are integrated in the anterior cingulate. These findings may have implications for understanding the neural dynamics underlying emotional distractibility on attentional tasks in affective disorders.

Neural Correlates of Person Recognition

Learning & Memory (Cold Spring Harbor, N.Y.). Jul-Aug, 2003  |  Pubmed ID: 12888543

Rapidly identifying known individuals is an essential skill in human society. To elucidate the neural basis of this skill, we monitored brain activity while experimental participants demonstrated their ability to recognize people on the basis of viewing their faces. Each participant first memorized the faces of 20 individuals who were not known to the participants in advance. Each face was presented along with a voice simulating the individual speaking their name and a biographical fact. Following this learning procedure, the associated verbal information could be recalled accurately in response to each face. These learned faces were subsequently viewed together with new faces in a memory task. Subjects made a yes-no recognition decision in response to each face while also covertly retrieving the person-specific information associated with each learned face. Brain activity that accompanied this retrieval of person-specific information was contrasted to that when new faces were processed. Functional magnetic resonance imaging in 10 participants showed that several brain regions were activated during blocks of learned faces, including left hippocampus, left middle temporal gyrus, left insula, and bilateral cerebellum. Recordings of event-related brain potentials in 10 other participants tracked the time course of face processing and showed that learned faces engaged neural activity responsible for person recognition 300-600 msec after face onset. Collectively, these results suggest that the visual input of a recently learned face can rapidly trigger retrieval of associated person-specific information through reactivation of distributed cortical networks linked via hippocampal connections.

Emotional Memory Functions of the Human Amygdala

Current Neurology and Neuroscience Reports. Sep, 2003  |  Pubmed ID: 12914677

Dynamic Perception of Facial Affect and Identity in the Human Brain

Cerebral Cortex (New York, N.Y. : 1991). Oct, 2003  |  Pubmed ID: 12967919

Functional magnetic resonance imaging (fMRI) was used to compare brain activation to static facial displays versus dynamic changes in facial identity or emotional expression. Static images depicted prototypical fearful, angry and neutral expressions. Identity morphs depicted identity changes from one person to another, always with neutral expressions. Emotion morphs depicted expression changes from neutral to fear or anger, creating the illusion that the actor was 'getting scared' or 'getting angry' in real-time. Brain regions implicated in processing facial affect, including the amygdala and fusiform gyrus, showed greater responses to dynamic versus static emotional expressions, especially for fear. Identity morphs activated a dorsal fronto-cingulo-parietal circuit and additional ventral areas, including the amygdala, that also responded to the emotion morphs. Activity in the superior temporal sulcus discriminated emotion morphs from identity morphs, extending its known role in processing biologically relevant motion. The results highlight the importance of temporal cues in the neural coding of facial displays.

Emotion-attention Network Interactions During a Visual Oddball Task

Brain Research. Cognitive Brain Research. Jun, 2004  |  Pubmed ID: 15130591

Emotional and attentional functions are known to be distributed along ventral and dorsal networks in the brain, respectively. However, the interactions between these systems remain to be specified. The present study used event-related functional magnetic resonance imaging (fMRI) to investigate how attentional focus can modulate the neural activity elicited by scenes that vary in emotional content. In a visual oddball task, aversive and neutral scenes were presented intermittently among circles and squares. The squares were frequent standard events, whereas the other novel stimulus categories occurred rarely. One experimental group [N=10] was instructed to count the circles, whereas another group [N=12] counted the emotional scenes. A main effect of emotion was found in the amygdala (AMG) and ventral frontotemporal cortices. In these regions, activation was significantly greater for emotional than neutral stimuli but was invariant to attentional focus. A main effect of attentional focus was found in dorsal frontoparietal cortices, whose activity signaled task-relevant target events irrespective of emotional content. The only brain region that was sensitive to both emotion and attentional focus was the anterior cingulate gyrus (ACG). When circles were task-relevant, the ACG responded equally to circle targets and distracting emotional scenes. The ACG response to emotional scenes increased when they were task-relevant, and the response to circles concomitantly decreased. These findings support and extend prominent network theories of emotion-attention interactions that highlight the integrative role played by the anterior cingulate.

Interaction Between the Amygdala and the Medial Temporal Lobe Memory System Predicts Better Memory for Emotional Events

Neuron. Jun, 2004  |  Pubmed ID: 15182723

Emotional events are remembered better than neutral events possibly because the amygdala enhances the function of medial temporal lobe (MTL) memory system (modulation hypothesis). Although this hypothesis has been supported by much animal research, evidence from humans has been scarce and indirect. We investigated this issue using event-related fMRI during encoding of emotional and neutral pictures. Memory performance after scanning showed a retention advantage for emotional pictures. Successful encoding activity in the amygdala and MTL memory structures was greater and more strongly correlated for emotional than for neutral pictures. Moreover, a double dissociation was found along the longitudinal axis of the MTL memory system: activity in anterior regions predicted memory for emotional items, whereas activity in posterior regions predicted memory for neutral items. These results provide direct evidence for the modulation hypothesis in humans and reveal a functional specialization within the MTL regarding the effects of emotion on memory formation.

Dissociable Effects of Arousal and Valence on Prefrontal Activity Indexing Emotional Evaluation and Subsequent Memory: an Event-related FMRI Study

NeuroImage. Sep, 2004  |  Pubmed ID: 15325353

Prefrontal cortex (PFC) activity associated with emotional evaluation and subsequent memory was investigated with event-related functional MRI (fMRI). Participants were scanned while rating the pleasantness of emotionally positive, negative, and neutral pictures, and memory for the pictures was tested after scanning. Emotional evaluation was measured by comparing activity during the picture rating task relative to baseline, and successful encoding was measured by comparing activity for subsequently remembered versus forgotten pictures (Dm effect). The effect of arousal on these measures was indicated by greater activity for both positive and negative pictures than for neutral ones, and the effect of valence was indicated by differences in activity between positive and negative pictures. The study yielded three main results. First, consistent with the valence hypothesis, specific regions in left dorsolateral PFC were more activated for positive than for negative picture evaluation, whereas regions in right ventrolateral PFC showed the converse pattern. Second, dorsomedial PFC activity was sensitive to emotional arousal, whereas ventromedial PFC activity was sensitive to positive valence, consistent with evidence linking these regions, respectively, to emotional processing and self-awareness or appetitive behavior. Finally, successful encoding (Dm) activity in left ventrolateral and dorsolateral PFC was greater for arousing than for neutral pictures. This finding suggests that the enhancing effect of emotion on memory formation is partly due to an augmentation of PFC-mediated strategic, semantic, and working memory operations. These results underscore the critical role of PFC in emotional evaluation and memory, and disentangle the effects of arousal and valence across PFC regions associated with different cognitive functions.

Impact of Healthy Aging on Awareness and Fear Conditioning

Behavioral Neuroscience. Oct, 2004  |  Pubmed ID: 15506873

Fear conditioning has provided a useful model system for studying associative emotional learning, but the impact of healthy aging has gone relatively unexplored. The present study investigated fear conditioning across the adult life span in humans. A delay discrimination task was employed using visual conditioned stimuli and an auditory unconditioned stimulus. Awareness of the reinforcement contingencies was assessed in a postexperimental interview. Compared with young adult participants, middle-aged and older adults displayed reductions in unconditioned responding, discriminant conditioning, and contingency awareness. When awareness and overall arousability were taken into consideration, there were no residual effects of aging on conditioning. These results highlight the importance of considering the influence of declarative knowledge when interpreting age-associated changes in discriminative conditioned learning.

Brain Activity During Episodic Retrieval of Autobiographical and Laboratory Events: an FMRI Study Using a Novel Photo Paradigm

Journal of Cognitive Neuroscience. Nov, 2004  |  Pubmed ID: 15622612

Functional neuroimaging studies of episodic memory retrieval generally measure brain activity while participants remember items encountered in the laboratory ("controlled laboratory condition") or events from their own life ("open autobiographical condition"). Differences in activation between these conditions may reflect differences in retrieval processes, memory remoteness, emotional content, retrieval success, self-referential processing, visual/spatial memory, and recollection. To clarify the nature of these differences, a functional MRI study was conducted using a novel "photo paradigm," which allows greater control over the autobiographical condition, including a measure of retrieval accuracy. Undergraduate students took photos in specified campus locations ("controlled autobiographical condition"), viewed in the laboratory similar photos taken by other participants (controlled laboratory condition), and were then scanned while recognizing the two kinds of photos. Both conditions activated a common episodic memory network that included medial temporal and prefrontal regions. Compared with the controlled laboratory condition, the controlled autobiographical condition elicited greater activity in regions associated with self-referential processing (medial prefrontal cortex), visual/spatial memory (visual and parahippocampal regions), and recollection (hippocampus). The photo paradigm provides a way of investigating the functional neuroanatomy of real-life episodic memory under rigorous experimental control.

Emotional Intensity Predicts Autobiographical Memory Experience

Memory & Cognition. Oct, 2004  |  Pubmed ID: 15813494

College students generated autobiographical memories from distinct emotional categories that varied in valence (positive vs. negative) and intensity (high vs. low). They then rated various perceptual, cognitive, and emotional properties for each memory. The distribution of these emotional memories favored a vector model over a circumplex model. For memories of all specific emotions, intensity accounted for significantly more variance in autobiographical memory characteristics than did valence or age of the memory. In two additional experiments, we examined multiple memories of emotions of high intensity and positive or negative valence and of positive valence and high or low intensity. Intensity was a more consistent predictor of autobiographical memory properties than was valence or the age of the memory in these experiments as well. The general effects of emotion on autobiographical memory properties are due primarily to intensity differences in emotional experience, not to benefits or detriments associated with a specific valence.

Reinstatement of Conditioned Fear in Humans is Context Dependent and Impaired in Amnesia

Behavioral Neuroscience. Jun, 2005  |  Pubmed ID: 15998188

A contextual reinstatement procedure was developed to assess the contributions of environmental cues and hippocampal function in the recovery of conditioned fear following extinction in humans. Experiment 1 showed context specificity in the recovery of extinguished skin conductance responses after presentations of an auditory unconditioned stimulus. Experiment 2 demonstrated that fear recovery did not generalize to an explicitly unpaired conditioned stimulus. Experiment 3 replicated the context dependency of fear recovery with a shock as an unconditioned stimulus. Two amnesic patients failed to recover fear responses following reinstatement in the same context, despite showing initial fear acquisition. These results extend the known functions of the human hippocampus and highlight the importance of environmental contexts in regulating the expression of latent fear associations.

Remembering One Year Later: Role of the Amygdala and the Medial Temporal Lobe Memory System in Retrieving Emotional Memories

Proceedings of the National Academy of Sciences of the United States of America. Feb, 2005  |  Pubmed ID: 15703295

The memory-enhancing effect of emotion can be powerful and long-lasting. Most studies investigating the neural bases of this phenomenon have focused on encoding and early consolidation processes, and hence little is known regarding the contribution of retrieval processes, particularly after lengthy retention intervals. To address this issue, we used event-related functional MRI to measure neural activity during the retrieval of emotional and neutral pictures after a retention interval of 1 yr. Retrieval activity for emotional and neutral pictures was separately analyzed for successfully (hits) vs. unsuccessfully (misses) retrieved items and for responses based on recollection vs. familiarity. Recognition performance was better for emotional than for neutral pictures, and this effect was found only for recollection-based responses. Successful retrieval of emotional pictures elicited greater activity than successful retrieval of neutral pictures in the amygdala, entorhinal cortex, and hippocampus. Moreover, in the amygdala and hippocampus, the emotion effect was greater for recollection than for familiarity, whereas in the entorhinal cortex, it was similar for both forms of retrieval. These findings clarify the role of the amygdala and the medial temporal lobe memory regions in recollection and familiarity of emotional memory after lengthy retention intervals.

Co-activation of the Amygdala, Hippocampus and Inferior Frontal Gyrus During Autobiographical Memory Retrieval

Neuropsychologia. 2005  |  Pubmed ID: 15721179

Functional MRI was used to investigate the role of medial temporal lobe and inferior frontal lobe regions in autobiographical recall. Prior to scanning, participants generated cue words for 50 autobiographical memories and rated their phenomenological properties using our autobiographical memory questionnaire (AMQ). During scanning, the cue words were presented and participants pressed a button when they retrieved the associated memory. The autobiographical retrieval task was interleaved in an event-related design with a semantic retrieval task (category generation). Region-of-interest analyses showed greater activation of the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval relative to semantic retrieval. In addition, the left inferior frontal gyrus showed a more prolonged duration of activation in the semantic retrieval condition. A targeted correlational analysis revealed pronounced functional connectivity among the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval but not during semantic retrieval. These results support theories of autobiographical memory that hypothesize co-activation of frontotemporal areas during recollection of episodes from the personal past.

Amygdala Activation to Sad Pictures During High-field (4 Tesla) Functional Magnetic Resonance Imaging

Emotion (Washington, D.C.). Mar, 2005  |  Pubmed ID: 15755216

Fear-related processing in the amygdala has been well documented, but its role in signaling other emotions remains controversial. The authors recovered signal loss in the amygdala at high-field strength using an inward spiral pulse sequence and probed its response to pictures varying in their degree of portrayed sadness. These pictures were presented as intermittent task-irrelevant distractors during a concurrent visual oddball task. Relative to neutral distractors, sad distractors elicited greater activation along ventral brain regions, including the amygdala, fusiform gyrus, and inferior frontal gyrus. In contrast, oddball targets engaged dorsal sectors of frontal, parietal, and cingulate cortices. The amygdala's role in emotional evaluation thus extends to images of grief and despair as well as to those depicting violence and threat.

Emotional Enhancement of Perceptual Priming is Preserved in Aging and Early-stage Alzheimer's Disease

Neuropsychologia. 2005  |  Pubmed ID: 16154458

Perceptual priming for emotionally-negative and neutral scenes was tested in early-stage Alzheimer's disease (AD) patients and healthy younger, middle-aged and older adults. In the study phase, participants rated the scenes for their arousal properties. In the test phase, studied and novel scenes were initially presented subliminally, and the exposure duration was gradually increased until a valence categorization was made. The difference in exposure duration required to categorize novel versus studied items was the dependent measure of priming. Aversive content increased the magnitude of priming, an effect that was preserved in healthy aging and AD. Results from an immediate recognition memory test showed that the priming effects could not be attributable to enhanced explicit memory for the aversive scenes. These findings implicate a dissociation between the modulatory effect of emotion across implicit and explicit forms of memory in aging and early-stage AD.

Sex, Stress, and Fear: Individual Differences in Conditioned Learning

Cognitive, Affective & Behavioral Neuroscience. Jun, 2005  |  Pubmed ID: 16180625

It has long been recognized that humans vary in their conditionability, yet the factors that contribute to individual variation in emotional learning remain to be delineated. The goal of the present study was to investigate the relationship among sex, stress hormones, and fear conditioning in humans. Forty-five healthy adults (22 females) underwent differential delay conditioning, using fear-relevant conditioned stimuli and a shock unconditioned stimulus. Salivary cortisol samples were taken at baseline and after acquisition training and a 24-h-delayed retention test. The results showed that acquisition of conditioning significantly correlated with postacquisition cortisol levels in males, but not in females. This sex-specific relationship was found despite similar overall levels of conditioning, unconditioned responding, and cortisol. There was no effect of postacquisition cortisol on consolidation of fear learning in either sex. These findings have implications for the understanding of individual differences in fear acquisition and risk factors for the development of affective disorders.

Startle Modulation During Conscious Emotion Regulation is Arousal-dependent

Behavioral Neuroscience. Aug, 2005  |  Pubmed ID: 16187839

Conscious regulation of negative emotion has been shown to affect human eyeblink startle responses, but whether these results depend on modulation of arousal- or valence-based processes is unknown. The authors presented participants with negative, neutral, and positive pictures and directed them to enhance, maintain, and suppress emotional responses. On emotional picture trials, startle responses decreased as a function of cue in the following order: enhance > maintain > suppress. Analysis of negative and positive picture trials separately revealed similar patterns of startle modulation by emotion regulation. There were no effects of emotion regulation on neutral trials. Results indicate that arousal, not valence, may be critical to startle modulation via conscious emotion regulation.

Cognitive Neuroscience of Emotional Memory

Nature Reviews. Neuroscience. Jan, 2006  |  Pubmed ID: 16371950

Emotional events often attain a privileged status in memory. Cognitive neuroscientists have begun to elucidate the psychological and neural mechanisms underlying emotional retention advantages in the human brain. The amygdala is a brain structure that directly mediates aspects of emotional learning and facilitates memory operations in other regions, including the hippocampus and prefrontal cortex. Emotion-memory interactions occur at various stages of information processing, from the initial encoding and consolidation of memory traces to their long-term retrieval. Recent advances are revealing new insights into the reactivation of latent emotional associations and the recollection of personal episodes from the remote past.

Sequential Ordering of Morphed Faces and Facial Expressions Following Temporal Lobe Damage

Neuropsychologia. 2006  |  Pubmed ID: 16464481

A card ordering task was developed to evaluate the role of the temporal lobe in perceiving subtle featural displacements of faces that contribute to judgments of facial expression and identity. Individuals with varying degrees of temporal lobe damage and healthy controls were required to manually sort cards depicting morphs of facial expressions or facial identities so that the cards were sequentially ordered from one morph endpoint to another. Four morph progressions were used--three emotion morphs (neutral-to-anger, neutral-to-fear, and fear-to-anger) and an identity morph. Five exemplars were given per morph type. Debriefing verified that participants were using feature-level cues to sort the cards. A patient with bilateral amygdala damage due to epilepsy did not differ in her sorting abilities from unilateral temporal lobectomy patients or controls. In contrast, a post-encephalitic patient with widespread left temporal lobe damage showed impairments that were most marked on the fear-to-anger and identity sorts. These results show that amygdala-damaged individuals can use information contained in facial expressions to solve tasks that rely on feature-level analysis, which recruits processing in other temporal lobe regions involved in making fine featural distinctions.

Dissociation of Event-related Potentials Indexing Arousal and Semantic Cohesion During Emotional Word Encoding

Brain and Cognition. Oct, 2006  |  Pubmed ID: 16678953

Event-related potential (ERP) studies have shown that emotional stimuli elicit greater amplitude late positive-polarity potentials (LPPs) than neutral stimuli. This effect has been attributed to arousal, but emotional stimuli are also more semantically coherent than uncategorized neutral stimuli. ERPs were recorded during encoding of positive, negative, uncategorized neutral, and categorized neutral words. Differences in LPP amplitude elicited by emotional versus uncategorized neutral stimuli were evident from 450 to 1000 ms. From 450 to 700 ms, LPP effects at midline and right hemisphere frontal electrodes indexed arousal, whereas LPP effects at left hemisphere centro-parietal electrodes indexed semantic cohesion. This dissociation helps specify the processes underlying emotional stimulus encoding, and suggests the need to control for semantic cohesion in emotional information processing studies.

Mood Alters Amygdala Activation to Sad Distractors During an Attentional Task

Biological Psychiatry. Nov, 2006  |  Pubmed ID: 16713587

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.

Effects of Stress and Sex on Acquisition and Consolidation of Human Fear Conditioning

Learning & Memory (Cold Spring Harbor, N.Y.). Jul-Aug, 2006  |  Pubmed ID: 16847304

We examined the relationship between stress hormone (cortisol) release and acquisition and consolidation of conditioned fear learning in healthy adults. Participants underwent acquisition of differential fear conditioning, and consolidation was assessed in a 24-h delayed extinction test. The acquisition phase was immediately followed by an 11-min psychosocial stress period (arithmetic test combined with a public speech). Salivary cortisol was sampled at various time points before and after acquisition and retention of fear conditioning. Results showed two effects of endogenous cortisol. Post-acquisition cortisol correlated with fear acquisition in male but not female participants. In addition, post-acquisition cortisol correlated with consolidation of fear but only in those participants with high cortisol levels. We conclude that in the short term, a robust and sexually dimorphic relationship exists between fear learning and stress hormone levels. For those participants whose fear learning is accompanied by high stress hormone levels, a long-term relationship exists between cortisol release and memory consolidation. These short-term and long-term effects may relate to the differential involvement of mineralocorticoid and glucocorticoid receptor subtypes, respectively. The findings have implications for understanding the role of stress, sex, and hormones in different stages of fear learning and memory.

Reinstatement of Conditioned Fear and the Hippocampus: an Attentional-associative Model

Behavioural Brain Research. Feb, 2007  |  Pubmed ID: 17178163

An existing attentional-associative model of classical conditioning [Schmajuk N, Lam Y, Gray JA. Latent inhibition: a neural network approach. J Exp Psychol: Anim Behav Process 1996;22:321-49] is applied to the description of reinstatement in animals and humans. According to the model, inhibitory associations between the context (CX) and unconditioned stimulus (US) are formed during extinction, which help preserve the association between the conditioned stimulus (CS) and the US. However, summation and retardation tests fail to reveal these associations because (a) the CX is not attended or (b) a CX-CS configural stimulus formed during extinction is both poorly attended and weakly active during testing. When US presentations and testing occur in the same context, reinstatement is the consequence of a decreased CX inhibition and the increased attention to the CS, which activates the remaining CS-US association. When US presentations occur in the context of extinction but the CS is tested in a different context, reinstatement results from an increased attention to the CS and the combination of CS-CX and CX-US excitatory associations. The assumption that associations between CSs are impaired following neurotoxic hippocampal lesions or in amnesia, is sufficient to describe absence of reinstatement in those cases. However, additional assumptions might be needed to describe the effect of hippocampal lesions on other postextinction manipulations.

Garner Interference Reveals Dependencies Between Emotional Expression and Gaze in Face Perception

Emotion (Washington, D.C.). May, 2007  |  Pubmed ID: 17516809

The relationship between facial expression and gaze processing was investigated with the Garner selective attention paradigm. In Experiment 1, participants performed expression judgments without interference from gaze, but expression interfered with gaze judgments. Experiment 2 replicated these results across different emotions. In both experiments, expression judgments occurred faster than gaze judgments, suggesting that expression was processed before gaze could interfere. In Experiments 3 and 4, the difficulty of the emotion discrimination was increased in two different ways. In both cases, gaze interfered with emotion judgments and vice versa. Furthermore, increasing the difficulty of the emotion discrimination resulted in gaze and expression interactions. Results indicate that expression and gaze interactions are modulated by discriminability. Whereas expression generally interferes with gaze judgments, gaze direction modulates expression processing only when facial emotion is difficult to discriminate.

Dissociable Effects of Conscious Emotion Regulation Strategies on Explicit and Implicit Memory

Emotion (Washington, D.C.). May, 2007  |  Pubmed ID: 17516813

The authors manipulated emotion regulation strategies at encoding and administered explicit and implicit memory tests. In Experiment 1, participants used reappraisal to enhance and decrease the personal relevance of unpleasant and neutral pictures. In Experiment 2, decrease cues were replaced with suppress cues that directed participants to inhibit emotion-expressive behavior. Across experiments, using reappraisal to enhance the personal relevance of pictures improved free recall. By contrast, attempting to suppress emotional displays tended to impair recall, especially compared to the enhance condition. Using reappraisal to decrease the personal relevance of pictures had different effects depending on picture type. Paired with unpleasant pictures, the decrease cue tended to improve recall. Paired with neutral stimuli, the decrease cue tended to impair recall. Emotion regulation did not affect perceptual priming. Results highlight dissociable effects of emotion regulation on explicit and implicit memory, as well as dissociations between regulation strategies with respect to explicit memory.

Perception of Dynamic Changes in Facial Affect and Identity in Autism

Social Cognitive and Affective Neuroscience. Jun, 2007  |  Pubmed ID: 18174910

Despite elegant behavioral descriptions of abnormalities for processing emotional facial expressions and biological motion in autism, identification of the neural mechanisms underlying these abnormalities remains a critical and largely unmet challenge. We compared brain activity with dynamic and static facial expressions in participants with and without high-functioning autism using event-related functional magnetic resonance imaging (fMRI) and three classes of face stimuli-emotion morphs (fearful and angry), identity morphs and static images (fearful, angry and neutral). We observed reduced activity in the amygdala (AMY) and fusiform gyrus (FFG) to dynamic emotional expressions in people with autism. There was also a lack of modulation by dynamic compared with static emotional expressions of social brain regions including the AMY, posterior superior temporal sulcus (STS) region and FFG. We observed equivalent emotion and identity morph-evoked activity in participants with and without autism in a region corresponding to the expected location of the more generally motion-sensitive area MT or V5. We conclude that dysfunctions in key components of the human face processing system including the AMY, FFG and posterior STS region are present in individuals with high-functioning autism, and this dysfunction might contribute to the deficits in processing emotional facial expressions.

Beyond Fear Emotional Memory Mechanisms in the Human Brain

Current Directions in Psychological Science. 2007  |  Pubmed ID: 18604284

Neurobiological accounts of emotional memory have been derived largely from animal models investigating the encoding and retention of memories for events that signal threat. This literature has implicated the amygdala, a structure in the brain's temporal lobe, in the learning and consolidation of fear memories. Its role in fear conditioning has been confirmed, but the human amygdala also interacts with cortical regions to mediate other aspects of emotional memory. These include the encoding and consolidation of pleasant and unpleasant arousing events into long-term memory, the narrowing of focus on central emotional information, the retrieval of prior emotional events and contexts, and the subjective experience of recollection and emotional intensity during retrieval. Along with other mechanisms that do not involve the amygdala, these functions ensure that significant life events leave a lasting impression in memory.

Happy and Fearful Emotion in Cues and Targets Modulate Event-related Potential Indices of Gaze-directed Attentional Orienting

Social Cognitive and Affective Neuroscience. Dec, 2007  |  Pubmed ID: 18626515

The goal of the present study was to characterize the effects of valence in facial cues and object targets on event-related potential (ERPs) indices of gaze-directed orienting. Participants were shown faces at fixation that concurrently displayed dynamic gaze shifts and expression changes from neutral to fearful or happy emotions. Emotionally-salient target objects subsequently appeared in the periphery and were spatially congruent or incongruent with the gaze direction. ERPs were time-locked to target presentation. Three sequential ERP components were modulated by happy emotion, indicating a progression from an expression effect to a gaze-by-expression interaction to a target emotion effect. These effects included larger P1 amplitude over contralateral occipital sites for targets following happy faces, larger centrally distributed N1 amplitude for targets following happy faces with leftward gaze, and faster P3 latency for positive targets. In addition, parietally distributed P3 amplitude was reduced for validly cued targets following fearful expressions. Results are consistent with accounts of attentional broadening and motivational approach by happy emotion, and facilitation of spatially directed attention in the presence of fearful cues. The findings have implications for understanding how socioemotional signals in faces interact with each other and with emotional features of objects in the environment to alter attentional processes.

Quantifying Deficits in the Perception of Fear and Anger in Morphed Facial Expressions After Bilateral Amygdala Damage

Neuropsychologia. Jan, 2007  |  Pubmed ID: 16806315

Amygdala damage has been associated with impairments in perceiving facial expressions of fear. However, deficits in perceiving other emotions, such as anger, and deficits in perceiving emotion blends have not been definitively established. One possibility is that methods used to index expression perception are susceptible to heuristic use, which may obscure impairments. To examine this, we adapted a task used to examine categorical perception of morphed facial expressions [Etcoff, N. L., & Magee, J. J. (1992). Categorical perception of facial expressions. Cognition, 44(3), 227-240]. In one version of the task, expressions were categorized with unlimited time constraints. In the other, expressions were presented with limited exposure durations to tap more automatic aspects of processing. Three morph progressions were employed: neutral to anger, neutral to fear, and fear to anger. Both tasks were administered to a participant with bilateral amygdala damage (S.P.), age- and education-matched controls, and young controls. The second task was also administered to unilateral temporal lobectomy patients. In the first version, S.P. showed impairments relative to normal controls on the neutral-to-anger and fear-to-anger morphs, but not on the neutral-to-fear morph. However, reaction times suggested that speed-accuracy tradeoffs could account for results. In the second version, S.P. showed impairments on all morph types relative to all other subject groups. A third experiment showed that this deficit did not extend to the perception of morphed identities. These results imply that when heuristics use is discouraged on tasks utilizing subtle emotion transitions, deficits in the perception of anger and anger/fear blends, as well as fear, are evident with bilateral amygdala damage.

Development of Emotional Facial Recognition in Late Childhood and Adolescence

Developmental Science. Sep, 2007  |  Pubmed ID: 17683341

The ability to interpret emotions in facial expressions is crucial for social functioning across the lifespan. Facial expression recognition develops rapidly during infancy and improves with age during the preschool years. However, the developmental trajectory from late childhood to adulthood is less clear. We tested older children, adolescents and adults on a two-alternative forced-choice discrimination task using morphed faces that varied in emotional content. Actors appeared to pose expressions that changed incrementally along three progressions: neutral-to-fear, neutral-to-anger, and fear-to-anger. Across all three morph types, adults displayed more sensitivity to subtle changes in emotional expression than children and adolescents. Fear morphs and fear-to-anger blends showed a linear developmental trajectory, whereas anger morphs showed a quadratic trend, increasing sharply from adolescents to adults. The results provide evidence for late developmental changes in emotional expression recognition with some specificity in the time course for distinct emotions.

Anorexia Nervosa and Autism Spectrum Disorders: Guided Investigation of Social Cognitive Endophenotypes

Psychological Bulletin. Nov, 2007  |  Pubmed ID: 17967091

Death by suicide occurs in a disproportionate percentage of individuals with anorexia nervosa (AN), with a standardized mortality ratio indicating a 57-fold greater risk of death from suicide relative to an age-matched cohort. Longitudinal studies indicate impaired social functioning increases risk for fatal outcomes, while social impairment persists following recovery. Study of social cognition in AN may elucidate impaired processes that may influence therapeutic efficacy. Symptoms of autism spectrum disorders (ASD) are overrepresented in those who evidence a chronic course. Relative to that in AN, social information processing in ASD is well characterized and may inform systematic study in AN. This article (a) reviews impaired interpersonal processes in AN, (b) compares the phenotype of AN with that of ASD, (c) highlights deficits of social cognitive disturbance in ASD relative to AN, and (d) proposes a new framework to understand the interaction of individuals with AN with their social context.

Neural Systems for Executive and Emotional Processing Are Modulated by Symptoms of Posttraumatic Stress Disorder in Iraq War Veterans

Psychiatry Research. Jan, 2008  |  Pubmed ID: 18093809

The symptom-provocation paradigms generally used in neuroimaging studies of posttraumatic stress disorder (PTSD) have placed high demands on emotion processing but lacked cognitive processing, thereby limiting the ability to assess alterations in neural systems that subserve executive functions and their interactions with emotion processing. Thirty-nine veterans from Iraq and Afghanistan underwent functional magnetic resonance imaging while exposed to emotional combat-related and neutral civilian scenes interleaved with an executive processing task. Contrast activation maps were regressed against PTSD symptoms as measured by the Davidson Trauma Scale. Activation for emotional compared with neutral stimuli was highly positively correlated with level of PTSD symptoms in ventral frontolimbic regions, notably the ventromedial prefrontal cortex, inferior frontal gyrus, and ventral anterior cingulate gyrus. Conversely, activation for the executive task was negatively correlated with PTSD symptoms in the dorsal executive network, notably the middle frontal gyrus, dorsal anterior cingulate gyrus, and inferior parietal lobule. Thus, there is a strong link between the subjectively assessed behavioral phenomenology of PTSD and objective neurobiological markers. These findings extend the largely symptom provocation-based functional neuroanatomy to provide evidence that interrelated executive and emotional processing systems of the brain are differentially affected by PTSD symptomatology in recently deployed war veterans.

Fear Relevancy, Strategy Use, and Probabilistic Learning of Cue-outcome Associations

Learning & Memory (Cold Spring Harbor, N.Y.). Oct, 2008  |  Pubmed ID: 18832564

The goal of this study was to determine how the fear relevancy of outcomes during probabilistic classification learning affects behavior and strategy use. Novel variants of the "weather prediction" task were created, in which cue cards predicted either looming fearful or neutral outcomes in a between-groups design. Strategy use was examined by goodness-of-fit estimates of response patterns across trial blocks to mathematical models of simple, complex, and nonidentifiable strategies. Participants in the emotional condition who were fearful of the outcomes had greater skin conductance responses compared with controls and performed worse, used suboptimal strategies, and had less insight into the predictive cue features during initial learning. In contrast, nonfearful participants in the emotional condition used more optimal strategies than the other groups by the end of the two training days. Results have implications for understanding how individual differences in fear relevancy alter the impact of emotion on feedback-based learning.

The Short and Long of It: Neural Correlates of Temporal-order Memory for Autobiographical Events

Journal of Cognitive Neuroscience. Jul, 2008  |  Pubmed ID: 18284345

Previous functional neuroimaging studies of temporal-order memory have investigated memory for laboratory stimuli that are causally unrelated and poor in sensory detail. In contrast, the present functional magnetic resonance imaging (fMRI) study investigated temporal-order memory for autobiographical events that were causally interconnected and rich in sensory detail. Participants took photographs at many campus locations over a period of several hours, and the following day they were scanned while making temporal-order judgments to pairs of photographs from different locations. By manipulating the temporal lag between the two locations in each trial, we compared the neural correlates associated with reconstruction processes, which we hypothesized depended on recollection and contribute mainly to short lags, and distance processes, which we hypothesized to depend on familiarity and contribute mainly to longer lags. Consistent with our hypotheses, parametric fMRI analyses linked shorter lags to activations in regions previously associated with recollection (left prefrontal, parahippocampal, precuneus, and visual cortices), and longer lags with regions previously associated with familiarity (right prefrontal cortex). The hemispheric asymmetry in prefrontal cortex activity fits very well with evidence and theories regarding the contributions of the left versus right prefrontal cortex to memory (recollection vs. familiarity processes) and cognition (systematic vs. heuristic processes). In sum, using a novel photo-paradigm, this study provided the first evidence regarding the neural correlates of temporal-order for autobiographical events.

Age-dependent Reduction of Amygdala Volume in Bipolar Disorder

Psychiatry Research. May, 2008  |  Pubmed ID: 18407469

The amygdala is hypothesized to play a critical role in mood regulation, yet its involvement in bipolar disorder remains unclear. The aim of the present study was to compare measurements of amygdala volumes in a relatively large sample of bipolar disorder patients and healthy controls ranging in age from 18 to 49 years. Subjects comprised 54 adult patients meeting DSM-IV criteria for bipolar disorder and 41 healthy controls matched for age, sex, and education. Magnetic resonance imaging (1.5 T) was performed to obtain volumetric measurements of the amygdala using a manual region-of-interest tracing method with software that allowed simultaneous visualization of the amygdala in three orthogonal planes. The anterior head of the hippocampus was removed in the sagittal plane prior to amygdala volumetry measurement. Multiple regression analysis was computed on amygdala volume measurements as a function of diagnosis, age, sex, and cerebral volume. Bipolar patients showed an age-related reduction of amygdala volume, but controls did not. Among bipolar subjects, amygdala volume was unrelated to medication history. There were no significant hemispheric or sex interactions with the main effects. Results support a role for amygdala dysfunction in bipolar disorder which appears most robustly in older relative to younger adult patients. Differential aging effects in bipolar disorder may compromise amygdala integrity and contribute to mood dysregulation.

Prefrontal Mechanisms for Executive Control over Emotional Distraction Are Altered in Major Depression

Psychiatry Research. Jul, 2008  |  Pubmed ID: 18455373

A dysfunction in the interaction between executive function and mood regulation has been proposed as the pathophysiology of depression. However, few studies have investigated the alteration in brain systems related to executive control over emotional distraction in depression. To address this issue, 19 patients with major depressive disorder (MDD) and 20 healthy controls were scanned using functional magnetic resonance imaging. Participants performed an emotional oddball task in which infrequently presented circle targets required detection while sad and neutral pictures were irrelevant novel distractors. Hemodynamic responses were compared for targets, sad distractors, and for targets that followed sad or neutral distractors (Target-after-Sad and Target-after-Neutral). Patients with MDD revealed attenuated activation overall to targets in executive brain regions. Behaviorally, MDD patients were slower in response to Target-after-Sad than Target-after-Neutra stimuli. Patients also revealed a reversed activation pattern from controls in response to this contrast in the left anterior cingulate, insula, right inferior frontal gyrus (IFG), and bilateral middle frontal gyrus. Those patients who engaged the right IFG more during Target-after-Neutral stimuli responded faster to targets, confirming a role of this region in coping with emotional distraction. The results provide direct evidence of an alteration in the neural systems that interplay cognition with mood in MDD.

The Spatiotemporal Dynamics of Autobiographical Memory: Neural Correlates of Recall, Emotional Intensity, and Reliving

Cerebral Cortex (New York, N.Y. : 1991). Jan, 2008  |  Pubmed ID: 17548799

We sought to map the time course of autobiographical memory retrieval, including brain regions that mediate phenomenological experiences of reliving and emotional intensity. Participants recalled personal memories to auditory word cues during event-related functional magnetic resonance imaging (fMRI). Participants pressed a button when a memory was accessed, maintained and elaborated the memory, and then gave subjective ratings of emotion and reliving. A novel fMRI approach based on timing differences capitalized on the protracted reconstructive process of autobiographical memory to segregate brain areas contributing to initial access and later elaboration and maintenance of episodic memories. The initial period engaged hippocampal, retrosplenial, and medial and right prefrontal activity, whereas the later period recruited visual, precuneus, and left prefrontal activity. Emotional intensity ratings were correlated with activity in several regions, including the amygdala and the hippocampus during the initial period. Reliving ratings were correlated with activity in visual cortex and ventromedial and inferior prefrontal regions during the later period. Frontopolar cortex was the only brain region sensitive to emotional intensity across both periods. Results were confirmed by time-locked averages of the fMRI signal. The findings indicate dynamic recruitment of emotion-, memory-, and sensory-related brain regions during remembering and their dissociable contributions to phenomenological features of the memories.

Alterations in the Neural Circuitry for Emotion and Attention Associated with Posttraumatic Stress Symptomatology

Psychiatry Research. Apr, 2009  |  Pubmed ID: 19237269

Information processing models of posttraumatic stress disorder (PTSD) suggest that PTSD is characterized by preferential allocation of attentional resources to potentially threatening stimuli. However, few studies have examined the neural pattern underlying attention and emotion in association with PTSD symptomatology. In the present study, combat veterans with PTSD symptomatology engaged in an emotional oddball task while undergoing functional magnetic resonance imaging (fMRI). Veterans were classified into a high or low symptomatology group based on their scores on the Davidson Trauma Scale (DTS). Participants discriminated infrequent target stimuli (circles) from frequent standards (squares) while emotional and neutral distractors were presented infrequently and irregularly. Results revealed that participants with greater PTSD symptomatology showed enhanced neural activity in ventral-limbic and dorsal regions for emotional stimuli and attenuated activity in dorsolateral prefrontal and parietal regions for attention targets. In the anterior cingulate gyrus, participants with fewer PTSD symptoms showed equivalent responses to attentional and emotional stimuli while the high symptom group showed greater activation for negative emotional stimuli. Taken together, the results suggest that hyperresponsive ventral-limbic activity coupled with altered dorsal-attention and anterior cingulate function may be a neural marker of attention bias in PTSD.

The Role of Trauma-related Distractors on Neural Systems for Working Memory and Emotion Processing in Posttraumatic Stress Disorder

Journal of Psychiatric Research. May, 2009  |  Pubmed ID: 19091328

The relevance of emotional stimuli to threat and survival confers a privileged role in their processing. In PTSD, the ability of trauma-related information to divert attention is especially pronounced. Information unrelated to the trauma may also be highly distracting when it shares perceptual features with trauma material. Our goal was to study how trauma-related environmental cues modulate working memory networks in PTSD. We examined neural activity in participants performing a visual working memory task while distracted by task-irrelevant trauma and non-trauma material. Recent post-9/11 veterans were divided into a PTSD group (n=22) and a trauma-exposed control group (n=20) based on the Davidson trauma scale. Using fMRI, we measured hemodynamic change in response to emotional (trauma-related) and neutral distraction presented during the active maintenance period of a delayed-response working memory task. The goal was to examine differences in functional networks associated with working memory (dorsolateral prefrontal cortex and lateral parietal cortex) and emotion processing (amygdala, ventrolateral prefrontal cortex, and fusiform gyrus). The PTSD group showed markedly different neural activity compared to the trauma-exposed control group in response to task-irrelevant visual distractors. Enhanced activity in ventral emotion processing regions was associated with trauma distractors in the PTSD group, whereas activity in brain regions associated with working memory and attention regions was disrupted by distractor stimuli independent of trauma content. Neural evidence for the impact of distraction on working memory is consistent with PTSD symptoms of hypervigilance and general distractibility during goal-directed cognitive processing.

A Comparison of Automated Segmentation and Manual Tracing for Quantifying Hippocampal and Amygdala Volumes

NeuroImage. Apr, 2009  |  Pubmed ID: 19162198

Large databases of high-resolution structural MR images are being assembled to quantitatively examine the relationships between brain anatomy, disease progression, treatment regimens, and genetic influences upon brain structure. Quantifying brain structures in such large databases cannot be practically accomplished by expert neuroanatomists using hand-tracing. Rather, this research will depend upon automated methods that reliably and accurately segment and quantify dozens of brain regions. At present, there is little guidance available to help clinical research groups in choosing such tools. Thus, our goal was to compare the performance of two popular and fully automated tools, FSL/FIRST and FreeSurfer, to expert hand tracing in the measurement of the hippocampus and amygdala. Volumes derived from each automated measurement were compared to hand tracing for percent volume overlap, percent volume difference, across-sample correlation, and 3-D group-level shape analysis. In addition, sample size estimates for conducting between-group studies were computed for a range of effect sizes. Compared to hand tracing, hippocampal measurements with FreeSurfer exhibited greater volume overlap, smaller volume difference, and higher correlation than FIRST, and sample size estimates with FreeSurfer were closer to hand tracing. Amygdala measurement with FreeSurfer was also more highly correlated to hand tracing than FIRST, but exhibited a greater volume difference than FIRST. Both techniques had comparable volume overlap and similar sample size estimates. Compared to hand tracing, a 3-D shape analysis of the hippocampus showed FreeSurfer was more accurate than FIRST, particularly in the head and tail. However, FIRST more accurately represented the amygdala shape than FreeSurfer, which inflated its anterior and posterior surfaces.

Event-related Potentials Reveal Temporal Staging of Dynamic Facial Expression and Gaze Shift Effects on Attentional Orienting

Social Neuroscience. 2009  |  Pubmed ID: 19274577

Multiple sources of information from the face guide attention during social interaction. The present study modified the Posner cueing paradigm to investigate how dynamic changes in emotional expression and eye gaze in faces affect the neural processing of subsequent target stimuli. Event-related potentials (ERPs) were recorded while participants viewed centrally presented face displays in which gaze direction (left, direct, right) and facial expression (fearful, neutral) covaried in a fully crossed design. Gaze direction was not predictive of peripheral target location. ERP analysis revealed several sequential effects, including: (1) an early enhancement of target processing following fearful faces (P1); (2) an interaction between expression and gaze (N1), with enhanced target processing following fearful faces with rightward gaze; and (3) an interaction between gaze and target location (P3), with enhanced processing for invalidly cued left visual field targets. Behaviorally, participants responded faster to targets following fearful faces and targets presented in the right visual field, in concordance with the P1 and N1 effects, respectively. The findings indicate that two nonverbal social cues-facial expression and gaze direction-modulate attentional orienting across different temporal stages of processing. Results have implications for understanding the mental chronometry of shared attention and social referencing.

Generalization of Conditioned Fear Along a Dimension of Increasing Fear Intensity

Learning & Memory (Cold Spring Harbor, N.Y.). Jul, 2009  |  Pubmed ID: 19553384

The present study investigated the extent to which fear generalization in humans is determined by the amount of fear intensity in nonconditioned stimuli relative to a perceptually similar conditioned stimulus. Stimuli consisted of graded emotionally expressive faces of the same identity morphed between neutral and fearful endpoints. Two experimental groups underwent discriminative fear conditioning between a face stimulus of 55% fear intensity (conditioned stimulus, CS+), reinforced with an electric shock, and a second stimulus that was unreinforced (CS-). In Experiment 1 the CS- was a relatively neutral face stimulus, while in Experiment 2 the CS- was the most fear-intense stimulus. Before and following fear conditioning, skin conductance responses (SCR) were recorded to different morph values along the neutral-to-fear dimension. Both experimental groups showed gradients of generalization following fear conditioning that increased with the fear intensity of the stimulus. In Experiment 1 a peak shift in SCRs extended to the most fear-intense stimulus. In contrast, generalization to the most fear-intense stimulus was reduced in Experiment 2, suggesting that discriminative fear learning procedures can attenuate fear generalization. Together, the findings indicate that fear generalization is broadly tuned and sensitive to the amount of fear intensity in nonconditioned stimuli, but that fear generalization can come under stimulus control. These results reveal a novel form of fear generalization in humans that is not merely based on physical similarity to a conditioned exemplar, and may have implications for understanding generalization processes in anxiety disorders characterized by heightened sensitivity to nonthreatening stimuli.

Rebuttal to Hasan and Pedraza in Comments and Controversies: "Improving the Reliability of Manual and Automated Methods for Hippocampal and Amygdala Volume Measurements"

NeuroImage. Nov, 2009  |  Pubmed ID: 19616634

Here we address the critiques offered by Hasan and Pedraza to our recently published manuscript comparing the performance of two automated segmentation programs, FSL/FIRST and FreeSurfer (Morey R, Petty C, Xu Y, Pannu Hayes J, Wagner H, Lewis D, LaBar K, Styner M, McCarthy G. (2009): A comparison of automated segmentation and manual tracing for quantifying of hippocampal and amygdala volumes. Neuroimage 45:855-866). We provide an assessment and discussion of their specific critiques. Hasan and Pedraza bring up some important points concerning our omission of sample demographic features and inclusion of left and right hemisphere volumes as independent measures in correlational analyses. We present additional data on demographic attributes of our sample and correlations analyzed separately on left and right hemispheres of the amygdala and hippocampus. While their commentary aids the reader to more critically asses our study, it falls short of substantiating that our omissions ought to lead readers to significantly revise their interpretations. Further research will help to disentangle the advantages and limitations of the various freely-available automated segmentation software packages.

Delayed Extinction Attenuates Conditioned Fear Renewal and Spontaneous Recovery in Humans

Behavioral Neuroscience. Aug, 2009  |  Pubmed ID: 19634943

This study investigated whether the retention interval after an aversive learning experience influences the return of fear after extinction training. After fear conditioning, participants underwent extinction training either 5 min or 1 day later and in either the same room (same context) or a different room (context shift). The next day, conditioned fear was tested in the original room. When extinction took place immediately, fear renewal was robust and prolonged for context-shift participants, and spontaneous recovery was observed in the same-context participants. Delayed extinction, by contrast, yielded a brief form of fear renewal that reextinguished within the testing session for context-shift participants, and there was no spontaneous recovery in the same-context participants. The authors conclude that the passage of time allows for memory consolidation processes to promote the formation of distinct yet flexible emotional memory traces that confer an ability to recall extinction, even in an alternate context, and minimize the return of fear. Furthermore, immediate extinction can yield spontaneous recovery and prolong fear renewal. These findings have potential implications for ameliorating fear relapse in anxiety disorders.

Dominance, Politics, and Physiology: Voters' Testosterone Changes on the Night of the 2008 United States Presidential Election

PloS One. 2009  |  Pubmed ID: 19844583

Political elections are dominance competitions. When men win a dominance competition, their testosterone levels rise or remain stable to resist a circadian decline; and when they lose, their testosterone levels fall. However, it is unknown whether this pattern of testosterone change extends beyond interpersonal competitions to the vicarious experience of winning or losing in the context of political elections. Women's testosterone responses to dominance competition outcomes are understudied, and to date, a clear pattern of testosterone changes in response to winning and losing dominance competitions has not emerged.

Stressful Politics: Voters' Cortisol Responses to the Outcome of the 2008 United States Presidential Election

Psychoneuroendocrinology. Jun, 2010  |  Pubmed ID: 19962831

Social subordination can be biologically stressful; when mammals lose dominance contests they have acute increases in the stress hormone cortisol. However, human studies of the effect of dominance contest outcomes on cortisol changes have had inconsistent results. Moreover, human studies have been limited to face-to-face competitions and have heretofore never examined cortisol responses to shifts in political dominance hierarchies. The present study investigated voters' cortisol responses to the outcome of the 2008 United States Presidential election. 183 participants at two research sites (Michigan and North Carolina) provided saliva samples at several time points before and after the announcement of the winner on Election Night. Radioimmunoassay was used to measure levels of cortisol in the saliva samples. In North Carolina, John McCain voters (losers) had increases in post-outcome cortisol levels, whereas Barack Obama voters (winners) had stable post-outcome cortisol levels. The present research provides novel evidence that societal shifts in political dominance can impact biological stress responses in voters whose political party becomes socio-politically subordinate.

Mental Hoop Diaries: Emotional Memories of a College Basketball Game in Rival Fans

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. Feb, 2010  |  Pubmed ID: 20147540

The rivalry between the men's basketball teams of Duke University and the University of North Carolina-Chapel Hill (UNC) is one of the most storied traditions in college sports. A subculture of students at each university form social bonds with fellow fans, develop expertise in college basketball rules, team statistics, and individual players, and self-identify as a member of a fan group. The present study capitalized on the high personal investment of these fans and the strong affective tenor of a Duke-UNC basketball game to examine the neural correlates of emotional memory retrieval for a complex sporting event. Male fans watched a competitive, archived game in a social setting. During a subsequent functional magnetic resonance imaging session, participants viewed video clips depicting individual plays of the game that ended with the ball being released toward the basket. For each play, participants recalled whether or not the shot went into the basket. Hemodynamic signal changes time locked to correct memory decisions were analyzed as a function of emotional intensity and valence, according to the fan's perspective. Results showed intensity-modulated retrieval activity in midline cortical structures, sensorimotor cortex, the striatum, and the medial temporal lobe, including the amygdala. Positively valent memories specifically recruited processing in dorsal frontoparietal regions, and additional activity in the insula and medial temporal lobe for positively valent shots recalled with high confidence. This novel paradigm reveals how brain regions implicated in emotion, memory retrieval, visuomotor imagery, and social cognition contribute to the recollection of specific plays in the mind of a sports fan.

Component Neural Systems for the Creation of Emotional Memories During Free Viewing of a Complex, Real-World Event

Frontiers in Human Neuroscience. 2010  |  Pubmed ID: 20508750

To investigate the neural systems that contribute to the formation of complex, self-relevant emotional memories, dedicated fans of rival college basketball teams watched a competitive game while undergoing functional magnetic resonance imaging (fMRI). During a subsequent recognition memory task, participants were shown video clips depicting plays of the game, stemming either from previously-viewed game segments (targets) or from non-viewed portions of the same game (foils). After an old-new judgment, participants provided emotional valence and intensity ratings of the clips. A data driven approach was first used to decompose the fMRI signal acquired during free viewing of the game into spatially independent components. Correlations were then calculated between the identified components and post-scanning emotion ratings for successfully encoded targets. Two components were correlated with intensity ratings, including temporal lobe regions implicated in memory and emotional functions, such as the hippocampus and amygdala, as well as a midline fronto-cingulo-parietal network implicated in social cognition and self-relevant processing. These data were supported by a general linear model analysis, which revealed additional valence effects in fronto-striatal-insular regions when plays were divided into positive and negative events according to the fan's perspective. Overall, these findings contribute to our understanding of how emotional factors impact distributed neural systems to successfully encode dynamic, personally-relevant event sequences.

FMRI Studies of Successful Emotional Memory Encoding: A Quantitative Meta-analysis

Neuropsychologia. Oct, 2010  |  Pubmed ID: 20688087

Over the past decade, fMRI techniques have been increasingly used to interrogate the neural correlates of successful emotional memory encoding. These investigations have typically aimed to either characterize the contributions of the amygdala and medial temporal lobe (MTL) memory system, replicating results in animals, or delineate the neural correlates of specific behavioral phenomena. It has remained difficult, however, to synthesize these findings into a systems neuroscience account of how networks across the whole-brain support the enhancing effects of emotion on memory encoding. To this end, the present study employed a meta-analytic approach using activation likelihood estimates to assess the anatomical specificity and reliability of event-related fMRI activations related to successful memory encoding for emotional versus neutral information. The meta-analysis revealed consistent clusters within bilateral amygdala, anterior hippocampus, anterior and posterior parahippocampal gyrus, the ventral visual stream, left lateral prefrontal cortex and right ventral parietal cortex. The results within the amygdala and MTL support a wealth of findings from the animal literature linking these regions to arousal-mediated memory effects. The consistency of findings in cortical targets, including the visual, prefrontal, and parietal cortices, underscores the importance of generating hypotheses regarding their participation in emotional memory formation. In particular, we propose that the amygdala interacts with these structures to promote enhancements in perceptual processing, semantic elaboration, and attention, which serve to benefit subsequent memory for emotional material. These findings may motivate future research on emotional modulation of widespread neural systems and the implications of this modulation for cognition.

Brain-derived Neurotrophic Factor Val66met Polymorphism and Hippocampal Activation During Episodic Encoding and Retrieval Tasks

Hippocampus. May, 2010  |  Pubmed ID: 20865733

Brain-derived neurotrophic factor (BDNF) is a neurotrophin which has been shown to regulate cell survival and proliferation, as well as synaptic growth and hippocampal long-term potentiation. A naturally occurring single nucleotide polymorphism in the human BDNF gene (val66met) has been associated with altered intercellular trafficking and regulated secretion of BDNF in met compared to val carriers. Additionally, previous studies have found a relationship between the BDNF val66met genotype and functional activity in the hippocampus during episodic and working memory tasks in healthy young adults. Specifically, studies have found that met carriers exhibit both poorer performance and reduced neural activity within the medial temporal lobe (MTL) when performing episodic memory tasks. However, these studies have not been well replicated and have not considered the role of behavioral differences in the interpretation of neural differences. The current study sought to control for cognitive performance in investigating the role of the BDNF val66met genotype on neural activity associated with episodic memory. Across item and relational memory tests, met carriers exhibited increased MTL activation during both encoding and retrieval stages, compared to noncarriers. The results suggest that met carriers are able to recruit MTL activity to support successful memory processes, and reductions in cognitive performance observed in prior studies are not a ubiquitous effect associated with variants of the BDNF val66met genotype. © 2010 Wiley-Liss, Inc.

Staying Cool when Things Get Hot: Emotion Regulation Modulates Neural Mechanisms of Memory Encoding

Frontiers in Human Neuroscience. 2010  |  Pubmed ID: 21212840

During times of emotional stress, individuals often engage in emotion regulation to reduce the experiential and physiological impact of negative emotions. Interestingly, emotion regulation strategies also influence memory encoding of the event. Cognitive reappraisal is associated with enhanced memory while expressive suppression is associated with impaired explicit memory of the emotional event. However, the mechanism by which these emotion regulation strategies affect memory is unclear. We used event-related fMRI to investigate the neural mechanisms that give rise to memory formation during emotion regulation. Twenty-five participants viewed negative pictures while alternately engaging in cognitive reappraisal, expressive suppression, or passive viewing. As part of the subsequent memory design, participants returned to the laboratory two weeks later for a surprise memory test. Behavioral results showed a reduction in negative affect and a retention advantage for reappraised stimuli relative to the other conditions. Imaging results showed that successful encoding during reappraisal was uniquely associated with greater co-activation of the left inferior frontal gyrus, amygdala, and hippocampus, suggesting a possible role for elaborative encoding of negative memories. This study provides neurobehavioral evidence that engaging in cognitive reappraisal is advantageous to both affective and mnemonic processes.

COMT Val108/158 Met Genotype Affects Neural but Not Cognitive Processing in Healthy Individuals

Cerebral Cortex (New York, N.Y. : 1991). Mar, 2010  |  Pubmed ID: 19641018

The relationship between cognition and a functional polymorphism in the catechol-O-methlytransferase (COMT) gene, val108/158met, is one of debate in the literature. Furthermore, based on the dopaminergic differences associated with the COMT val108/158met genotype, neural differences during cognition may be present, regardless of genotypic differences in cognitive performance. To investigate these issues the current study aimed to 1) examine the effects of COMT genotype using a large sample of healthy individuals (n = 496-1218) and multiple cognitive measures, and using a subset of the sample (n = 22), 2) examine whether COMT genotype effects medial temporal lobe (MTL) and frontal activity during successful relational memory processing, and 3) investigate group differences in functional connectivity associated with successful relational memory processing. Results revealed no significant group difference in cognitive performance between COMT genotypes in any of the 19 cognitive measures. However, in the subset sample, COMT val homozygotes exhibited significantly decreased MTL and increased prefrontal activity during both successful relational encoding and retrieval, and reduced connectivity between these regions compared with met homozygotes. Taken together, the results suggest that although the COMT val108/158met genotype has no effect on cognitive behavioral measures in healthy individuals, it is associated with differences in neural process underlying cognitive output.

Neurobehavioral Mechanisms of Human Fear Generalization

NeuroImage. Apr, 2011  |  Pubmed ID: 21256233

While much research has elucidated the neurobiology of fear learning, the neural systems supporting the generalization of learned fear are unknown. Using functional magnetic resonance imaging (fMRI), we show that regions involved in the acquisition of fear support the generalization of fear to stimuli that are similar to a learned threat, but vary in fear intensity value. Behaviorally, subjects retrospectively misidentified a learned threat as a more intense stimulus and expressed greater skin conductance responses (SCR) to generalized stimuli of high intensity. Brain activity related to intensity-based fear generalization was observed in the striatum, insula, thalamus/periacqueductal gray, and subgenual cingulate cortex. The psychophysiological expression of generalized fear correlated with amygdala activity, and connectivity between the amygdala and extrastriate visual cortex was correlated with individual differences in trait anxiety. These findings reveal the brain regions and functional networks involved in flexibly responding to stimuli that resemble a learned threat. These regions may comprise an intensity-based fear generalization circuit that underlies retrospective biases in threat value estimation and overgeneralization of fear in anxiety disorders.

Conceptual Similarity Promotes Generalization of Higher Order Fear Learning

Learning & Memory (Cold Spring Harbor, N.Y.). 2011  |  Pubmed ID: 21330378

We tested the hypothesis that conceptual similarity promotes generalization of conditioned fear. Using a sensory preconditioning procedure, three groups of subjects learned an association between two cues that were conceptually similar, unrelated, or mismatched. Next, one of the cues was paired with a shock. The other cue was then reintroduced to test for fear generalization, as measured by the skin conductance response. Results showed enhanced fear generalization that correlated with trait anxiety levels in the group that learned an association between conceptually similar stimuli. These findings suggest that conceptual representations of conditional stimuli influence human fear learning processes.

Low- and High-testosterone Individuals Exhibit Decreased Aversion to Economic Risk

Psychological Science. Apr, 2011  |  Pubmed ID: 21393575

Testosterone is positively associated with risk-taking behavior in social domains (e.g., crime, physical aggression). However, the scant research linking testosterone to economic risk preferences presents inconsistent findings. We examined the relationship between endogenous testosterone and individuals' economic preferences (i.e., risk preference, ambiguity preference, and loss aversion) in a large sample (N = 298) of men and women. We found that endogenous testosterone levels have a significant U-shaped association with individuals' risk and ambiguity preferences, but not loss aversion. Specifically, individuals with low or high levels of testosterone (more than 1.5 SD from the mean for their gender) were risk and ambiguity neutral, whereas individuals with intermediate levels of testosterone were risk and ambiguity averse. This relationship was highly similar in men and women. In contrast to received wisdom regarding testosterone and risk, the present data provide the first robust evidence for a nonlinear association between economic preferences and levels of endogenous testosterone.

Reprint Of: FMRI Studies of Successful Emotional Memory Encoding: a Quantitative Meta-analysis

Neuropsychologia. Mar, 2011  |  Pubmed ID: 21414466

Over the past decade, fMRI techniques have been increasingly used to interrogate the neural correlates of successful emotional memory encoding. These investigations have typically aimed to either characterize the contributions of the amygdala and medial temporal lobe (MTL) memory system, replicating results in animals, or delineate the neural correlates of specific behavioral phenomena. It has remained difficult, however, to synthesize these findings into a systems neuroscience account of how networks across the whole-brain support the enhancing effects of emotion on memory encoding. To this end, the present study employed a meta-analytic approach using activation likelihood estimates to assess the anatomical specificity and reliability of event-related fMRI activations related to successful memory encoding for emotional versus neutral information. The meta-analysis revealed consistent clusters within bilateral amygdala, anterior hippocampus, anterior and posterior parahippocampal gyrus, the ventral visual stream, left lateral prefrontal cortex and right ventral parietal cortex. The results within the amygdala and MTL support a wealth of findings from the animal literature linking these regions to arousal-mediated memory effects. The consistency of findings in cortical targets, including the visual, prefrontal, and parietal cortices, underscores the importance of generating hypotheses regarding their participation in emotional memory formation. In particular, we propose that the amygdala interacts with these structures to promote enhancements in perceptual processing, semantic elaboration, and attention, which serve to benefit subsequent memory for emotional material. These findings may motivate future research on emotional modulation of widespread neural systems and the implications of this modulation for cognition.

Neurocognitive Mechanisms of Fear Conditioning and Vulnerability to Anxiety

Frontiers in Human Neuroscience. 2011  |  Pubmed ID: 21519378

Hemodynamic Signals of Mixed Messages During a Social Exchange

Neuroreport. Jun, 2011  |  Pubmed ID: 21602650

This study used functional magnetic resonance imaging to characterize hemodynamic activation patterns recruited when the participants viewed mixed social communicative messages during a common interpersonal exchange. Mixed messages were defined as conflicting sequences of biological motion and facial affect signals that are unexpected within a particular social context (e.g. observing the reception of a gift). Across four social vignettes, valenced facial expressions were crossed with rejecting and accepting gestures in a virtual avatar responding to presentation of a gift from the participant. The results indicate that conflicting facial affect and gesture activated superior temporal sulcus, a region implicated in expectancy violations, as well as inferior frontal gyrus and putamen. Scenarios conveying rejection differentially activated the insula and putamen, regions implicated in embodied cognition, and motivated learning, as well as frontoparietal cortex. Characterizing how meaning is inferred from integration of conflicting nonverbal communicative cues is essential to understand nuances and complexities of human exchange.

Anticipatory Anxiety Hinders Detection of a Second Target in Dual-target Search

Psychological Science. Jul, 2011  |  Pubmed ID: 21670427

Professional visual searches (e.g., baggage screenings, military searches, radiological examinations) are often conducted in high-pressure environments and require focus on multiple visual targets. Yet laboratory studies of visual search tend to be conducted in emotionally neutral settings with only one possible target per display. In the experiment reported here, we looked to better emulate high-pressure search conditions by presenting searchers with arrays that contained between zero and two targets while inducing anticipatory anxiety via a threat-of-shock paradigm. Under conditions of anticipatory anxiety, dual-target performance was negatively affected, but single-target performance and time on task were unaffected. These results suggest that multiple-target searches may be a more sensitive instrument to measure the effect of environmental factors on visual cognition than single-target searches are. Further, the effect of anticipatory anxiety was modulated by individual differences in state anxiety levels of participants prior to the experiment. These results have implications for both the laboratory study of visual search and the management and assessment of professional searchers.

Cracking the Almond (Commentary on Prévost Et Al.)

The European Journal of Neuroscience. Jul, 2011  |  Pubmed ID: 21722206

Structural Magnetic Resonance Imaging in Bipolar Disorder: an International Collaborative Mega-analysis of Individual Adult Patient Data

Biological Psychiatry. Feb, 2011  |  Pubmed ID: 21030008

There is substantial inconsistency in results of brain structural magnetic resonance imaging studies in adult bipolar disorder. This is likely consequent upon limited statistical power of studies together with their clinical and methodological heterogeneity. The current study was undertaken to perform an international collaborative mega-analysis of regional volumetric measurements of individual patient and healthy subject data, to optimize statistical power, detect case-control differences, assess the association of psychotropic medication usage with brain structural variation, and detect other possible sources of heterogeneity.

Reduced Hippocampal and Amygdala Activity Predicts Memory Distortions for Trauma Reminders in Combat-related PTSD

Journal of Psychiatric Research. May, 2011  |  Pubmed ID: 21047644

Neurobiological models of posttraumatic stress disorder (PTSD) suggest that altered activity in the medial temporal lobes (MTL) during encoding of traumatic memories contribute to the development and maintenance of the disorder. However, there is little direct evidence in the PTSD literature to support these models. The goal of the present study was to examine MTL activity during trauma encoding in combat veterans using the subsequent memory paradigm. Fifteen combat veterans diagnosed with PTSD and 14 trauma-exposed control participants viewed trauma-related and neutral pictures while undergoing event-related fMRI. Participants returned one week after scanning for a recognition memory test. Region-of-interest (ROI) and voxel-wise whole brain analyses were conducted to examine the neural correlates of successful memory encoding. Patients with PTSD showed greater false alarm rates for novel lures than the trauma-exposed control group, suggesting reliance on gist-based representations in lieu of encoding contextual details. Imaging analyses revealed reduced activity in the amygdala and hippocampus in PTSD patients during successful encoding of trauma-related stimuli. Reduction in left hippocampal activity was associated with high arousal symptoms on the Clinician-Administered PTSD Scale (CAPS). The behavioral false alarm rate for traumatic stimuli co-varied with activity in the bilateral precuneus. These results support neurobiological theories positing reduced hippocampal activity under conditions of high stress and arousal. Reduction in MTL activity for successfully encoded stimuli and increased precuneus activity may underlie reduced stimulus-specific encoding and greater gist memory in patients with PTSD, leading to maintenance of the disorder.

Unpredictability and Uncertainty in Anxiety: a New Direction for Emotional Timing Research

Frontiers in Integrative Neuroscience. 2011  |  Pubmed ID: 21954380

Level of Processing Modulates the Neural Correlates of Emotional Memory Formation

Journal of Cognitive Neuroscience. Apr, 2011  |  Pubmed ID: 20350176

Emotion is known to influence multiple aspects of memory formation, including the initial encoding of the memory trace and its consolidation over time. However, the neural mechanisms whereby emotion impacts memory encoding remain largely unexplored. The present study used a levels-of-processing manipulation to characterize the impact of emotion on encoding with and without the influence of elaborative processes. Participants viewed emotionally negative, neutral, and positive scenes under two conditions: a shallow condition focused on the perceptual features of the scenes and a deep condition that queried their semantic meaning. Recognition memory was tested 2 days later. Results showed that emotional memory enhancements were greatest in the shallow condition. fMRI analyses revealed that the right amygdala predicted subsequent emotional memory in the shallow more than deep condition, whereas the right ventrolateral PFC demonstrated the reverse pattern. Furthermore, the association of these regions with the hippocampus was modulated by valence: the amygdala-hippocampal link was strongest for negative stimuli, whereas the prefrontal-hippocampal link was strongest for positive stimuli. Taken together, these results suggest two distinct activation patterns underlying emotional memory formation: an amygdala component that promotes memory during shallow encoding, especially for negative information, and a prefrontal component that provides extra benefits during deep encoding, especially for positive information.

Cognitive and Neural Contributors to Emotion Regulation in Aging

Social Cognitive and Affective Neuroscience. Apr, 2011  |  Pubmed ID: 20385663

Older adults, compared to younger adults, focus on emotional well-being. While the lifespan trajectory of emotional processing and its regulation has been characterized behaviorally, few studies have investigated the underlying neural mechanisms. Here, older adults (range: 59-73 years) and younger adults (range: 19-33 years) participated in a cognitive reappraisal task during functional magnetic resonance imaging (fMRI) scanning. On each trial, participants viewed positive, negative or neutral pictures and either naturally experienced the image ('Experience' condition) or attempted to detach themselves from the image ('Reappraise' condition). Across both age groups, cognitive reappraisal activated prefrontal regions similar to those reported in prior studies of emotion regulation, while emotional experience activated the bilateral amygdala. Psychophysiological interaction analyses revealed that the left inferior frontal gyrus (IFG) and amygdala demonstrated greater inverse connectivity during the 'Reappraise' condition relative to the 'Experience' condition. The only regions exhibiting significant age differences were the left IFG and the left superior temporal gyrus, for which greater regulation-related activation was observed in younger adults. Controlling for age, increased performance on measures of cognition predicted greater regulation-related decreases in amygdala activation. Thus, while older and younger adults use similar brain structures for emotion regulation and experience, the functional efficacy of those structures depends on underlying cognitive ability.

Is All Motivation Good for Learning? Dissociable Influences of Approach and Avoidance Motivation in Declarative Memory

Learning & Memory (Cold Spring Harbor, N.Y.). 2011  |  Pubmed ID: 22021253

The present study investigated the effects of approach versus avoidance motivation on declarative learning. Human participants navigated a virtual reality version of the Morris water task, a classic spatial memory paradigm, adapted to permit the experimental manipulation of motivation during learning. During this task, participants were instructed to navigate to correct platforms while avoiding incorrect platforms. To manipulate motivational states participants were either rewarded for navigating to correct locations (approach) or punished for navigating to incorrect platforms (avoidance). Participants' skin conductance levels (SCLs) were recorded during navigation to investigate the role of physiological arousal in motivated learning. Behavioral results revealed that, overall, approach motivation enhanced and avoidance motivation impaired memory performance compared to nonmotivated spatial learning. This advantage was evident across several performance indices, including accuracy, learning rate, path length, and proximity to platform locations during probe trials. SCL analysis revealed three key findings. First, within subjects, arousal interacted with approach motivation, such that high arousal on a given trial was associated with performance deficits. In addition, across subjects, high arousal negated or reversed the benefits of approach motivation. Finally, low-performing, highly aroused participants showed SCL responses similar to those of avoidance-motivation participants, suggesting that for these individuals, opportunities for reward may evoke states of learning similar to those typically evoked by threats of punishment. These results provide a novel characterization of how approach and avoidance motivation influence declarative memory and indicate a critical and selective role for arousal in determining how reinforcement influences goal-oriented learning.

Revealing Context-specific Conditioned Fear Memories with Full Immersion Virtual Reality

Frontiers in Behavioral Neuroscience. 2011  |  Pubmed ID: 22069384

The extinction of conditioned fear is known to be context-specific and is often considered more contextually bound than the fear memory itself (Bouton, 2004). Yet, recent findings in rodents have challenged the notion that contextual fear retention is initially generalized. The context-specificity of a cued fear memory to the learning context has not been addressed in the human literature largely due to limitations in methodology. Here we adapt a novel technology to test the context-specificity of cued fear conditioning using full immersion 3-D virtual reality (VR). During acquisition training, healthy participants navigated through virtual environments containing dynamic snake and spider conditioned stimuli (CSs), one of which was paired with electrical wrist stimulation. During a 24-h delayed retention test, one group returned to the same context as acquisition training whereas another group experienced the CSs in a novel context. Unconditioned stimulus expectancy ratings were assayed on-line during fear acquisition as an index of contingency awareness. Skin conductance responses time-locked to CS onset were the dependent measure of cued fear, and skin conductance levels during the interstimulus interval were an index of context fear. Findings indicate that early in acquisition training, participants express contingency awareness as well as differential contextual fear, whereas differential cued fear emerged later in acquisition. During the retention test, differential cued fear retention was enhanced in the group who returned to the same context as acquisition training relative to the context shift group. The results extend recent rodent work to illustrate differences in cued and context fear acquisition and the contextual specificity of recent fear memories. Findings support the use of full immersion VR as a novel tool in cognitive neuroscience to bridge rodent models of contextual phenomena underlying human clinical disorders.

Neural Systems for Guilt from Actions Affecting Self Versus Others

NeuroImage. Jan, 2012  |  Pubmed ID: 22230947

Guilt is a core emotion governing social behavior by promoting compliance with social norms or self-imposed standards. The goal of this study was to contrast guilty responses to actions that affect self versus others, since actions with social consequences are hypothesized to yield greater guilty feelings due to adopting the perspective and subjective emotional experience of others. Sixteen participants were presented with brief hypothetical scenarios in which the participant's actions resulted in harmful consequences to self (guilt-self) or to others (guilt-other) during functional MRI. Participants felt more intense guilt for guilt-other than guilt-self and guilt-neutral scenarios. Guilt scenarios revealed distinct regions of activity correlated with intensity of guilt, social consequences of actions, and the interaction of guilt by social consequence. Guilt intensity was associated with activation of the dorsomedial PFC, superior frontal gyrus, supramarginal gyrus, and anterior inferior frontal gyrus. Guilt accompanied by social consequences was associated with greater activation than without social consequences in the ventromedial and dorsomedial PFC, precuneus, posterior cingulate, and posterior superior temporal sulcus. Finally, the interaction analysis highlighted select regions that were more strongly correlated with guilt intensity as a function of social consequence, including the left anterior inferior frontal gyrus, left ventromedial PFC, and left anterior inferior parietal cortex. Our results suggest these regions intensify guilt where harm to others may incur a greater social cost.

Role of Conceptual Knowledge in Learning and Retention of Conditioned Fear

Biological Psychology. Feb, 2012  |  Pubmed ID: 22118937

Associating sensory cues with aversive outcomes is a relatively basic process shared across species. Yet higher-order cognitive processes likely contribute to associative fear learning in many circumstances, especially in humans. Here we ask whether fears can be acquired based on conceptual knowledge of object categories, and whether such concept-based fear conditioning leads to enhanced memory representations for conditioned objects. Participants were presented with a heterogeneous collection of images of animals and tools. Objects from one category were reinforced by an electrical shock, whereas the other category was never reinforced. Results confirmed concept-based fear learning through subjective report of shock expectancy, heightened skin conductance responses, and enhanced 24h recognition memory for items from the conditioned category. These results provide novel evidence that conditioned fear can generalize through knowledge of object concepts, and sheds light on the persistent nature of fear memories and category-based fear responses symptomatic of some anxiety disorders.

Neurocognitive Mechanisms of Gaze-expression Interactions in Face Processing and Social Attention

Neuropsychologia. Jan, 2012  |  Pubmed ID: 22285906

The face conveys a rich source of non-verbal information used during social communication. While research has revealed how specific facial channels such as emotional expression are processed, little is known about the prioritization and integration of multiple cues in the face during dyadic exchanges. Classic models of face perception have emphasized the segregation of dynamic vs. static facial features along independent information processing pathways. Here we review recent behavioral and neuroscientific evidence suggesting that within the dynamic stream, concurrent changes in eye gaze and emotional expression can yield early independent effects on face judgments and covert shifts of visuospatial attention. These effects are partially segregated within initial visual afferent processing volleys, but are subsequently integrated in limbic regions such as the amygdala or via reentrant visual processing volleys. This spatiotemporal pattern may help to resolve otherwise perplexing discrepancies across behavioral studies of emotional influences on gaze-directed attentional cueing. Theoretical explanations of gaze-expression interactions are discussed, with special consideration of speed-of-processing (discriminability) and contextual (ambiguity) accounts. Future research in this area promises to reveal the mental chronometry of face processing and interpersonal attention, with implications for understanding how social referencing develops in infancy and is impaired in autism and other disorders of social cognition.

Fear-relevant Outcomes Modulate the Neural Correlates of Probabilistic Classification Learning

NeuroImage. Jan, 2012  |  Pubmed ID: 21827859

Although much work has implicated the contributions of frontostriatal and medial temporal lobe (MTL) systems during probabilistic classification learning, the impact of emotion on these learning circuits is unknown. We used a modified version of the weather prediction task in which two participant groups were scanned with identical neutral cue cards probabilistically linked to either emotional (snake/spider) or neutral (mushroom/flower) outcomes. Owing to the differences in visual information shown as outcomes, analyses were restricted to the cue phase of the trials. Learning rates did not differ between the two groups, although the Emotional group was more likely to use complex strategies and to respond more slowly during initial learning. The Emotional group had reduced frontostriatal and MTL activation relative to the Neutral group, especially for participants who scored higher on snake/spider phobia questionnaires. Accurate performance was more tied to medial prefrontal activity in the Emotional group early in training, and to MTL activity in the Neutral group later in training. Trial-by-trial fluctuations in functional connectivity between the caudate and MTL were also reduced in the Emotional group compared to the Neutral group. Across groups, reaction time indexed a switch in learning systems, with faster trials mediated by the caudate and slower trials mediated by the MTL and frontal lobe. The extent to which the caudate was activated early in training predicted later performance improvements. These results reveal insights into how emotional outcomes modulate procedural learning systems, and the dynamics of MTL-striatal engagement across training trials.