The perception of fearful and happy facial expression is modulated by anxiety: an event-related potential study

Anxiety is supposed to interfere with cognitive and emotional processing and high level of trait-anxiety has been associated with an attentional bias for fearful faces, even in sub-clinical anxiety. On the basis of the Spielberger State and Trait Anxiety Inventory (STAI), twenty students were grouped as low vs. high anxious. Pictures from the Ekman and Friesen series were used in an event-related potentials study to investigate the neurophysiological correlates of the emotional processing of fear and happiness in sub-clinical anxiety. Subjects were confronted with a visual oddball design, in which they had to detect, as quickly as possible, deviant happy or fearful faces amongst a train of standard stimuli (neutral faces). Anxiety does not modify early perceptual (N100, P100, N170, VPP) or attentional (N2b) component, but later components are affected. Indeed, high anxious subjects are faster to detect deviant faces as suggested by earlier reaction times and P3b component. However, they show a reduced ability to process the emotional content of faces, this deficit being indexed by a decreased N300 component. Indeed, N300 is supposed to be particularly sensitive to affective features of stimuli rather than to physical characteristics. We propose that the earlier P3b observed in high anxious subjects could be interpreted as a way to overcome the deficient emotional appraisal by a more salient conscious processing.     

When does size not matter? Effects of stimulus size on affective modulation

Motivationally relevant stimuli have been shown to receive prioritized processing compared to neutral stimuli at distinct processing stages. This effect has been related to the evolutionary importance of rapidly detecting dangers and potential rewards and has been shown to be modulated by the distance between an organism and a faced stimulus. Similarly, recent studies showed degrees of emotional modulation of autonomic responses and subjective arousal ratings depending on stimulus size. In the present study, affective modulation of pictures presented in different sizes was investigated by measuring event-related potentials during a two-choice categorization task. Results showed significant emotional modulation across all sizes at both earlier and later stages of processing. Moreover, affective modulation of earlier processes was reduced in smaller compared to larger sizes, whereas no changes in affective modulation were observed at later stages.     

Time course of affective processing bias in major depression: An ERP study

The current study investigated the time course of the affective processing bias in major depressive disorder (MDD) in a visual three-stimulus semantic oddball task using event-related potentials (ERPs). MDD patients showed decreased P1 latency over right posterior regions to negative relative to positive target stimuli, reflecting a very early onset of the negativity bias in emotional perception. Compared to controls, MDD patients showed enlarged anterior P2 amplitude to positive target stimuli, reflecting an affective bias in the early attentional stages of processing. In addition, MDD patients showed relatively high N2 and reduced P3 amplitudes to negative compared with positive target stimuli, as well as marginally reduced N2 amplitude to positive target stimuli compared with controls. This suggests that the negativity bias also occurs during later strategic evaluation stages. Therefore, the present study extended previous findings by demonstrating that the affective processing bias in MDD begins in the early stages of perceptual processing and continues at later cognitive stages.     

Event-related potentials (ERPs) to hemifield presentations of emotional stimuli: differences between depressed patients and healthy adults in P3 amplitude and asymmetry.

Depression may involve dysfunction of right parietotemporal cortex, a region activated during perception of affective stimuli. To further test this hypothesis, event-related brain potentials (ERPs) were measured in a paradigm previously shown to produce ERP asymmetries to affective stimuli over parietal sites in healthy adults. Pictures of patients with dermatological diseases showing disordered or healed facial areas before (negative) or after (neutral) surgical treatment were briefly exposed for 250 ms to either the left or right hemifield. ERPs of 30 unmedicated, unipolar depressed patients and 16 healthy adults, all right-handed, were recorded from 30 electrodes. A principal components analysis extracted factors which closely corresponded to distinctive ERP components previously reported for this task (N1, N2, early P3, late P3, slow wave). Significant effects of emotional content, i.e. enhanced amplitudes to negative than neutral stimuli, were found for early and late P3. Control subjects showed significant hemispheric asymmetries of emotional processing for late P3 (peak latency 460 ms), with the largest emotional content effects over the right parietal region. In striking contrast to control subjects, depressed patients did not show an increase in late P3 for negative compared to neutral stimuli over either hemisphere and had smaller late P3 amplitude than control subjects. Patients did, however, show larger early P3 (peak latency 330 ms) to negative than neutral stimuli. Results suggest intact early discrimination but abnormal late appraisal of affective content in depression, which may arise from selective inhibition of right parietal regions integral for perceiving and evaluating emotional stimuli.     

Time course of attentional modulations on automatic emotional processing

In a previous study using event-related potentials (ERPs) [S. Doallo, S. Rodríguez Holguín, F. Cadaveira, Attentional load affects automatic emotional processing: evidence from event-related potentials, Neuroreport 17 (2006) 1797-1801], we reported that differential responses to unattended peripheral affective pictures, as reflected by N1-P2 modulations at posterior regions, are modulated by attentional load at fixation. Here, new analyses of these data were performed to evaluate whether a sustained, broadly distributed, negative shift in the unattended pictures ERP waveforms, which displayed larger amplitudes for emotional stimuli, reflects an additional differential response to the emotional content. Under low-load conditions, unpleasant (versus neutral) pictures elicited greater negativities in the 80-140 ms latency range over frontocentral sites and more centroparietally distributed from 200 to 280 ms. These findings provide further evidence of the time course of emotional processing at unattended locations and its modulation by attentional load.     

Effects of aging on functional connectivity of the amygdala during negative evaluation: a network analysis of fMRI data

Previous evidence has suggested both preserved emotional function in aging and age-related differences in emotional processing, but the neural networks underlying such processing alterations in the context of preserved affective function are not clear. Using event-related fMRI, we scanned young and older adults while they made valence ratings for emotional pictures. Behavioral results showed a similar pattern of emotional evaluation, but older adults experienced negatively valenced pictures as being less negative. Consistent with behavioral findings, we identified common activity in the right amygdala, but age-related differences in the functional connectivity of this region with the rest of the brain. Compared to young adults, older adults had greater functional connectivity between the right amygdala and ventral anterior cingulate cortex, possibly reflecting increased emotional regulation. Conversely, older adults showed decreased functional connectivity with posterior brain regions, likely reflecting decreased perceptual processing. Thus, age-related differences in evaluating negatively valenced stimuli might reflect decreased perceptual processing of these stimuli, as well as the engagement of control processes that inhibit the response to negative emotion.     

Brain potentials in affective picture processing: covariation with autonomic arousal and affective report

Emotionally arousing picture stimuli evoked scalp-recorded event-related potentials. A late, slow positive voltage change was observed, which was significantly larger for affective than neutral stimuli. This positive shift began 200-300 ms after picture onset, reached its maximum amplitude approximately 1 s after picture onset, and was sustained for most of a 6-s picture presentation period. The positive increase was not related to local probability of content type, but was accentuated for pictures that prompted increased autonomic responses and reports of greater affective arousal (e.g. erotic or violent content). These results suggest that the late positive wave indicates a selective processing of emotional stimuli, reflecting the activation of motivational systems in the brain.     

How negative affect influences neural control processes underlying the resolution of cognitive interference: an event-related fMRI study

In this event-related fMRI study, we sought to investigate the influence of negative affect on the processing of two kinds of cognitive interference: Stroop-interference and oddball interference. For our purpose, we adopted an oddball variant of the Stroop task in which Stroop-interference and oddball interference conditions were created by presenting incongruent and rarely occurring word meanings, respectively. Immediately preceding the target stimuli, we presented pictures of the International Affective Picture System which were either emotionally negative and arousing or emotionally neutral, providing two affective conditions under which the cognitive task was administered. Both the behavioral and the neuroimaging data exhibited an interaction effect between emotional and cognitive condition. First, the emotion induction selectively impaired behavioral performance on interference trials while behavioral measures on non-interference trials were roughly identical in both emotional conditions. Second, in the negative emotional condition there was incremental interference-related activation in control-related regions (fronto-parietal cortices). Taken together, findings suggest that negative affect specifically disturbs the neural control processes that in a neutral affective state allow to select task-relevant information and to shield its processing from task-irrelevant distraction. Accordingly, agents in a negative affective state have to exert enhanced control efforts to resolve cognitive interference. Additional connectivity analyses revealed that a negative coupling between lateral PFC on the one hand and amygdala and OFC on the other is related to enhanced interference resolution which can be tentatively interpreted as evidence that emotional regulation is an integrated part of an agent's efforts to preserve cognitive performance in affective situations.     

Neural response to emotional pictures is unaffected by concurrent task difficulty: an event-related potential study

The late positive potential (LPP) is an event-related potential that is enhanced when viewing arousing (pleasant and unpleasant) pictures compared to neutral pictures. The affective modulation of the LPP is believed to reflect the increased attention to, and perceptual processing of, emotional stimuli. The present study examined whether concurrent task difficulty (performing mathematics) would modulate the LPP while participants viewed emotionally arousing stimuli. Results indicated that the LPP was larger following pleasant and unpleasant stimuli than it was following neutral stimuli; moreover, the magnitude of this increase was not influenced by concurrent task difficulty. This finding suggests that the affective modulation of neural activity during picture viewing is relatively automatic and is insusceptible to competing task demands. Results are further discussed in terms of the LPP's role in motivated attention and implications for research on emotion regulation.     

Perceived social support moderates neural reactivity to emotionally valenced stimuli during pregnancy

Reactivity to emotional information, measurable at the level of neural activity using event-related potentials, is linked to symptoms of affective disorders. Behavioral evidence suggests that contextual factors, such as social support, can alter emotional reactivity such that affective responding is normalized when social support is high. This possibility remains largely untested at the neural level, specifically through approaches that can offer insight into the mechanistic processes contributing to individual differences in emotional reactivity. Yet, such knowledge could be useful for prevention and intervention efforts, particularly with groups at risk for increased emotional reactivity, such as pregnant mothers for whom emotional distress predicts both maternal and child outcomes. Expectant mothers took part in a longitudinal study that tested whether the late positive potential (LPP), a neural index of reactivity to emotional information, was moderated by maternal perceptions of social support. In the third trimester of pregnancy, lower perceived social support was associated with an absence of a traditional LPP effect, which differentiates valenced from neutral stimuli. Findings suggest that perceptions of social support may normalize emotional processing at the neural level and highlight the potential importance of social support modulation of emotional reactivity during times of known biological change.     

Judgments of emotion in words and faces: ERP correlates

Visual event-related potentials (ERPs) to two types of stimuli (faces and words) were analyzed to determine the effects of the perceived emotional connotations of the stimuli (positive, neutral, or negative) in 10 right-handed normal functioning adult males. Principal component analysis (PCA) of the ERPs reveals 5 factors accounting for over 90% of the ERP waveform variance for both faces and words. In the facial data, two ERP components varied in amplitude according to the perceived emotional connotation of the stimulus. For the P3 component, neutrally rated stimuli produced significantly larger amplitudes than stimuli rated as positive or negative. This effect was lateralized to the left hemisphere. A later positive component, the slow wave (448-616 ms), manifested complementary effects, i.e. faces perceived as positive and negative produced larger amplitudes than those perceived as neutral over the right hemisphere. The verbal stimuli did not result in significant main effects for perceived emotional connotation, but produced subtle connotation-related differences in slow wave topography. Hemispheric asymmetries, unrelated to affective connotation, were evident in the verbal data, manifesting different patterns of lateralization depending on the ERP component. The results suggest that differential processing of emotional connotation affects ERP waveforms and that the effects can be understood in terms of ERP components known to be associated with more general aspects of cognitive processing.     

Affective modulation of brain potentials to painful and nonpainful stimuli

In accordance with the emotional priming hypothesis, emotions seem to modulate pain perception and pain tolerance thresholds. To further evaluate this association, event-related brain potentials (ERPs) elicited by painful and nonpainful electrical stimuli during processing of positive, neutral, and negative valenced pictures were recorded from 30 healthy volunteers. Valence of pictures affected pain ratings and the N150 elicited by painful stimuli, with lowest amplitudes for positive pictures and highest amplitudes for negative pictures. The P260 elicited by painful and nonpainful stimuli was modulated by arousal with reduced amplitudes with arousing (positive or negative) compared to neutral pictures. N150 amplitudes varying with picture valence seem to reflect an affective modulation of pain perception whereas P260 amplitudes varying with picture arousal rather reflect non-pain-specific attentional processes.     

Modulation of spinal reflexes: Arousal, pleasure, action

The human startle reflex is reliably modulated by the affective valence of foreground pictures, with larger reflexes elicited when viewing unpleasant relative to pleasant scenes. If this modulation is due to priming of the defensive startle reflex by an aversive foreground, a different pattern should occur for a reflex that is not inherently defensive in nature. In the current study, affective modulation was investigated using the spinal tendinous (T) reflex, which is well documented as sensitive to differences in arousal and is involved in actions that are both appetitively and defensively motivated. As such, T reflexes elicited during unpleasant pictures were not expected to be augmented relative to those elicited in the context of pleasant pictures. Results showed that T reflexes were facilitated during processing of arousing stimuli – either pleasant or unpleasant relative to low-arousal neutral materials. These effects of emotional stimuli on T-reflex amplitude are consistent with the hypothesis that motivational priming underlies affective reflex modulation.     

Emotion and psychopathy: startling new insights

Abnormal affective response in psychopaths is conceptualized within a broad theory of emotion that emphasizes reciprocal appetitive and defensive motivational systems. The startle response is proposed as a specific measure of the directional component of emotional activation. I review the literature that indicates that criminal psychopaths do not show the expected potentiation of the startle reflex that normally occurs during processing of aversive stimuli such as unpleasant photographs or punishment cues. Evidence is presented to demonstrate that this deviant response pattern is specific to individuals who display the classic affective symptoms of psychopathy. The core emotional deviation in psychopathy could be a deficit in fear response, which is defined as a failure of aversive cues to prime normal defensive actions. This emotional deficit may represent an extreme variant of normal temperament.     

The interplay between attention and long-term memory in affective habituation

Previous studies have consistently shown that repeated exposure to emotional stimuli leads to a reduction in cortical and autonomic responses (affective habituation). These findings emerge from studies conducted within a single experimental session, preventing the possibility of disentangling short-term from long-term habituation effects. The present study investigated whether affective habituation reflects a short-living learning process, or a more stable change involving long-term memory. Participants went through a first habituation phase consisting of 80 repetitions of the same set of emotional and neutral pictures, when event-related potentials and oscillatory activity were measured (Session 1). Crucially, after a 1-day interval, the same participants were exposed to a second habituation phase with the same stimuli that had been seen before. Results showed that the attenuation of the late positive potential (LPP) affective modulation prompted throughout repetitions of Session 1 remained unchanged after a 1-day interval, and this between-session habituation effect, which was specific to repeated exemplars, was consistent across different emotional contents. Alpha desynchronization was clearly enhanced for pictures of erotica and mutilation and this modulatory pattern remained fairly stable over repetitions. Altogether, these findings suggest that LPP affective habituation is not a short-living learning process, but, rather, reflects a strengthened long-term memory representation of specific repeated stimuli.     

Additive effects of serotonin transporter and tryptophan hydroxylase-2 gene variation on neural correlates of affective processing

Individual differences in brain response to emotional stimuli have previously been associated with gene variations within the serotonin transporter (5-HTT) and tryptophan hydroxylase-2 (TPH2) genes. We recently reported that these two genes exhibit an additive effect, based on recordings of event-related potentials (ERPs) from individuals viewing emotional scenes. The current study was designed to replicate and extent this initial report in an independent study sample, and use functional magnetic resonance imaging (fMRI) to identify specific neural loci that may mediate the 5-HTT-TPH2 additive effect. Furthermore, we sought to obtain convergent evidence for a gene-gene additive effect by collecting fMRI data from the same individuals engaged in two different cognitive-affective tasks, using emotional and neutral facial expressions and word stimuli. We found evidence for an additive effect of 5-HTT-TPH2 genotype, which was most robust in the putamen, a region rich in both 5-HTT and TPH2 protein, but was also observed in the amygdala at a less stringent threshold, and in other cortical regions. The additive effect was more robust effect for visuospatial than for verbal stimuli, and more robust for negatively than for positively valenced stimuli. These findings confirm and extend the additive effect of two critical genes in the serotonergic regulation of neural processing of affective stimuli, and identify the striatum as a critical site where is gene-gene regulation takes place.     

Emotional deficit in subjects with psychopathic tendencies as assessed by the Minnesota Multiphasic Personality Inventory-2: an event-related potentials study

On the basis of the Minnesota Multiphasic Personality Inventory-2 (MMPI-2), 20 male students were separated in low (LP) versus high (HP) subjects with psychopathic tendencies. Pictures from the Ekman and Friesen series were used in an event-related potentials study to investigate the neurophysiological correlates of the emotional deficit described in psychopathy. Subjects were confronted with a visual oddball design, in which they had to detect, as quickly as possible, deviant happy, sad or fearful faces amongst a train of standard stimuli (neutral faces). Behavioral results suggest that LP subjects are more efficient in the detection of emotional deviant faces. This emotional deficit was neurophysiologically indexed, in HP subjects, by a decreased N300 component, which is supposed to be particularly sensitive to affective features of stimuli rather than to physical characteristics. Moreover, independently of the nature of the deviant faces, the P3a component shows an earlier latency in the HP group as compared to the LP group, whereas the reversed pattern was shown for the P3b component. Implications of these results were discussed.     

Parietal processing of affect and cognition: cerebral organization in strongly lateralized left-handed subjects

Cerebral organization for the processing of emotion and for the differential processing of affect and cognition have thus far been studied exclusively in right-handed subjects. Since handedness is related to cerebral dominance, our understanding of the neural processing of affect and cognition should be further enhanced by the study of sinistral subjects. The present study examined the differential parietal processing of emotional stimuli under affective and cognitive conditions in left-handed subjects. The subjects were exposed to emotional stimuli presented under affective and cognitive instructional sets and to neutral, control stimuli. Bilateral electroencephalographic and bilateral electrodermal activity data showed greater responsiveness to emotional than neutral stimuli but no differential lateralization of affect and cognition in left-handed subjects. Results also showed that female subjects were more responsive to emotional stimuli than male subjects. Results were interpreted as suggesting that left-handed individuals may be less strongly lateralized for the differential processing of affect and cognition than right-handed individuals and that females may have greater focal organization for the processing of emotion than males.     

Selective influence of the menstrual cycle on perception of stimuli with reproductive significance: an event-related potential study

In this study, we examined changes in the event-related potential (ERP) to stimuli with and without reproductive significance occurring during the menstrual cycle. Eleven spontaneously cycling women were tested during three menstrual phases (menses, ovulatory phase, luteal phase) differing in plasma concentrations of gonadal hormones. ERPs were recorded while subjects were presented with slides showing pictures from four different stimulus categories (sexual stimuli, babies, people occupied with body care, ordinary people). Slides were presented randomly in the context of two tasks, requiring either affective processing (i.e., to judge the emotional content of a slide as positive, neutral, or negative) or structural processing (i.e., to estimate the number of parallel thin lines inserted in each picture). Menstrual phase primarily affected a late positive component (LPC) peaking 550-600 ms poststimulus. The effects were as follows: (i) During the ovulatory phase, amplitude of the LPC to sexual stimuli was larger than that evoked by the other stimulus categories. (ii) This relationship was not apparent during the other menstrual phases or (iii) during the ovulatory phase when the task required structural processing. The ovulatory increase in LPC positivity to sexual stimuli suggests a greater valence of these stimuli during a phase of increased sexual desire. The data indicate a specific effect of the menstrual cycle on the processing of sexual stimuli that increases with deeper emotional processing.     

Event-related P3a and P3b in response to unpredictable emotional stimuli

In natural situations, unpredictable events processing often interacts with the ongoing cognitive activities. In a similar manner, the insertion of deviant unpredictable stimuli into a classical oddball task evokes both the P3a and P3b event-related potentials (ERPs) components that are, respectively, thought to index reallocation of attentional resources or inhibitory process and memory updating mechanism. This study aims at characterising the influence of the emotional arousal and valence of a deviant and unpredictable non-target stimulus on these components. ERPs were recorded from 28 sites during a visual three-stimulus oddball paradigm. Unpleasant, neutral and pleasant pictures served as non-target unpredictable items and subjects were asked to realize a perceptually difficult standard/target discrimination task. A temporal principal component analysis (PCA) allowed us to show that non-target pictures elicited both a P3a and a P3b. Moreover, the P3b component was modulated by the emotional arousal and the valence of the pictures. Thus, the memory updating process may be modulated by the affective arousal and valence of unpredictable disturbing stimuli, even if the task does not require any explicit emotional categorization.