ARTICLE UPDATE - Influence of attention to somatic information on emotional and autonomic responses.

Murakami H, Ohira H, Matsunaga M, Kimura K.

Perceptual Motor Skills, 108, 531-539

The present study aimed to investigate the dissociable effects of two forms of self-focus on emotional and autonomic responses. One form is suppression, which includes the suppression of heart rate and self-evaluation of performance. The other is observation, which includes attention to one's own heart rate with no suppression and no evaluation. 26 undergraduate and graduate students from the Nagoya University campus (13 men, 13 women), ages 18 to 24 years (M = 20.7, SD = 1.6) were recruited. Participants were provided with their own heart rate as feedback for 5 min., during which participants conducted a self-focus manipulation. Several days after the experimental session for one condition, the same participants conducted another experimental session for the other condition. Instruction to suppress enhanced physiological arousal and subsequent negative emotions; however, instruction to observe did not increase physiological arousal or negative emotions.

ARTICLE UPDATE - Worry tendencies predict brain activation during aversive imagery.

Schienle A, Schäfer A, Pignanelli R, Vaitl D.

Neuroscience Letters, in press

Because of its abstract nature, worrying might function as an avoidance response in order to cognitively disengage from fearful imagery. The present functional magnetic resonance imaging study investigated neural correlates of aversive imagery and their association with worry tendencies, as measured by the Penn State Worry Questionnaire (PSWQ). Nineteen healthy women first viewed, and subsequently imagined pictures from two categories, 'threat' and 'happiness'. Worry tendencies were negatively correlated with brain activation in the anterior cingulate cortex, the prefrontal cortex (dorsolateral, dorsomedial, ventrolateral), the parietal cortex and the insula. These negative correlations between PSWQ scores and localized brain activation were specific for aversive imagery. Moreover, activation in the abovementioned regions was positively associated with the experienced vividness of both pleasant and unpleasant mental pictures. As the identified brain regions are involved in emotion regulation, vivid imagery and memory retrieval, a lowered activity in high PSWQ scorers might be associated with cognitive disengagement from aversive imagery as well as insufficient refresh rates of mental pictures. Our preliminary findings encourage future imagery studies on generalized anxiety disorder patients, as one of the main symptoms of this disorder is excessive worrying.

ARTICLE UPDATE - In search of specificity: functional MRI in the study of emotional experience.

Schienle A, Schäfer A.

International Journal of Psychophysiology, 73, 22-26.

The growing availability of functional magnetic resonance imaging (fMRI) with its property of high spatial resolution has energized the search for specific neural substrates of basic emotions and their feeling components. In the present article, we address the question as to whether recent fMRI studies on primary affective experiences have truly helped to pinpoint emotion-specific areas in the human brain or whether these studies are afflicted with methodological problems which make such inferences difficult. As one approach for improvement, we suggest the combination of fMRI with methods characterized by high temporal resolution, such as electroencephalography (EEG). Simultaneous recoding allows the correlation of temporally specific EEG components (e.g., the late positive potential) with regional blood-oxygen-level-dependent (BOLD) signals during affective experiences. Combined information on the source as well as the exact temporal pattern of a neural affective response will help to improve our understanding of emotion-specific brain activation.

ARTICLE UPDATE - Emotions in motion: Dynamic compared to static facial expressions of disgust and happiness reveal more widespread emotion-specific ac

Emotions in motion: Dynamic compared to static facial expressions of disgust and happiness reveal more widespread emotion-specific activations.

Brain Research, in press

In social contexts, facial expressions are dynamic in nature and vary rapidly in relation to situational requirements. However, there are very few fMRI studies using dynamic emotional stimuli. The aim of this study was (1) to introduce and evaluate a new stimulus database of static and dynamic emotional facial expressions according to arousal and recognizability investigated by a rating by both participants of the present fMRI study and by an external sample of 30 healthy women, (2) to examine the neural networks involved in emotion perception of static and dynamic facial stimuli separately, and (3) to examine the impact of motion on the emotional processing of dynamic compared to static face stimuli. A total of 16 females participated in the present fMRI study performing a passive emotion perception task including static and dynamic faces of neutral, happy and disgusted expressions. Comparing dynamic stimuli to static faces indicated enhanced emotion-specific brain activation patterns in the parahippocampal gyrus (PHG) including the amygdala (AMG), fusiform gyrus (FG), superior temporal gyrus (STG), inferior frontal gyrus (IFG), and occipital and orbitofrontal cortex (OFC). These regions have been discussed to be associated with emotional memory encoding, the perception of threat, facial identity, biological motion, the mirror neuron system, an increase of emotional arousal, and reward processing, respectively. Post hoc ratings of the dynamic stimuli revealed a better recognizability in comparison to the static stimuli. In conclusion, dynamic facial expressions might provide a more appropriate approach to examine the processing of emotional face perception than static stimuli.

ARTICLE UPDATE - Emotion and space. Lateralized emotional word detection depends on line bisection bias.

Tamagni C, Mantei T, Brugger P.

Neuroscience, in press

There is converging evidence, from various independent areas of neuroscience, for a functional specialization of the left and right cerebral hemispheres for positive and negative emotions, respectively ("valence theory" of emotional processing). One subfield, however, has produced mixed results, i.e. work on the detection of parafoveally presented positively or negatively emotional words by healthy subjects. Right or left visual field advantages were described and interpreted as reflecting the superiority of either the left hemisphere (LH) for linguistic material, or of the right hemisphere (RH) for highly emotional stimuli. Here we show that 48 healthy, right-handed participants' performance on a lateralized lexical decision task depends on their individual inclination to bisect a line to the left or right of the objective center. Only those with a bisection bias to the right showed the LH advantage for word detection known from the neuropsychological literature. Negative emotional words were processed with comparable accuracy in the two visual fields. However, a recognition advantage for negative over positive emotional words was found exclusively for those participants with a leftward line bisection bias. These results suggest that in work on functional hemispheric differences state variables like stimulus lateralization and word emotionality may be less decisive than the trait variable of lateral hemispatial attention. We propose a cautious reconsideration of the concept of "hemisphericity", which once emphasized individual differences in baseline hemispheric arousal, but was later dismissed in a reaction to oversimplifications in popular science accounts.

ARTICLE UPDATE - EEG coherence in humans: relationship with success in recognizing emotions in the voice.

Kislova OO, Rusalova MN.

Neuroscience and Behavioral Physiology, in press

EEG recordings from two groups of subjects - with high and low levels of recognition of emotions from voices were made. Comparisons were performed of the numbers of pairs of leads with different levels of coherence in baseline conditions and on recognition of emotions in six standard frequency ranges and in individual bands with 1-Hz steps. Significant differences were seen between groups 1 and 2 both in baseline conditions and during recognition of emotions: in most cases, coherence was greater in subjects with poor recognition of emotions from voices.

ARTICLE UPDATE - Prolonged reduction of electrocortical activity predicts correct performance during rapid serial visual processing.

Keil A, Heim S.

Psychophysiology, in press

Abstract When two targets are shown in a rapid temporal stream of distractors, performance for the second target (T2) is typically reduced when presented between 200 and 500 ms after the first (T1). The present study used the steady-state visual evoked potential (ssVEP), a continuous index of electrocortical facilitation, to compare brain responses in trials with correct versus incorrect T2 responses. We found a reduction of the electrocortical response following T1 in trials with correct T2 identification. By contrast, incorrect T2 trials were characterized by enhanced electrocortical amplitude. Amplitude attenuation predictive of successful T2 report was sustained over time, suggesting a reduction of resources allocated to the distractor stream in correct trials. Across intertarget intervals, T2 performance was a linear function of the ssVEP amplitude reduction in correct trials, weighted by the stimulus onset asynchrony.

PUBLICATION - Affective learning enhances activity and functional connectivity in early visual cortex.

Damaraju E, Huang YM, Barrett LF, Pessoa L.

Neuropsychologia, in press

This study examined the impact of task-irrelevant affective information on early visual processing regions V1-V4. Fearful and neutral faces presented with rings of different colors were used as stimuli. During the conditioning phase, fearful faces presented with a certain ring color (e.g., black) were paired with mild electrical stimulation. Neutral faces shown with rings of that color, as well as fearful or neutral faces shown with another ring color (e.g., white), were never paired with shock. Our findings revealed that fearful faces evoked enhanced blood oxygen level dependent (BOLD) responses in V1 and V4 compared to neutral faces. Faces embedded in a color ring that was paired with shock (e.g., black) evoked greater BOLD responses in V1-V4 compared to a ring color that was never paired with shock (e.g., white). Finally, BOLD responses in early visual cortex were tightly interrelated (i.e., correlated) during an affectively potent context (i.e., ring color) but not during a neutral one, suggesting that increased functional integration was present with affective learning. Taken together, the results suggest that task-irrelevant affective information not only influences evoked responses in early, retinotopically organized visual cortex, but also determines the pattern of responses across early visual cortex.

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ARTICLE UPDATE - Embodiment of emotion concepts.

Niedenthal PM, Winkielman P, Mondillon L, Vermeulen N.

Journal of Personality and Social Psychology, 96, 1120-1136

Theories of embodied cognition hold that higher cognitive processes operate on perceptual symbols and that concept use involves partial reactivations of the sensory-motor states that occur during experience with the world. On this view, the processing of emotion knowledge involves a (partial) reexperience of an emotion, but only when access to the sensory basis of emotion knowledge is required by the task. In 2 experiments, participants judged emotional and neutral concepts corresponding to concrete objects (Experiment 1) and abstract states (Experiment 2) while facial electromyographic activity was recorded from the cheek, brow, eye, and nose regions. Results of both studies show embodiment of specific emotions in an emotion-focused but not a perceptual-focused processing task on the same words. A follow up in Experiment 3, which blocked selective facial expressions, suggests a causal, rather than simply a correlational, role for embodiment in emotion word processing. Experiment 4, using a property generation task, provided support for the conclusion that emotions embodied in conceptual tasks are context-dependent situated simulations rather than associated emotional reactions. Implications for theories of embodied simulation and for emotion theories are discussed.

ARTICLE UPDATE - Early and late temporo-spatial effects of contextual interference during perception of facial affect.

Frühholz S, Fehr T, Herrmann M.

International Journal of Psychophysiology, in press

Contextual features during recognition of facial affect are assumed to modulate the temporal course of emotional face processing. Here, we simultaneously presented colored backgrounds during valence categorizations of facial expressions. Subjects incidentally learned to perceive negative, neutral and positive expressions within a specific colored context. Subsequently, subjects made fast valence judgments while presented with the same face-color-combinations as in the first run (congruent trials) or with different face-color-combinations (incongruent trials). Incongruent trials induced significantly increased response latencies and significantly decreased performance accuracy. Contextual incongruent information during processing of neutral expressions modulated the P1 and the early posterior negativity (EPN) both localized in occipito-temporal areas. Contextual congruent information during emotional face perception revealed an emotion-related modulation of the P1 for positive expressions and of the N170 and the EPN for negative expressions. Highest amplitude of the N170 was found for negative expressions in a negatively associated context and the N170 amplitude varied with the amount of overall negative information. Incongruent trials with negative expressions elicited a parietal negativity which was localized to superior parietal cortex and which most likely represents a posterior manifestation of the N450 as an indicator of conflict processing. A sustained activation of the late LPP over parietal cortex for all incongruent trials might reflect enhanced engagement with facial expression during task conditions of contextual interference. In conclusion, whereas early components seem to be sensitive to the emotional valence of facial expression in specific contexts, late components seem to subserve interference resolution during emotional face processing.

ARTICLE UPDATE - The Interrelations between Verbal Working Memory and Visual Selection of Emotional Faces.

Grecucci A, Soto D, Rumiati RI, Humphreys GW, Rotshtein P.

The Journal of Cognitive Neuroscience, in press

Working memory (WM) and visual selection processes interact in a reciprocal fashion based on overlapping representations abstracted from the physical characteristics of stimuli. Here, we assessed the neural basis of this interaction using facial expressions that conveyed emotion information. Participants memorized an emotional word for a later recognition test and then searched for a face of a particular gender presented in a display with two faces that differed in gender and expression. The relation between the emotional word and the expressions of the target and distractor faces was varied. RTs for the memory test were faster when the target face matched the emotional word held in WM (on valid trials) relative to when the emotional word matched the expression of the distractor (on invalid trials). There was also enhanced activation on valid compared with invalid trials in the lateral orbital gyrus, superior frontal polar (BA 10), lateral occipital sulcus, and pulvinar. Re-presentation of the WM stimulus in the search display led to the earlier onset of activity in the superior and inferior frontal gyri and the anterior hippocampus irrespective of the search validity of the re-presented stimulus. The data indicate that the middle temporal and prefrontal cortices are sensitive to the reappearance of stimuli that are held in WM, whereas a fronto-thalamic occipital network is sensitive to the behavioral significance of the match between WM and targets for selection. We conclude that these networks are modulated by high-level matches between the contents of WM, the behavioral goals, and our current sensory input.

ARTICLE UPDATE - Decoding of Emotional Information in Voice-Sensitive Cortices.

Ethofer T, Van De Ville D, Scherer K, Vuilleumier P.

Current Biology, in press

The ability to correctly interpret emotional signals from others is crucial for successful social interaction. Previous neuroimaging studies showed that voice-sensitive auditory areas [1-3] activate to a broad spectrum of vocally expressed emotions more than to neutral speech melody (prosody). However, this enhanced response occurs irrespective of the specific emotion category, making it impossible to distinguish different vocal emotions with conventional analyses [4-8]. Here, we presented pseudowords spoken in five prosodic categories (anger, sadness, neutral, relief, joy) during event-related functional magnetic resonance imaging (fMRI), then employed multivariate pattern analysis [9, 10] to discriminate between these categories on the basis of the spatial response pattern within the auditory cortex. Our results demonstrate successful decoding of vocal emotions from fMRI responses in bilateral voice-sensitive areas, which could not be obtained by using averaged response amplitudes only. Pairwise comparisons showed that each category could be classified against all other alternatives, indicating for each emotion a specific spatial signature that generalized across speakers. These results demonstrate for the first time that emotional information is represented by distinct spatial patterns that can be decoded from brain activity in modality-specific cortical areas.

ARTICLE UPDATE - Transmission of facial expressions of emotion co-evolved with their efficient decoding in the brain: behavioral and brain evidence.

Schyns PG, Petro LS, Smith ML.

PlosOne

Competent social organisms will read the social signals of their peers. In primates, the face has evolved to transmit the organism's internal emotional state. Adaptive action suggests that the brain of the receiver has co-evolved to efficiently decode expression signals. Here, we review and integrate the evidence for this hypothesis. With a computational approach, we co-examined facial expressions as signals for data transmission and the brain as receiver and decoder of these signals. First, we show in a model observer that facial expressions form a lowly correlated signal set. Second, using time-resolved EEG data, we show how the brain uses spatial frequency information impinging on the retina to decorrelate expression categories. Between 140 to 200 ms following stimulus onset, independently in the left and right hemispheres, an information processing mechanism starts locally with encoding the eye, irrespective of expression, followed by a zooming out to processing the entire face, followed by a zooming back in to diagnostic features (e.g. the opened eyes in "fear", the mouth in "happy"). A model categorizer demonstrates that at 200 ms, the left and right brain have represented enough information to predict behavioral categorization performance.

ARTICLE UPDATE - Involvement of medial prefrontal cortex in emotion during feedback presentation.

Jimura K, Konishi S, Asari T, Miyashita Y.

Neuroreport, in press

It has been suggested that the posterior medial prefrontal cortex (pMPFC) implements cognitive functions involved during negative feedback processing. It has also been suggested that the presentation of the feedback elicits emotional processes. This functional MRI study examined whether pMPFC was associated with the emotional component in feedback processing. Participants were exposed to feedback while performing a version of a motion prediction task. The pMPFC was activated during negative feedback presentation and emotion-related activity was extracted from the pMPFC activation through parametric imaging analysis. It was found that the emotional pMPFC activity was greater in participants who scored higher on depressive mood scales. The results suggest that pMPFC also implements feedback-related emotional functions, which individually vary depending on depressive moods.

ARTICLE UPDATE - Social Anxiety and Anger Identification: Bubbles Reveal Differential Use of Facial Information With Low Spatial Frequencies.

Langner O, Becker ES, Rinck M.

Psychological Science, in press

We investigated the facial information that socially anxious and nonanxious individuals utilize to judge emotions. Using a reversed-correlation technique, we presented participants with face images that were masked with random bubble patterns. These patterns determined which parts of the face were visible in specific spatial-frequency bands. This masking allowed us to establish which locations and spatial frequencies were helping participants to successfully discriminate angry faces from neutral ones. Although socially anxious individuals performed as well as nonanxious individuals on the emotion-discrimination task, they did not utilize the same facial information for the task. The fine details (high spatial frequencies) around the eyes were discriminative for both groups, but only socially anxious participants additionally processed rough configural information (low spatial frequencies).

ARTICLE UPDATE - Emotion Improves and Impairs Early Vision

Bocanegra BR, Zeelenberg R.

Psychological Science, in press

Recent studies indicate that emotion enhances early vision, but the generality of this finding remains unknown. Do the benefits of emotion extend to all basic aspects of vision, or are they limited in scope? Our results show that the brief presentation of a fearful face, compared with a neutral face, enhances sensitivity for the orientation of subsequently presented low-spatial-frequency stimuli, but diminishes orientation sensitivity for high-spatial-frequency stimuli. This is the first demonstration that emotion not only improves but also impairs low-level vision. The selective low-spatial-frequency benefits are consistent with the idea that emotion enhances magnocellular processing. Additionally, we suggest that the high-spatial-frequency deficits are due to inhibitory interactions between magnocellular and parvocellular pathways. Our results suggest an emotion-induced trade-off in visual processing, rather than a general improvement. This trade-off may benefit perceptual dimensions that are relevant for survival at the expense of those that are less relevant.

ARTICLE UPDATE - Binding and Inhibition in Episodic Memory -Cognitive, Emotional, and Neural Processes.

Bäuml KH, Pastötter B, Hanslmayr S.

Neuroscience and Biobehavioral Reviews, in press

The goal-directed use of human memory requires that irrelevant or unpleasant memories are, at least temporarily, reduced in their accessibility and memory for more relevant or pleasant information is enhanced, thus making memory more efficient. There is evidence that, in memory, inhibitory processes operate to serve this function. Results from three experimental paradigms are reviewed in which the action of intentionally and unintentionally recruited inhibitory processes has been suggested. The findings provide evidence on representational preconditions for the action of inhibitory processes, specifying binding structures in which inhibitory processes may be triggered and binding structures in which inhibitory processes are generally not observed. The findings also provide evidence on how inhibition a ffects memory representations, including changes at the memory unit level and changes in the binding between single units. Finally, current knowledge on the interplay between inhibition and emotion and on possible neural correlates of inhibitory processes is reviewed.

ARTICLE UPDATE - Coarse threat images reveal theta oscillations in the amygdala: A magnetoencephalography study.

Maratos FA, Mogg K, Bradley BP, Rippon G, Senior C.

Cognitive, Affective & Behavioral Neuroscience, 9, 133-143

Neurocognitive models propose a specialized neural system for processing threat-related information, in which the amygdala plays a key role in the analysis of threat cues. fMRI research indicates that the amygdala is sensitive to coarse visual threat relevant information-for example, low spatial frequency (LSF) fearful faces. However, fMRI cannot determine the temporal or spectral characteristics of neural responses. Consequently, we used magnetoencephalography to explore spatiotemporal patterns of activity in the amygdala and cortical regions with blurry (LSF) and normal angry, fearful, and neutral faces. Results demonstrated differences in amygdala activity between LSF threat-related and LSF neutral faces (50-250 msec after face onset). These differences were evident in the theta range (4-8 Hz) and were accompanied by power changes within visual and frontal regions. Our results support the view that the amygdala is involved in the early processing of coarse threat related information and that theta is important in integrating activity within emotion-processing networks.

ARTICLE UPDATE - Do tests of executive functioning predict ability to downregulate emotions spontaneously and when instructed to suppress?

Gyurak A, Goodkind MS, Madan A, Kramer JH, Miller BL, Levenson RW.

Cognitive, Affective & Behavioral Neuroscience, 9, 144-152

Behavioral regulation is a hallmark feature of executive functioning (EF). The present study investigated whether commonly used neuropsychological test measures of EF (i.e., working memory, Stroop, trail making, and verbal fluency) were related to ability to downregulate emotion both spontaneously and when instructed to suppress emotional expressions. To ensure a wide range of EF, 24 frontotemporal lobar degeneration patients, 7 Alzheimer's patients, and 17 neurologically normal controls participated. Participants were exposed to an acoustic startle stimulus (single aversive noise burst) under three conditions: (1) unwarned, (2) warned with no instructions (to measure spontaneous emotion downregulation), and (3) warned with instructions to suppress (to measure instructed emotion downregulation). Results indicated that higher verbal fluency scores were related to greater emotion regulation (operationalized as reduction in body movement and emotional facial behavior when warned of the impending startle) in both regulation conditions. No relationships were found between emotion regulation in these conditions and the other EF measures. We conclude that, of four commonly used measures of EF, verbal fluency best indexes the complex processes of monitoring, evaluation, and control necessary for successful emotion regulation, both spontaneously and following instructions to suppress.

ARTICLE UPDATE - Contingency learning in human fear conditioning involves the ventral striatum.

Klucken T, Tabbert K, Schweckendiek J, Merz CJ, Kagerer S, Vaitl D, Stark R.

Human Brain Mapping, in press

The ability to detect and learn contingencies between fearful stimuli and their predictive cues is an important capacity to cope with the environment. Contingency awareness refers to the ability to verbalize the relationships between conditioned and unconditioned stimuli. Although there is a heated debate about the influence of contingency awareness on conditioned fear responses, neural correlates behind the formation process of contingency awareness have gained only little attention in human fear conditioning. Recent animal studies indicate that the ventral striatum (VS) could be involved in this process, but in human studies the VS is mostly associated with positive emotions. To examine this question, we reanalyzed four recently published classical fear conditioning studies (n = 117) with respect to the VS at three distinct levels of contingency awareness: subjects, who did not learn the contingencies (unaware), subjects, who learned the contingencies during the experiment (learned aware) and subjects, who were informed about the contingencies in advance (instructed aware). The results showed significantly increased activations in the left and right VS in learned aware compared to unaware subjects. Interestingly, this activation pattern was only found in learned but not in instructed aware subjects. We assume that the VS is not involved when contingency awareness does not develop during conditioning or when contingency awareness is unambiguously induced already prior to conditioning. VS involvement seems to be important for the transition from a contingency unaware to a contingency aware state. Implications for fear conditioning models as well as for the contingency awareness debate are discussed. Hum Brain Mapp, 2009.

ARTICLE UPDATE - Unmasking emotion: Exposure duration and emotional engagement.

Codispoti M, Mazzetti M, Bradley MM.

Psychophysiology, in press

Effects of exposure duration on emotional reactivity were investigated in two experiments that parametrically varied the duration of exposure to affective pictures from 25-6000 ms in the presence or absence of a visual mask. Evaluative, facial, autonomic, and cortical responses were measured. Results demonstrated that, in the absence of a visual mask (Experiment 1), emotional content modulated evaluative ratings, cortical, autonomic, and facial changes even with very brief exposures, and there was little evidence that emotional engagement increased with longer exposure. When information persistence was reduced by a visual mask (Experiment 2), differences as a function of hedonic content were absent for all measures when exposure duration was 25 ms but statistically reliable when exposure duration was 80 ms. Between 25-80 ms, individual differences in discriminability were critical in observing affective reactions to masked pictures.

ARTICLE UPDATE - Sleep promotes the neural reorganization of remote emotional memory.

Sterpenich V, Albouy G, Darsaud A, Schmidt C, Vandewalle G, Dang Vu TT, Desseilles M, Phillips C, Degueldre C, Balteau E, Collette F, Luxen A, Maquet P.

The Journal of Neuroscience, 16, 5143-5152

Sleep promotes memory consolidation, a process by which fresh and labile memories are reorganized into stable memories. Emotional memories are usually better remembered than neutral ones, even at long retention delays. In this study, we assessed the influence of sleep during the night after encoding onto the neural correlates of recollection of emotional memories 6 months later. After incidental encoding of emotional and neutral pictures, one-half of the subjects were allowed to sleep, whereas the others were totally sleep deprived, on the first postencoding night. During subsequent retest, functional magnetic resonance imaging sessions taking place 3 d and 6 months later, subjects made recognition memory judgments about the previously studied and new pictures. Between these retest sessions, all participants slept as usual at home. At 6 month retest, recollection was associated with significantly larger responses in subjects allowed to sleep than in sleep-deprived subjects, in the ventral medial prefrontal cortex (vMPFC) and the precuneus, two areas involved in memory retrieval, as well as in the extended amygdala and the occipital cortex, two regions the response of which was modulated by emotion at encoding. Moreover, the functional connectivity was enhanced between the vMPFC and the precuneus, as well as between the extended amygdala, the vMPFC, and the occipital cortex in the sleep group relative to the sleep-deprived group. These results suggest that sleep during the first postencoding night profoundly influences the long-term systems-level consolidation of emotional memory and modifies the functional segregation and integration associated with recollection in the long term.

ARTICLE UPDATE - Reason, emotion and decision-making: risk and reward computation with feeling.

Quartz SR.

Trends in Cognitive Sciences, in press

Many models of judgment and decision-making posit distinct cognitive and emotional contributions to decision-making under uncertainty. Cognitive processes typically involve exact computations according to a cost-benefit calculus, whereas emotional processes typically involve approximate, heuristic processes that deliver rapid evaluations without mental effort. However, it remains largely unknown what specific parameters of uncertain decision the brain encodes, the extent to which these parameters correspond to various decision-making frameworks, and their correspondence to emotional and rational processes. Here, I review research suggesting that emotional processes encode in a precise quantitative manner the basic parameters of financial decision theory, indicating a reorientation of emotional and cognitive contributions to risky choice.

ARTICLE UPDATE - Neural correlates of affective picture processing - a depth ERP study.

Brázdil M, Roman R, Urbánek T, Chládek J, Spok D, Mareček R, Mikl M, Jurák P, Halámek J, Daniel P, Rektor I.

Neuroimage, in press

Using functional neuroimaging techniques (PET and fMRI), various cortical, limbic, and paralimbic structures have been identified in the last decade as neural substrates of human emotion. In this study we used a novel approach (intracerebral recordings of event-related potentials) to add to our knowledge of specific brain regions involved in affective picture processing. Ten intractable epileptic patients undergoing pre-surgical depth electrode recording viewed pleasant, neutral, and unpleasant pictures and intracerebral event-related potentials (ERPs) were recorded. A total of 752 cortical and subcortical sites were investigated. Significant differences in ERPs to unpleasant as compared to neutral or pleasant pictures were frequently and consistently observed in recordings from various brain areas - the mesial temporal cortex (the amygdala, the hippocampus, the temporal pole), the lateral temporal cortex, the mesial prefrontal cortex (ACC and the medial frontal gyrus), and the lateral prefrontal cortex. Interestingly, the mean latencies of responses to emotional stimuli were somewhat shorter in the frontal lobe structures (with evidently earlier activation within lateral prefrontal areas when compared to mesial prefrontal cortex) and longer in the temporal lobe regions. These differences, however, were not significant. Additional clearly positive findings were observed in some rarely investigated regions - in the posterior parietal cortex, the precuneus, and the insula. An approximately equivalent number of positive findings was revealed in the left and right hemisphere structures. These results are in agreement with a multisystem model of human emotion, distributed far beyond the typical limbic system and substantially comprising lateral aspects of both frontal lobes as well.

ARTICLE UPDATE - I'll Know What You're Like When I See How You Feel.

Ames DR, Johar GV.

Psychological Science, in press

Accumulating evidence suggests that targets' displays of emotion shape perceivers' impression of those targets. Prior research has highlighted generalization effects, such as an angry display prompting an impression of hostility. In two studies, we went beyond generalization to examine the interaction of displays and behaviors, finding new evidence of augmenting effects (behavior-correspondent inferences are stronger when behavior is accompanied by positive affect) and discounting effects (such inferences are weaker when behavior is accompanied by negative affect). Thus, the same display can have different effects on impressions depending on the behavior it accompanies. We found evidence that these effects are mediated by ascribed intentions and that they have a boundary: When behaviors and affective displays are repeated, the augmenting and discounting power of displays appears to wane.

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ARTICLE UPDATE - Laterality effect on emotional faces processing: ALE meta-analysis of evidence.

Fusar-Poli P, Placentino A, Carletti F, Allen P, Landi P, Abbamonte M, Barale F, Perez J, McGuire P, Politi PL.

Neuroscience Letters, in press

Recognizing emotion from facial expressions draws on diverse psychological processes implemented in a large array of neural structures. Two major theories of cerebral lateralization of emotional perception have been proposed: (i) the Right-Hemisphere Hypothesis (RHH) and (ii) the Valence-Specific Hypothesis (VSH). To test these lateralization models we conducted a large voxel-based meta-analysis of current functional magnetic resonance imaging (fMRI) studies employing emotional faces paradigms in healthy volunteers. Two independent researchers conducted separate comprehensive PUBMED (1990-May 2008) searches to find all functional magnetic resonance imaging studies using a variant of the emotional faces paradigm in healthy subjects. Out of the 551 originally identified studies, 105 studies met inclusion criteria. The overall database consisted of 1785 brain coordinates which yield an overall sample of 1600 healthy subjects. We found no support for the hypothesis of overall right-lateralization of emotional processing. Conversely, across all emotional conditions the parahippocampal gyrus and amygdala, fusiform gyrus, lingual gyrus, precuneus, inferior and middle occipital gyrus, posterior cingulated, middle temporal gyrus, inferior frontal and superior frontal gyri were activated bilaterally (p=0.001). There was a valence-specific lateralization of brain response during negative emotions processing in the left amygdala (p=0.001). Significant interactions between the approach and avoidance dimensions and prefrontal response were observed (p=0.001).

ARTICLE UPDATE - Neural systems of visual attention responding to emotional gestures.

Flaisch T, Schupp HT, Renner B, Junghöfer M.

Neuroimage, in press

Humans are the only species known to use symbolic gestures for communication. This affords a unique medium for nonverbal emotional communication with a distinct theoretical status compared to facial expressions and other biologically evolved nonverbal emotion signals. While a frown is a frown all around the world, the relation of emotional gestures to their referents is arbitrary and varies from culture to culture. The present studies examined whether such culturally based emotion displays guide visual attention processes. In two experiments, participants passively viewed symbolic hand gestures with positive, negative and neutral emotional meaning. In Experiment 1, functional magnetic resonance imaging (fMRI) measurements showed that gestures of insult and approval enhance activity in selected bilateral visual-associative brain regions devoted to object perception. In Experiment 2, dense sensor event-related brain potential recordings (ERP) revealed that emotional hand gestures are differentially processed already 150 ms poststimulus. Thus, the present studies provide converging neuroscientific evidence that emotional gestures provoke the cardinal signatures of selective visual attention regarding brain structures and temporal dynamics previously shown for emotional face and body expressions. It is concluded that emotionally charged gestures are efficient in shaping selective attention processes already at the level of stimulus perception.

ARTICLE UPDATE - Links between rapid ERP responses to fearful faces and conscious awareness.

Eimer M, Kiss M, Holmes A.

Journal of Neurophysiology, 2, 165-181

To study links between rapid ERP responses to fearful faces and conscious awareness, a backward-masking paradigm was employed where fearful or neutral target faces were presented for different durations and were followed by a neutral face mask. Participants had to report target face expression on each trial. When masked faces were clearly visible (200 ms duration), an early frontal positivity, a later more broadly distributed positivity, and a temporo-occipital negativity were elicited by fearful relative to neutral faces, confirming findings from previous studies with unmasked faces. These emotion-specific effects were also triggered when masked faces were presented for only 17 ms, but only on trials where fearful faces were successfully detected. When masked faces were shown for 50 ms, a smaller but reliable frontal positivity was also elicited by undetected fearful faces. These results demonstrate that early ERP responses to fearful faces are linked to observers' subjective conscious awareness of such faces, as reflected by their perceptual reports. They suggest that frontal brain regions involved in the construction of conscious representations of facial expression are activated at very short latencies.

ARTICLE UPDATE - Segregated and integrated coding of reward and punishment in the cingulate cortex.

Fujiwara J, Tobler PN, Taira M, Iijima T, Tsutsui KI.

Journal of Neurophysiology, in press

Affective stimuli fall into two major classes, reward and punishment, both of which are processed by the cingulate cortex. However, it is unclear whether the positive and negative affective values of monetary reward and punishment are processed by separate or common subregions of the cingulate cortex. We performed a functional magnetic resonance imaging (fMRI) study using a free-choice task, and compared cingulate activations for different levels of monetary gain and loss. Gain-specific activation (increasing activation for increasing gain, but no activation change in relation to loss) occurred mainly in the anterior part of the anterior cingulate and in the posterior cingulate cortex. Conversely, loss-specific activation (increasing activation for increasing loss, but no activation change in relation to gain) occurred in-between these areas, in the middle and posterior part of the anterior cingulate. Integrated coding of gain and loss (increasing activation throughout the full range, from biggest loss to biggest gain) occurred in the dorsal part of the anterior cingulate, at the border with medial prefrontal cortex. Finally, unspecific activation increases to both gains and losses (increasing activation to increasing gains and increasing losses, possibly reflecting attention) occurred in dorsal and middle regions of the cingulate cortex. Together, these results suggest separate and common coding of monetary reward and punishment in distinct subregions of the cingulate cortex. Further meta-analysis suggested that the presently found reward- and punishment-specific areas overlapped with those processing positive and negative emotions, respectively.

ARTICLE UPDATE - Between- and within-ear congruency and laterality effects in an auditory semantic/emotional prosody conflict task.

Techentin C, Voyer D, Klein RM.

Brain and Cognition, in press

The present study investigated the influence of within- and between-ear congruency on interference and laterality effects in an auditory semantic/prosodic conflict task. Participants were presented dichotically with words (e.g., mad, sad, glad) pronounced in either congruent or incongruent emotional tones (e.g., angry, happy, or sad) and identified a target word or emotion under one of two conditions. In the within-ear condition, the congruent or incongruent dimensions were bound within a single stimulus and therefore, presented to the same ear. In the between-ear condition, the two dimensions were split between two stimuli and, therefore, presented in separate ears. Findings indicated interference in both conditions. However, the expected right ear advantage (EA) for words and left EA for emotions were obtained only in the between-ear condition. Factors involved in producing interference and laterality effects in dichotic listening tasks are discussed.

ARTICLE UPDATE - Enhanced post-learning memory consolidation is influenced by arousal predisposition and emotion regulation but not by stimulus valenc

Nielson KA, Lorber W.

Neurobiology of Learning & Memory, in press

Emotionally arousing stimuli are more memorable than neutral ones and arousal induced after learning enhances later retrieval. However, there is as yet little study of how stimulus qualities might interact with induced arousal and how individual differences might influence the modulation of memory. Thus, the present study examined the effect of arousal induced after learning on memory for words that varied in both arousal and valence quality, as well as the influence of three individual differences factors that are known to influence arousal response: emotional suppression, emotional reappraisal, and arousal predisposition. Seventy-six adults (57 female) viewed and rated 60 words that normatively ranged from high to low in arousal and valence. Ten minutes later, they viewed a 3-min comedic or neutral video clip. Arousal induced after learning enhanced one-week delayed memory, spanning the lengthy task without preference for word type or serial position, contrasting with reports of arousal effects interacting with stimulus qualities. Importantly, being predisposed to arousal led to greater enhancement of long-term memory modulation, while the use of emotional reappraisal, which reduces arousal responding, inhibited the ability of arousal to induce memory enhancement. Thus, individual differences that influence arousal responding can contribute to or interfere with memory modulation.

ARTICLE UPDATE - How do emotion and motivation direct executive control?

Pessoa L.

Trends in Cognitive Science, in press

Emotion and motivation have crucial roles in determining human behavior. Yet, how they interact with cognitive control functions is less understood. Here, the basic elements of a conceptual framework for understanding how they interact are introduced. More broadly, the 'dual competition' framework proposes that emotion and motivation affect both perceptual and executive competition. In particular, the anterior cingulate cortex is hypothesized to be engaged in attentional/effortful control mechanisms and to interact with several other brain structures, including the amygdala and nucleus accumbens, in integrating affectively significant signals with control signals in prefrontal cortex. An implication of the proposal is that emotion and motivation can either enhance or impair behavioral performance depending on how they interact with control functions.

ARTICLE UPDATE - Nonautomatic emotion perception in a dual-task situation.

Tomasik D, Ruthruff E, Allen PA, Lien MC.

Psychological Bulletin & Review, 16, 282-288

Are emotions perceived automatically? Two psychological refractory period experiments were conducted to ascertain whether emotion perception requires central attentional resources. Task 1 required an auditory discrimination (tone vs. noise), whereas Task 2 required a discrimination between happy and angry faces. The difficulty of Task 2 was manipulated by varying the degree of emotional expression. The stimulus onset asynchrony (SOA) between Task 1 and Task 2 was also varied. Experiment 1 revealed additive effects of SOA and Task 2 emotion-perception difficulty. Experiment 2 replicated the additive relationship with a stronger manipulation of emotion-perception difficulty. According to locus-of-slack logic, our participants did not process emotional expressions while central resources were devoted to Task 1. We conclude that emotion perception is not fully automatic.

ARTICLE UPDATE - Emotion perception in emotionless face images suggests a norm-based representation.

Neth D, Martinez AM.

Journal of Vision, 9, 5.1 - 5.11

Perception of facial expressions of emotion is generally assumed to correspond to underlying muscle movement. However, it is often observed that some individuals have sadder or angrier faces, even for neutral, motionless faces. Here, we report on one such effect caused by simple static configural changes. In particular, we show four variations in the relative vertical position of the nose, mouth, eyes, and eyebrows that affect the perception of emotion in neutral faces. The first two configurations make the vertical distance between the eyes and mouth shorter than average, resulting in the perception of an angrier face. The other two configurations make this distance larger than average, resulting in the perception of sadness. These perceptions increase with the amount of configural change, suggesting a representation based on variations from a norm (prototypical) face.

ARTICLE UPDATE - Flawless visual short-term memory for facial emotional expressions.

Bankó EM, Gál V, Vidnyánszky Z.

Journal of Vision, 9, 12.1-12.13

Facial emotions are important cues of human social interactions. Emotional expressions are continuously changing and thus should be monitored, memorized, and compared from time to time during social intercourse. However, it is not known how efficiently emotional expressions can be stored in short-term memory. Here we show that emotion discrimination is not impaired when the faces to be compared are separated by several seconds, requiring storage of fine-grained emotion-related information in short-term memory. Likewise, we found no significant effect of increasing the delay between the sample and the test face in the case of facial identity discrimination. Furthermore, a second experiment conducted on a large subject sample (N = 160) revealed flawless short-term memory for both facial emotions and facial identity also when observers performed the discrimination tasks only twice with novel faces. We also performed an fMRI experiment, which confirmed that discrimination of fine-grained emotional expressions in our experimental paradigm involved processing of high-level facial emotional attributes. Significantly stronger fMRI responses were found in a cortical network--including the posterior superior temporal sulcus--that is known to be involved in processing of facial emotional expression during emotion discrimination than during identity discrimination. These findings reveal flawless, high-resolution visual short-term memory for emotional expressions, which might underlie efficient monitoring of continuously changing facial emotions.

ARTICLE UPDATE - The role of mirror neurons in processing vocal emotions: evidence from psychophysiological data.

Ramachandra V, Depalma N, Lisiewski S.

International Journal of Neuroscience, 119,681-690

Recent evidence suggests that the mirror neuron system may serve as a common neural substrate for processing motor, linguistic, emotional, and other higher-level cognitive information. The current study employed psychophysiological methods to elucidate the role of this system in processing vocal emotions. Skin conductance and heart rate were measured for 25 undergraduate students while they were both listening to emotional vocalizations and also thinking (internal production) about them. The results revealed changes in skin conductance response and heart rate during both "listening" and "thinking" conditions. This suggests an active role of the mirror neuron system in processing vocal emotions.

ARTICLE UPDATE - Specific and common brain regions involved in the perception of faces and bodies and the representation of their emotional expression

van de Riet WA, Grezes J, de Gelder B.

Social Neuroscience, 4, 101 - 120

Many studies provide support for the role of the fusiform gyrus in face recognition and its sensitivity to emotional expressions. Recently, category-specific representation was also observed for neutral human bodies in the middle temporal/middle occipital gyrus (extrastriate body area) but it is not clear whether this area is also sensitive to emotional bodily expressions. Besides these areas, other regions that process the affective information carried by the face and the body may be common and/or specific to the face or the body. To clarify these issues we performed a systematic comparison of how the whole brain processes faces and bodies and how their affective information is represented. Participants categorized emotional facial and bodily expressions while brain activity was measured using functional magnetic resonance imaging. Our results show that, first, the amygdala and the fusiform gyrus are sensitive to recognition of facial and bodily fear signals. Secondly, the extrastriate body area-area V5/MT is specifically involved in processing bodies without being sensitive to the emotion displayed. Thirdly, other important areas such as the superior temporal sulcus, the parietal lobe and subcortical structures represent selectively facial and bodily expressions. Finally, some face/body differences in activation are a function of the emotion expressed.

ARTICLE UPDATE - Effects of anxiety and attention control on processing pictorial and linguistic emotional information.

Reinholdt-Dunne ML, Mogg K, Bradley BP.

Behavioral Research & Therapy, in press

This study investigated the role of executive attention control in modulating selective processing of emotional information in anxiety. It was hypothesized that the combination of high anxiety and poor attention control would be associated with greater difficulty in ignoring task-irrelevant threat-related information. The study included both faces and words as stimuli. Cognitive interference effects were assessed using two emotional Stroop tasks: one with angry, fearful, happy and neutral faces, and one with threat-related, positive, and neutral words. An objective measure of attention control was obtained from the Attention network task. There were four participant groups with high/low trait anxiety and high/low attention control. Results indicated that the combination of high anxiety and poor attention control was associated with greater cognitive interference by emotional faces (including angry faces), compared to neutral faces. This interference effect was not evident in participants with high anxiety and high attentional control, or in low-anxious individuals. There was no evidence of associations between anxiety, attention control, and the interference effect of emotional words. Results indicate that high anxiety and poor attention control together predict enhanced processing of emotionally salient information, such as angry facial expressions. Implications for models of emotion processing are discussed.

ARTICLE UPDATE - Time course and task dependence of emotion effects in word processing.

Schacht A, Sommer W.

Cognitive, Affective and Behavioral Neuroscience, 8, 28-43

The emotional content of stimuli influences cognitive performance. In two experiments, we investigated the time course and mechanisms of emotional influences on visual word processing in various tasks by recording event-related brain potentials (ERPs). The stimuli were verbs of positive, negative, and neutral valence. In Experiment 1, where lexical decisions had to be performed on single verbs, both positive and negative verbs were processed more quickly than neutral verbs and elicited a distinct ERP component, starting around 370 msec. In Experiment 2, the verbs were embedded in a semantic context provided by single nouns. Likewise, structural, lexical, and semantic decisions for positive verbs were accelerated, and an ERP effect with a scalp distribution comparable to that in Experiment 1 now started about 200 msec earlier. These effects may signal an automatic allocation of attentional resources to emotionally arousing words, since they were not modulated by different task demands. In contrast, a later ERP effect of emotion was restricted to lexical and semantic decisions and, thus, appears to indicate more elaborated, task-dependent processing of emotional words.

ARTICLE UPDATE - Noradrenergic enhancement of amygdala responses to fear.

Onur OA, Walter H, Schlaepfer TE, Rehme AK, Schmidt C, Keysers C, Maier W, Hurlemann R.

Social Cognitive & Affective Neuroscience, in press

Multiple lines of evidence implicate the basolateral amygdala (BLA) and the noradrenergic (norepinephrine, NE) system in responding to stressful stimuli such as fear signals, suggesting hyperfunction of both in the development of stress-related pathologies including anxiety disorders. However, no causative link between elevated NE neurotransmission and BLA hyperresponsiveness to fear signals has been established to date in humans. To determine whether or not increased noradrenergic tone enhances BLA responses to fear signals, we used functional magnetic resonance imaging (fMRI) and a strategy of pharmacologically potentiating NE neurotransmission in healthy volunteers. 18 subjects were scanned two times on a facial emotion paradigm and given either a single-dose placebo or 4 mg of the selective NE reuptake inhibitor reboxetine 2 h prior to an fMRI session. We found that reboxetine induced an amygdala response bias towards fear signals that did not exist at placebo baseline. This pharmacological effect was probabilistically mapped to the BLA. Extrapolation of our data to conditions of traumatic stress suggests that disinhibited endogenous NE signaling could serve as a crucial etiological contributor to post-traumatic stress disorder (PTSD) by eliciting exaggerated BLA responses to fear signals.

ARTICLE UPDATE - The set switching function of nonclinical dissociators under negative emotion.

Chiu CD, Yeh YY, Huang YM, Wu YC, Chiu YC.

Journal of Abnormal Psychology, 118, 214-222

Rapid switching may underlie the disruption of some integrated thought processes that characterize dissociation in both nonclinical and clinical populations. We investigated the set switching function under negative emotion with three groups of nonclinical participants that had different degrees of dissociation proneness. In the experiment, participants judged whether the digit in a predefined target color was odd or even on the preswitch trials. In a perseverance condition, participants were required to switch to a new target color while the previous target color became the distractor color. In a learned irrelevance condition, the previously ignored color became the new target color. The results showed that the three groups did not differ in focusing attention in the preswitch trials, for set switching in the baseline condition (in which emotion was not engaged), or for switching in the learned irrelevance condition under negative emotion. However, high dissociators under negative emotion showed faster switching in the perseverance condition. This enhanced ability to divert attention to a new mental set under negative emotion may be a coping strategy related to cognitive symptoms in dissociative disorders.

ARTICLE UPDATE - Influence of intermixed emotion-relevant trials on the affective Simon effect.

Zhang Y, Proctor RW.

Experimental Psychology, 55, 409-416

"Good" and "bad" vocal responses are faster when an irrelevant emotional stimulus feature corresponds with the response than when it does not, a phenomenon known as the affective Simon effect. Two experiments investigated how this effect was influenced by an intermixed emotion-relevant evaluation task. In Experiment 1, four schematic faces (friendly, happy, hostile, sad) were used for the affective Simon task and four different images (bird, heart, gun, ghost) for the evaluation task, whereas in Experiment 2 the schematic faces were used for both tasks. Mixed-compatible emotion-relevant trials increased the affective Simon effect in both experiments, but mixed-incompatible emotion-relevant trials did not influence it. Also, the advantage of the compatible mapping over the incompatible mapping increased in mixed conditions rather than decreased. These results differ from those obtained when visual-manual tasks for which location is relevant and irrelevant are mixed. They confirm that enhancement of the affective Simon effect when the Simon task is mixed with a compatible emotion-relevant task is due to increased salience of the affective valence.

ARTICLE UPDATE - Motivated and controlled attention to emotion: Time-course of the late positive potential.

Hajcak G, Dunning JP, Foti D.

Clinical Neurophysiology, in press

OBJECTIVE: The present study examined the time-course of automatic and controlled modulation of the late positive potential (LPP) during emotional picture viewing. METHODS: Participants (N=32) viewed neutral and unpleasant stimuli for 6000ms; at 3000ms, one of two tones signaled participants to attend either to a more or less arousing portion of the picture. The time-course of the LPP was examined both during the passive viewing and directed attention portions of the trial using the method proposed by Guthrie and Buchwald [Guthrie D, Buchwald JS. Significance testing of difference potentials. Psychophysiology 1991;28(2):240-4]. RESULTS: During passive viewing, the LPP became reliably larger following the presentation of unpleasant pictures from 160ms onward; the magnitude of the LPP became reliably smaller beginning 620ms after participants were instructed to attend to the less arousing aspects of unpleasant pictures - and this difference was maintained throughout the duration of the trial. CONCLUSIONS: The LPP reflects relatively automatic attention to emotional visual stimuli, but is also sensitive to manipulations of directed attention toward arousing versus neutral aspects of such stimuli. SIGNIFICANCE: These results shed further light on the time-course of emotional and cognitive modulation of the LPP, and suggest that the LPP reflects the relatively rapid and dynamic allocation of increased attention to emotional stimuli.

ARTICLE UPDATE - Mood influences supraspinal pain processing separately from attention.

Villemure C, Bushnell MC.

Journal of Neuroscience, 29, 705 - 715

Studies show that inducing a positive mood or diverting attention from pain decreases pain perception. Nevertheless, induction manipulations, such as viewing interesting movies or performing mathematical tasks, often influence both emotional and attentional states. Imaging studies have examined the neural basis of psychological pain modulation, but none has explicitly separated the effects of emotion and attention. Using odors to modulate mood and shift attention from pain, we previously showed that the perceptual consequences of changing mood differed from those of altering attention, with mood primarily altering pain unpleasantness and attention preferentially altering pain intensity. These findings suggest that brain circuits involved in pain modulation provoked by mood or attention are partially separable. Here we used functional magnetic resonance imaging to directly compare the neurocircuitry involved in mood- and attention-related pain modulation. We manipulated independently mood state and attention direction, using tasks involving heat pain and pleasant and unpleasant odors. Pleasant odors, independent of attentional focus, induced positive mood changes and decreased pain unpleasantness and pain-related activity within the anterior cingulate (ACC), medial thalamus, and primary and secondary somatosensory cortices. The effects of attentional state were less robust, with only the activity in anterior insular cortex (aIC) showing possible attentional modulation. Lateral inferior frontal cortex [LinfF; Brodmann's area (BA) 45/47] activity correlated with mood-related modulation, whereas superior posterior parietal (SPP; BA7) and entorhinal activity correlated with attention-related modulation. ACC activity covaried with LinfF and periacqueductal gray activity, whereas aIC activity covaried with SPP activity. These findings suggest that separate neuromodulatory circuits underlie emotional and attentional modulation of pain.

ARTICLE UPDATE - Attentional control of emotional distraction in rapid serial visual presentation.

Peers PV, Lawrence AD.

Emotion, 9, 140 - 145

Temperament research has highlighted the importance of attentional control in both emotion regulation and as a predictor of psychopathology. Enhanced susceptibility to emotional distraction is a key feature of mood disturbance. Whereas many studies have examined the influence of individual differences in anxiety on the disruptive effects of emotional distractors, individual differences in attentional control have been largely neglected. Here we examine, within healthy volunteers, the relative contributions of individual differences in self-reported anxiety and attentional control to distractibility caused by emotional or neutral faces distractors occurring prior to neutral face targets during rapid serial visual presentation. Participants with good attentional control were less affected by both neutral and emotional distractors than participants with poorer attentional control. More pronounced distraction deficits were seen for emotional relative to neutral distractors in individuals with poor attentional control. In contrast state anxiety was not associated with increased emotional distraction. Our findings suggest a protective role of attentional control mechanisms in minimizing the influence of emotional distraction.

ARTICLE UPDATE - The effects of emotional intensity on ERP correlates of recognition memory

Schaefer A, Fletcher K, Pottage CL, Alexander K, Brown C.

Neuroreport, 20, 319 - 324

The effects of negative emotional intensity on memory-related brain activity were tested by using human scalp event-related potentials (ERP). A neural index of memory function - the electrophysiological 'Old-New' effect - was obtained from participants undertaking a memory recognition test of previously studied ('old') and unstudied ('new') pictures of variable levels of negative emotional intensity. The magnitude of the old-new effect was compared across four different levels of linearly increasing stimulus emotional intensity. Results revealed an inverted-U-shaped effect of emotional intensity on the magnitude of ERP old-new differences starting at 300 ms after stimulus onset. These results suggest that moderate negative emotions can enhance memory brain function, whereas extreme levels of emotional intensity have the potential of inhibiting memory function. Results are discussed in terms of their implications for neurobiological and psychological models of emotion-memory interactions.

ARTICLE UPDATE- Orienting to threat: faster localization of fearful facial expressions and body postures revealed by saccadic eye movements.

Bannerman RL, Milders M, de Gelder B, Sahraie A.

Proceedings in Biological Science, in press

Most studies investigating speeded orientation towards threat have used manual responses. By measuring orienting behaviour using eye movements a more direct and ecologically valid measure of attention can be made. Here, we used a forced-choice saccadic and manual localization task to investigate the speed of discrimination for fearful and neutral body and face images. Fearful/neutral body or face pairs were bilaterally presented for either 20 or 500ms. Results showed faster saccadic orienting to fearful body and face emotions compared with neutral only at the shortest presentation time (20ms). For manual responses, faster discrimination of fearful bodies and faces was observed only at the longest duration (500ms). More errors were made when localizing neutral targets, suggesting that fearful bodies and faces may have captured attention automatically. Results were not attributable to low-level image properties as no threat bias, in terms of reaction time or accuracy, was observed for inverted presentation. Taken together, the results suggest faster localization of threat conveyed both by the face and the body within the oculomotor system. In addition, enhanced detection of fearful body postures suggests that we can readily recognize threat-related information conveyed by body postures in the absence of any face cues.

ARTICLE UPDATE - Dissociable neural effects of stimulus valence and preceding context during the inhibition of responses to emotional faces.

Schulz KP, Clerkin SM, Halperin JM, Newcorn JH, Tang CY, Fan J.

Human Brain Mapping, in press

Socially appropriate behavior requires the concurrent inhibition of actions that are inappropriate in the context. This self-regulatory function requires an interaction of inhibitory and emotional processes that recruits brain regions beyond those engaged by either processes alone. In this study, we isolated brain activity associated with response inhibition and emotional processing in 24 healthy adults using event-related functional magnetic resonance imaging (fMRI) and a go/no-go task that independently manipulated the context preceding no-go trials (ie, number of go trials) and the valence (ie, happy, sad, and neutral) of the face stimuli used as trial cues. Parallel quadratic trends were seen in correct inhibitions on no-go trials preceded by increasing numbers of go trials and associated activation for correct no-go trials in inferior frontal gyrus pars opercularis, pars triangularis, and pars orbitalis, temporoparietal junction, superior parietal lobule, and temporal sensory association cortices. Conversely, the comparison of happy versus neutral faces and sad versus neutral faces revealed valence-dependent activation in the amygdala, anterior insula cortex, and posterior midcingulate cortex. Further, an interaction between inhibition and emotion was seen in valence-dependent variations in the quadratic trend in no-go activation in the right inferior frontal gyrus and left posterior insula cortex. These results suggest that the inhibition of response to emotional cues involves the interaction of partly dissociable limbic and frontoparietal networks that encode emotional cues and use these cues to exert inhibitory control over the motor, attention, and sensory functions needed to perform the task, respectively.

ARTICLE UPDATE - Emotions in word and face processing: Early and late cortical responses.

Emotions in word and face processing: Early and late cortical responses.

Brain & Cognition, in press

Recent research suggests that emotion effects in word processing resemble those in other stimulus domains such as pictures or faces. The present study aims to provide more direct evidence for this notion by comparing emotion effects in word and face processing in a within-subject design. Event-related brain potentials (ERPs) were recorded as participants made decisions on the lexicality of emotionally positive, negative, and neutral German verbs or pseudowords, and on the integrity of intact happy, angry, and neutral faces or slightly distorted faces. Relative to neutral and negative stimuli both positive verbs and happy faces elicited posterior ERP negativities that were indistinguishable in scalp distribution and resembled the early posterior negativities reported by others. Importantly, these ERP modulations appeared at very different latencies. Therefore, it appears that similar brain systems reflect the decoding of both biological and symbolic emotional signals of positive valence, differing mainly in the speed of meaning access, which is more direct and faster for facial expressions than for words.

ARTICLE UPDATE - Functional connectivity of the human amygdala using resting state fMRI.

Roy AK, Shehzad Z, Margulies DS, Kelly AM, Uddin LQ, Gotimer K, Biswal BB, Castellanos FX, Milham MP.

Neuroimage, in press

The amygdala is composed of structurally and functionally distinct nuclei that contribute to the processing of emotion through interactions with other subcortical and cortical structures. While these circuits have been studied extensively in animals, human neuroimaging investigations of amygdala-based networks have typically considered the amygdala as a single structure, which likely masks contributions of individual amygdala subdivisions. The present study uses resting state functional magnetic resonance imaging (fMRI) to test whether distinct functional connectivity patterns, like those observed in animal studies, can be detected across three amygdala subdivisions: laterobasal, centromedial, and superficial. In a sample of 65 healthy adults, voxelwise regression analyses demonstrated positively-predicted ventral and negatively-predicted dorsal networks associated with the total amygdala, consistent with previous animal and human studies. Investigation of individual amygdala subdivisions revealed distinct differences in connectivity patterns within the amygdala and throughout the brain. Spontaneous activity in the laterobasal subdivision predicted activity in temporal and frontal regions, while activity in the centromedial nuclei predicted activity primarily in striatum. Activity in the superficial subdivision positively predicted activity throughout the limbic lobe. These findings suggest that resting state fMRI can be used to investigate human amygdala networks at a greater level of detail than previously appreciated, allowing for the further advancement of translational models.

ARTICLE UPDATE - Dissociable processes underlying decisions in the Iowa Gambling Task: A new integrative framework.

Stocco A, Fum D, Napoli A.

Behavioral and Brain Functions, in press

ABSTRACT: BACKGROUND: The Iowa Gambling Task (IGT) is a common paradigm used to study the interactions between emotions and decision making, yet little consensus exists on the cognitive process determining participants' decisions, what affects them, and how these processes interact with each other. A novel conceptual framework is proposed according to which behavior in the IGT reflects a balance between two dissociable processes; a cognitively demanding process that tracks each option's long-term payoff, and a lower-level, automatic process that is primarily sensitive to loss frequency and magnitude. METHODS: A behavioral experiment was carried out with a modified version of IGT. In this modified version, participants went through an additional phase of interaction, designed to measure performance without further learning, in which no feedback on individual decisions was given. A secondary distractor task was presented in either the first or the second phase of the experiment. Behavioral measures of performance tracking both payoff and frequency sensitivity in choices were collected throughout the experiment. RESULTS: Consistent with our framework, the results confirmed that: (a) the two competing cognitive processes can be dissociated; (b) that learning from decision outcomes requires central cognitive resources to estimate long-term payoff; and (c) that the decision phase itself can be carried out during an interfering task once learning has occurred. CONCLUSIONS: The experimental results support our novel description of the cognitive processes underlying performance in the Iowa Gambling Task. They also suggest that patients' impairments in this and other gambling paradigms can originate from a number of different causes, including a failure in allocating resources among cognitive strategies. This latter interpretation might be particularly useful in explaining the impairments of patients with ventromedial prefrontal cortex lesions and, by extension, the contribution of this brain region to human decision making.

ARTICLE UPDATE - The emotional blink: Adult age differences in visual attention to emotional information.

Langley LK, Rokke PD, Stark AC, Saville AL, Allen JL, Bagne AG.

Psycholoyg & Aging, 23, 873-885

To assess age differences in attention-emotion interactions, the authors asked young adults (ages 18-33 years) and older adults (ages 60-80 years) to identify target words in a rapid serial visual presentation (RSVP) task. The second of two target words was neutral or emotional in content (positive in Experiment 1, negative in Experiment 2). In general, the ability to identify targets from a word stream declined with age. Age differences specific to the attentional blink were greatly reduced when baseline detection accuracy was equated between groups. With regard to emotion effects, older adults showed enhanced identification of both positive and negative words relative to neutral words, whereas young adults showed enhanced identification of positive words and reduced identification of negative words. Together these findings suggest that the nature of attention-emotion interactions changes with age, but there was little support for a motivational shift consistent with emotional regulation goals at an early stage of cognitive processing.

ARTICLE UPDATE - Behavioral triggers of skin conductance responses and their neural correlates in the primate amygdala.

Laine CM, Spitler KM, Mosher CP, Gothard KM.

Journal of Neurophysiology, in press

The amygdala plays a crucial role in evaluating the emotional significance of stimuli and in transforming the results of this evaluation into appropriate autonomic responses. Lesion and stimulation studies suggest involvement of the amygdala in the generation of the skin conductance response (SCR), which is an indirect measure of autonomic activity that has been associated with both emotion and attention. It is unclear if this involvement marks an emotional reaction to an external stimulus, or sympathetic arousal regardless of its origin. We recorded skin conductance in parallel with single unit activity from the right amygdala of two rhesus monkeys during a rewarded image viewing task, and while the monkeys sat alone in a dimly lit room, drifting in and out of sleep. In both experimental conditions, we found similar SCR-related modulation of activity at the single unit and population level. This suggests that the amygdala contributes to the production or modulation of SCRs regardless of the source of sympathetic arousal.

ARTICLE UPDATE - Decoding face information in time, frequency and space from direct intracranial recordings of the human brain.

Tsuchiya N, Kawasaki H, Oya H, Howard MA 3rd, Adolphs R.

PLoS One, in press

Faces are processed by a neural system with distributed anatomical components, but the roles of these components remain unclear. A dominant theory of face perception postulates independent representations of invariant aspects of faces (e.g., identity) in ventral temporal cortex including the fusiform gyrus, and changeable aspects of faces (e.g., emotion) in lateral temporal cortex including the superior temporal sulcus. Here we recorded neuronal activity directly from the cortical surface in 9 neurosurgical subjects undergoing epilepsy monitoring while they viewed static and dynamic facial expressions. Applying novel decoding analyses to the power spectrogram of electrocorticograms (ECoG) from over 100 contacts in ventral and lateral temporal cortex, we found better representation of both invariant and changeable aspects of faces in ventral than lateral temporal cortex. Critical information for discriminating faces from geometric patterns was carried by power modulations between 50 to 150 Hz. For both static and dynamic face stimuli, we obtained a higher decoding performance in ventral than lateral temporal cortex. For discriminating fearful from happy expressions, critical information was carried by power modulation between 60-150 Hz and below 30 Hz, and again better decoded in ventral than lateral temporal cortex. Task-relevant attention improved decoding accuracy more than 10% across a wide frequency range in ventral but not at all in lateral temporal cortex. Spatial searchlight decoding showed that decoding performance was highest around the middle fusiform gyrus. Finally, we found that the right hemisphere, in general, showed superior decoding to the left hemisphere. Taken together, our results challenge the dominant model for independent face representation of invariant and changeable aspects: information about both face attributes was better decoded from a single region in the middle fusiform gyrus.

ARTICLE UPDATE - EEG-MEG evidence for early differential repetition effects for fearful, happy and neutral faces.

Morel S, Ponz A, Mercier M, Vuilleumier P, George N.

Brain Research, in press

To determine how emotional information modulates subsequent traces for repeated stimuli, we combined simultaneous electro-encephalography (EEG) and magneto-encephalography (MEG) measures during long-lag incidental repetition of fearful, happy, and neutral faces. Repetition effects were modulated by facial expression in three different time windows, starting as early as 40-50 ms in both EEG and MEG, then arising at the time of the N170/M170, and finally between 280-320 ms in MEG only. The very early repetition effect, observed at 40-50 ms over occipito-temporo-parietal regions, showed a different MEG topography according to the facial expression. This differential response to fearful, happy and neutral faces suggests the existence of very early discriminative visual processing of expressive faces, possibly based on the low-level physical features typical of different emotions. The N170 and M170 face-selective components both showed repetition enhancement selective to neutral faces, with greater amplitude for emotional than neutral faces on the first but not the second presentation. These differential repetition effects may reflect valence acquisition for the neutral faces due to repetition, and suggest a combined influence of emotion- and experience-related factors on the early stage of face encoding. Finally, later repetition effects consisted in enhanced M300 (MEG) between 280 and 320 ms for fearful relative to happy and neutral faces that occurred on the first presentation, but levelled out on the second presentation. This effect may correspond to the higher arousing value of fearful stimuli that might habituate with repetition. Our results reveal that multiple stages of face processing are affected by the repetition of emotional information.

ARTICLE UPDATE - Dissociable neural effects of stimulus valence and preceding context during the inhibition of responses to emotional faces.

Schulz KP, Clerkin SM, Halperin JM, Newcorn JH, Tang CY, Fan J.

Human Brain Mapping, in press

Socially appropriate behavior requires the concurrent inhibition of actions that are inappropriate in the context. This self-regulatory function requires an interaction of inhibitory and emotional processes that recruits brain regions beyond those engaged by either processes alone. In this study, we isolated brain activity associated with response inhibition and emotional processing in 24 healthy adults using event-related functional magnetic resonance imaging (fMRI) and a go/no-go task that independently manipulated the context preceding no-go trials (ie, number of go trials) and the valence (ie, happy, sad, and neutral) of the face stimuli used as trial cues. Parallel quadratic trends were seen in correct inhibitions on no-go trials preceded by increasing numbers of go trials and associated activation for correct no-go trials in inferior frontal gyrus pars opercularis, pars triangularis, and pars orbitalis, temporoparietal junction, superior parietal lobule, and temporal sensory association cortices. Conversely, the comparison of happy versus neutral faces and sad versus neutral faces revealed valence-dependent activation in the amygdala, anterior insula cortex, and posterior midcingulate cortex. Further, an interaction between inhibition and emotion was seen in valence-dependent variations in the quadratic trend in no-go activation in the right inferior frontal gyrus and left posterior insula cortex. These results suggest that the inhibition of response to emotional cues involves the interaction of partly dissociable limbic and frontoparietal networks that encode emotional cues and use these cues to exert inhibitory control over the motor, attention, and sensory functions needed to perform the task, respectively.

ARTICLE UPDATE - Working memory capacity and the self-regulation of emotional expression and experience.

Schmeichel BJ, Volokhov RN, Demaree HA.

Journal of Personality and Social Psychology, 95, 1526-1540

This research examined the relationship between individual differences in working memory capacity and the self-regulation of emotional expression and emotional experience. Four studies revealed that people higher in working memory capacity suppressed expressions of negative emotion (Study 1) and positive emotion (Study 2) better than did people lower in working memory capacity. Furthermore, compared to people lower in working memory capacity, people higher in capacity more capably appraised emotional stimuli in an unemotional manner and thereby experienced (Studies 3 and 4) and expressed (Study 4) less emotion in response to those stimuli. These findings indicate that cognitive ability contributes to the control of emotional responding.

ARTICLE UPDATE - Emotions in Go/NoGo conflicts.

Schacht A, Nigbur R, Sommer W.

Psychological Research, in press

On the basis of current emotion theories and functional and neurophysiological ties between the processing of conflicts and errors on the one hand and errors and emotions on the other hand we predicted that conflicts between prepotent Go responses and occasional NoGo trials in the Go/NoGo task would induce emotions. Skin conductance responses (SCRs), corrugator muscle activity, and startle blink responses were measured in three experiments requiring speeded Go responses intermixed with NoGo trials of different relative probability and in a choice reaction experiment serving as a control. NoGo trials affected several of these emotion-sensitive indicators as SCRs and startle blinks were reduced whereas corrugator activity was prolonged as compared to Go trials. From the pattern of findings we suggest that NoGo conflicts are not aversive. Instead, they appear to be appraised as obstructive for the response goal and as less action relevant than Go trials.

ARTICLE UPDATE - Visual Awareness, Emotion, and Gamma Band Synchronization.

Luo Q, Mitchell D, Cheng X, Mondillo K, McCaffrey D, Holroyd T, Carver F, Coppola R, Blair J.

Cerebral Cortex, in press

What makes us become aware? A popular hypothesis is that if cortical neurons fire in synchrony at a certain frequency band (gamma), we become aware of what they are representing. We tested this hypothesis adopting brain-imaging techniques with good spatiotemporal resolution and frequency-specific information. Specifically, we examined the degree to which increases in event-related synchronization (ERS) in the gamma band were associated with awareness of a stimulus (its detectability) and/or the emotional content of the stimulus. We observed increases in gamma band ERS within prefrontal-anterior cingulate, visual, parietal, posterior cingulate, and superior temporal cortices to stimuli available to conscious awareness. However, we also observed increases in gamma band ERS within the amygdala, visual, prefrontal, parietal, and posterior cingulate cortices to emotional relative to neutral stimuli, irrespective of their availability to conscious access. This suggests that increased gamma band ERS is related to, but not sufficient for, consciousness.

ARTICLE UPDATE - Attentional selectivity for emotional faces: Evidence from human electrophysiology.

Holmes A, Bradley BP, Kragh Nielsen M, Mogg K.

Psychophysiology, in press

Abstract This study investigated the temporal course of attentional biases for threat-related (angry) and positive (happy) facial expressions. Electrophysiological (event-related potential) and behavioral (reaction time [RT]) data were recorded while participants viewed pairs of faces (e.g., angry face paired with neutral face) shown for 500 ms and followed by a probe. Behavioral results indicated that RTs were faster to probes replacing emotional versus neutral faces, consistent with an attentional bias for emotional information. Electrophysiological results revealed that attentional orienting to threatening faces emerged earlier (early N2pc time window; 180-250 ms) than orienting to positive faces (after 250 ms), and that attention was sustained toward emotional faces during the 250-500-ms time window (late N2pc and SPCN components). These findings are consistent with models of attention and emotion that posit rapid attentional prioritization of threat.

ARTICLE UPDATE - See no evil: Directing visual attention within unpleasant images modulates the electrocortical response.

Dunning JP, Hajcak G.

Psychophysiology, in press

The late positive potential (LPP) is larger for emotional than neutral stimuli, and reflects increased attention to motivationally salient stimuli. Recent studies have shown that the LPP can also be modulated by stimulus meaning and task relevance. The present studies sought to determine whether the magnitude of the LPP can be manipulated by directing attention to more or less arousing aspects within an emotional stimulus. To this end, trials included a passive viewing and directed attention portion. In both Studies 1 and 2, unpleasant compared to neutral images were associated with an increased LPP during passive viewing; additionally, directing attention to non-arousing compared to highly arousing areas of unpleasant images resulted in a decreased LPP. Results are discussed in terms of the utility of using the LPP to understand emotion-cognition interactions, especially with regard to directed visual attention as an emotion regulation strategy.

ARTICLE UPDATE - Electrophysiological correlates of decreasing and increasing emotional responses to unpleasant pictures.

Moser JS, Krompinger JW, Dietz J, Simons RF.

Psychophysiology, in press

We examined event-related brain potential (ERP) modulations during the anticipation and processing of unpleasant pictures under instructions to cognitively decrease and increase negative emotion. Instructions to decrease and increase negative emotion modulated the ERP response to unpleasant pictures in the direction of emotional intensity beginning around 400 ms and lasting several seconds. Decrease, but not increase, instructions also elicited enhanced frontal negativity associated with orienting and preparation prior to unpleasant picture onset. Last, ERP modulation by unpleasant pictures began around 300 ms, just prior to regulation effects, suggesting that appraisal of emotion occurs before emotion regulation. Together, the current findings underscore the utility of ERPs in illuminating the time course of emotion modulation and regulation that may help to refine extant theoretical models.

ARTICLE UPDATE - Stereotype threat and executive resource depletion: Examining the influence of emotion regulation.

Johns M, Inzlicht M, Schmader T.

Journal of Experimental Psychology: General, 137, 691-705

Research shows that stereotype threat reduces performance by diminishing executive resources, but less is known about the psychological processes responsible for these impairments. The authors tested the idea that targets of stereotype threat try to regulate their emotions and that this regulation depletes executive resources, resulting in underperformance. Across 4 experiments, they provide converging evidence that targets of stereotype threat spontaneously attempt to control their expression of anxiety and that such emotion regulation depletes executive resources needed to perform well on tests of cognitive ability. They also demonstrate that providing threatened individuals with a means to effectively cope with negative emotions--by reappraising the situation or the meaning of their anxiety--can restore executive resources and improve test performance. They discuss these results within the framework of an integrated process model of stereotype threat, in which affective and cognitive processes interact to undermine performance.

ARTICLE UPDATE - Validation of affective and neutral sentence content for prosodic testing.

Russ JB, Gur RC, Bilker WB.

Behavior Research Methods, 40,935-939

Conducting a study of emotional prosody often requires that one have a valid set of stimuli for assessing perceived emotion in vocal intonation. In this study, we created a list of sentences with both affective and neutral content, and then validated them against rater opinion. Participants read sentences with content that implied happiness, sadness, anger, fear, or neutrality and rated how well they could imagine each sentence being expressed in each emotion. Coefficients of variation and intraclass correlations were calculated to narrow the list to affective sentences that had high agreement and neutral sentences that had low agreement. We found that raters could easily identify most emotional content and did not ascribe any unique emotion to most neutral content. We also found differences between the intensity of male and female ratings. The final list of sentences is available on the Internet (www.med.upenn.edu/bbl/) and can be recorded for use as stimuli for prosodic studies.

ARTICLE UPDATE - Exploring the motivational brain: effects of implicit power motivation on brain activation in response to facial expressions of emoti

Schultheiss OC, Wirth MM, Waugh CE, Stanton SJ, Meier EA, Reuter-Lorenz P.

Social, Cognitive, Affective Neuroscience, in press

This study tested the hypothesis that implicit power motivation (nPower), in interaction with power incentives, influences activation of brain systems mediating motivation. Twelve individuals low (lowest quartile) and 12 individuals high (highest quartile) in nPower, as assessed per content coding of picture stories, were selected from a larger initial participant pool and participated in a functional magnetic resonance imaging study during which they viewed high-dominance (angry faces), low-dominance (surprised faces) and control stimuli (neutral faces, gray squares) under oddball-task conditions. Consistent with hypotheses, high-power participants showed stronger activation in response to emotional faces in brain structures involved in emotion and motivation (insula, dorsal striatum, orbitofrontal cortex) than low-power participants.

ARTICLE UPDATE - Rapid influence of emotional scenes on encoding of facial expressions: an ERP study.

Righart R, de Gelder B.

Social, Cognitive, Affective Neuroscience, 3, 270-278

In daily life, we perceive a person's facial reaction as part of the natural environment surrounding it. Because most studies have investigated how facial expressions are recognized by using isolated faces, it is unclear what role the context plays. Although it has been observed that the N170 for facial expressions is modulated by the emotional context, it was not clear whether individuals use context information on this stage of processing to discriminate between facial expressions. The aim of the present study was to investigate how the early stages of face processing are affected by emotional scenes when explicit categorizations of fearful and happy facial expressions are made. Emotion effects were found for the N170, with larger amplitudes for faces in fearful scenes as compared to faces in happy and neutral scenes. Critically, N170 amplitudes were significantly increased for fearful faces in fearful scenes as compared to fearful faces in happy scenes and expressed in left-occipito-temporal scalp topography differences. Our results show that the information provided by the facial expression is combined with the scene context during the early stages of face processing

ARTICLE UPDATE - Emotional modulation of visual and motor areas by dynamic body expressions of anger.

Pichon S, de Gelder B, Grezes J.

Social Neuroscience, 3, 199-212

The ability to detect emotional meaning in others' behavior constitutes a central component of social competence. Expressions of anger in particular present salient signals that play a major role in the regulation of social interactions. Investigations of human anger signals have to date used still pictures of facial expressions but so far the neurobiological basis of bodily communication of anger remains largely unknown. Using functional magnetic resonance imaging, the present study investigated the neural bases involved in perceiving anger signals emanating from the whole body. Our study also investigates what the presence of dynamic information adds to the perception of body expressions of anger. Participants were scanned while viewing stimuli (stills or videos) of angry and neutral whole-body expressions. Whole-body expressions of anger elicit activity in regions including the amygdala and the lateral orbitofrontal cortex, which play a role in the affective evaluation of the stimuli. Importantly, the perception of dynamic body expressions of anger additionally engages the hypothalamus, the ventromedial prefrontal cortex, the temporal pole and the premotor cortex, brain regions that are coupled with autonomic reactions and motor responses related to defensive behaviors.

ARTICLE UPDATE - Effective connectivity between amygdala and orbitofrontal cortex differentiates the perception of facial expressions.

Liang X, Zebrowitz LA, Aharon I.

Social Neuroscience, in press

Emotion research is guided both by the view that emotions are points in a dimensional space, such as valence or approach-withdrawal, and by the view that emotions are discrete categories. We determined whether effective connectivity of amygdala with medial orbitofrontal cortex (MOFC) and lateral orbitofrontal cortex (LOFC) differentiates the perception of emotion faces in a manner consistent with the dimensional and/or categorical view. Greater effective connectivity from left MOFC to amygdala differentiated positive and neutral expressions from negatively valenced angry, disgust, and fear expressions. Greater effective connectivity from right LOFC to amygdala differentiated emotion expressions conducive to perceiver approach (happy, neutral, and fear) from angry expressions that elicit perceiver withdrawal. Finally, consistent with the categorical view, there were unique patterns of connectivity in response to fear, anger, and disgust, although not in response to happy expressions, which did not differ from neutral ones.

ARTICLE UPDATE - Both predator and prey: emotional arousal in threat and reward.

Löw A, Lang PJ, Smith JC, Bradley MM.

Psychological Science, 19, 865-873

This research examined the psychophysiology of emotional arousal anticipatory to potentially aversive and highly pleasant outcomes. Human brain reactions (event-related potentials) and body reactions (heart rate, skin conductance, the probe startle reflex) were assessed along motivational gradients determined by apparent distance from sites of potential punishment or reward. A predator-prey survival context was simulated using cues that signaled possible money rewards or possible losses; the cues appeared to loom progressively closer to the viewer, until a final step when a rapid key response could ensure reward or avoid a punishing loss. The observed anticipatory response patterns of heightened vigilance and physiological mobilization are consistent with the view that the physiology of emotion is founded on action dispositions that evolved in mammals to facilitate survival by dealing with threats or capturing life-sustaining rewards.