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 - 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 - 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 - Attention, emotion, and deactivation of default activity in inferior medial prefrontal cortex.

Geday J, Gjedde A.

Brain and Cognition, in press

Attention deactivates the inferior medial prefrontal cortex (IMPC), but it is uncertain if emotions can attenuate this deactivation. To test the extent to which common emotions interfere with attention, we measured changes of a blood flow index of brain activity in key areas of the IMPC with positron emission tomography (PET) of labeled water (H(2)(15)O) uptake in brain of 14 healthy subjects. The subjects performed either a less demanding or a more demanding task of attention while they watched neutral and emotive images of people in realistic indoor or outdoor situations. In the less demanding task, subjects used the index finger to press any key when a new image appeared. In the more demanding task, subjects chose the index or middle finger to press separate keys for outdoor and indoor scenes. Compared to the less demanding task, in a global search of all gray matter, the more demanding significantly lowered blood flow (rCBF) in left IMPC, left and right insula, and right amygdala, and significantly raised blood flow in motor cortex and right precuneus. Restricted searches of rCBF changes by emotion, at coordinates of significant effect in previous studies of the medial prefrontal and temporal cortices, revealed significant activation in the fusiform gyrus, independently of the task. In contrast, we found no effect of emotional content in the IMPC, where emotions failed to override the effect of the task. The results are consistent with a role of the IMPC in the selection among competitive inputs from multiple brain regions, as predicted by the theory of a default mode of brain function. The absent emotional interference with the deactivation of the default state suggests that the inferior prefrontal cortex continued to serve the attention rather than submit to the distraction.

ARTICLE UPDATE - The combined effect of gaze direction and facial expression on cueing spatial attention.

Pecchinenda A, Pes M, Ferlazzo F, Zoccolotti P.

Emotion, 8, 628-634

Empirical evidence shows an effect of gaze direction on cueing spatial attention, regardless of the emotional expression shown by a face, whereas a combined effect of gaze direction and facial expression has been observed on individuals' evaluative judgments. In 2 experiments, the authors investigated whether gaze direction and facial expression affect spatial attention depending upon the presence of an evaluative goal. Disgusted, fearful, happy, or neutral faces gazing left or right were followed by positive or negative target words presented either at the spatial location looked at by the face or at the opposite spatial location. Participants responded to target words based on affective valence (i.e., positive/negative) in Experiment 1 and on letter case (lowercase/uppercase) in Experiment 2. Results showed that participants responded much faster to targets presented at the spatial location looked at by disgusted or fearful faces but only in Experiment 1, when an evaluative task was used. The present findings clearly show that negative facial expressions enhance the attentional shifts due to eye-gaze direction, provided that there was an explicit evaluative goal present.

ARTICLE UPDATE - Visual search is not blind to emotion.

Gerritsen C, Frischen A, Blake A, Smilek D, Eastwood JD.

Perception and Psychophysics, 70, 1047-1059

A series of three visual search tasks revealed more efficient search for hostile than for peaceful faces among neutral face distractors. Given that this effect has been observed inconsistently in prior literature, meta-analytic methods were employed for evaluating data across three experiments in order to develop a more valid estimate of the potentially small effect size. Furthermore, in the present experiments, different emotional meanings were conditioned to identical faces across observers, thus eliminating confounds between the physical characteristics and the emotional valences of the face stimuli. On the basis of the present findings, we argue that the visual system is capable of determining a face's emotional valence before the face becomes the focus of attention, and that emotional valence can be used by the visual system to determine subsequent attention allocation. However, meta-analytic results indicate that emotional valence makes a relatively small contribution to search efficiency in the present search context.

ARTICLE UPDATE - The persistence of attention to emotion: Brain potentials during and after picture presentation.

Hajcak G, Olvet DM.

Emotion, 8, 250-255

Emotional stimuli have been shown to elicit increased perceptual processing and attentional allocation. The late positive potential (LPP) is a sustained P300-like component of the event-related potential that is enhanced after the presentation of pleasant and unpleasant pictures as compared with neutral pictures. In this study, the LPP was measured using dense array electroencephalograph both before and after pleasant, neutral, and unpleasant images to examine the time course of attentional allocation toward emotional stimuli. Results from 17 participants confirmed that the LPP was larger after emotional than neutral images and that this effect persisted for 800 ms after pleasant picture offset and at least 1,000 ms after unpleasant picture offset. The persistence of increased attention after unpleasant compared to pleasant stimuli is consistent with the existence of a negativity bias. Overall, these results indicate that attentional capture of emotion continues well beyond picture presentation and that this can be measured with the LPP. Implications and directions for future research are discussed.

ARTICLE UPDATE - Beyond fear: rapid spatial orienting toward positive emotional stimuli.

Brosch T, Sander D, Pourtois G, Scherer KR.

Psychological Science, 19, 362-370

There is much empirical evidence for modulation of attention by negative-particularly fear-relevant-emotional stimuli. This modulation is often explained in terms of a fear module. Appraisal theories of emotion posit a more general mechanism, predicting attention capture by stimuli that are relevant for the needs and goals of the organism, regardless of valence. To examine the brain-activation patterns underlying attentional modulation, we recorded event-related potentials from 20 subjects performing a dot-probe task in which the cues were fear-inducing and nurturance-inducing stimuli (i.e., anger faces and baby faces). Highly similar validity modulation was found for the P1 time-locked to target onset, indicating early attentional capture by both positive and negative emotional stimuli. Topographic segmentation analysis and source localization indicate that the same amplification process is involved whether attention orienting is triggered by negative, fear-relevant stimuli or positive, nurturance-relevant stimuli. These results confirm that biological relevance, and not exclusively fear, produces an automatic spatial orienting toward the location of a stimulus.

ARTICLE UPDATE - Selective Attention to Affective Value Alters How the Brain Processes Olfactory Stimuli.

Rolls ET, Grabenhorst F, Margot C, da Silva MA, Velazco MI.

Journal of Cognitive Neuroscience, in press

How does selective attention to affect influence sensory processing? In an functional magnetic resonance imaging investigation, when subjects were instructed to remember and rate the pleasantness of a jasmin odor, activations were greater in the medial orbito-frontal and pregenual cingulate cortex than when subjects were instructed to remember and rate the intensity of the odor. When the subjects were instructed to remember and rate the intensity, activations were greater in the inferior frontal gyrus. These top-down effects occurred not only during odor delivery but started in a preparation period after the instruction before odor delivery, and continued after termination of the odor in a short-term memory period. Thus, depending on the context in which odors are presented and whether affect is relevant, the brain prepares itself, responds to, and remembers an odor differently. These findings show that when attention is paid to affective value, the brain systems engaged to prepare for, represent, and remember a sensory stimulus are different from those engaged when attention is directed to the physical properties of a stimulus such as its intensity. This differential biasing of brain regions engaged in processing a sensory stimulus depending on whether the cognitive demand is for affect-related versus more sensory-related processing may be an important aspect of cognition and attention. This has many implications for understanding the effects not only of olfactory but also of other sensory stimuli.

ARTICLE UPDATE - Distinct effects of attention and affect on pain perception and somatosensory evoked potentials.

Kenntner-Mabiala R, Andreatta M, Wieser MJ, Mühlberger A, Pauli P.

Biological Psychology, in press

The influence of affect and attention on sensory and affective pain as well as on somatosensory evoked potentials in response to painful and nonpainful electrical stimuli was investigated in a single experimental design. Affect was induced by pictures from the International Affective Picture System; attention was manipulated by asking participants to focus attention either on the pictures or on the electrical stimuli. Sensory and affective pain ratings were generally lower during exposure to positive compared to negative and neutral pictures. Attention modulated only sensory pain ratings with lower ratings with an attention focus on pictures than with an attention focus on sensory pain. The N150 was modulated by picture valence, the P260 by picture arousal. Furthermore, the P260 was modulated by attention with highest amplitudes with an attention focus on the stimulus intensity. This study provides neurophysiological evidence that attention and affect have distinct effects on pain processing.

ARTICLE UPDATE - Immediate memory consequences of the effect of emotion on attention to pictures.

Talmi D, Anderson AK, Riggs L, Caplan JB, Moscovitch M.

Learning & Memory, 15, 172-182

Emotionally arousing stimuli are at once both highly attention grabbing and memorable. We examined whether emotional enhancement of memory (EEM) reflects an indirect effect of emotion on memory, mediated by enhanced attention to emotional items during encoding. We tested a critical prediction of the mediation hypothesis-that regions conjointly activated by emotion and attention would correlate with subsequent EEM. Participants were scanned with fMRI while they watched emotional or neutral pictures under instructions to attend to them a lot or a little, and were then given an immediate recognition test. A region in the left fusiform gyrus was activated by emotion, voluntary attention, and subsequent EEM. A functional network, different for each attention condition, connected this region and the amygdala, which was associated with emotion and EEM, but not with voluntary attention. These findings support an indirect cortical mediation account of immediate EEM that may complement a direct modulation model.

ARTICLE UPDATE - Emotional valence and arousal interact in attentional control.

Jefferies LN, Smilek D, Eich E, Enns JT.

Psychological Science, 19, 290-295

A recent study demonstrated that observers' ability to identify targets in a rapid visual sequence was enhanced when they simultaneously listened to happy music. In the study reported here, we examined how the emotion-attention relationship is influenced by changes in both mood valence (negative vs. positive) and arousal (low vs. high). We used a standard induction procedure to generate calm, happy, sad, and anxious moods in participants. Results for an attentional blink task showed no differences in first-target accuracy, but second-target accuracy was highest for participants with low arousal and negative affect (sad), lowest for those with strong arousal and negative affect (anxious), and intermediate for those with positive affect regardless of their arousal (calm, happy). We discuss implications of this valence-arousal interaction for the control of visual attention.

ARTICLE UPDATE - Brain activation and defensive response mobilization during sustained exposure to phobia-related and other affective pictures in spid

Wendt J, Lotze M, Weike AI, Hosten N, Hamm AO.

Psychophysiology, in press

This study explored defensive response mobilization as well as fMRI responses during sustained exposure to phobia-relevant stimuli. To test the specificity of affective physiology and brain activation, neutral and other affective stimuli were included. Phobia-specific startle potentiation was maintained and autonomic responses even increased during sustained phobic stimulation. Viewing of spider pictures also resulted in increased activation of the amygdala in spider-phobic participants. This effect, however, was not fear specific because other affective materials evoked comparable signal strength in the amygdala. In contrast, insula activation was specifically increased during sustained phobic exposure in phobic volunteers. These data suggest that the activation of the amygdala in fMRI studies primarily indexes the detection of motivationally relevant stimuli whereas the insula might be more specifically linked to defensive response mobilization.

ARTICLE UPDATE - As Time Goes By: Temporal Constraints on Emotional Activation of Inferior Medial Prefrontal Cortex

Jacob Geday, Ron Kupers and Albert Gjedde

Cerebral Cortex, 17, 2753-2759

To investigate the influence of stimulus duration on emotional processing, we measured changes of regional cerebral blood flow (rCBF) in 14 healthy subjects who viewed neutral or emotional images presented for 3 or 6 s. Presentation for 3 s reproduced the previous result of higher rCBF in inferior medial prefrontal cortex (IMPC) during neutral than emotional stimulation. Six-second presentation reverted this relationship, with activity in IMPC being higher during emotional stimulation. Prolonged stimulus presentation attenuated the rise of rCBF associated with emotions in left parietal cortex and cerebellar hemisphere. We speculate that the different rCBF during neutral and emotional stimulation for 6 s is a consequence of attention divided between the emotional stimuli and their associations. Thus, prefrontal activity rises when a cognitive task accompanies emotional stimulation because several cognitive processes compete for attention. The IMPC may serve the mechanism of attention underlying the concept of a default mode of brain function, selecting among competitive inputs from multiple brain regions rather than just processing emotions. The results emphasize the importance of implicit cognitive processing during emotional activation, however, unintended.

ARTICLE UPDATE - The Costs of Emotional Attention: Affective Processing Inhibits Subsequent Lexico-semantic Analysis

Niklas Ihssen, Sabine Heim and Andreas Keil

Journal of Cognitive Neuroscience, 19, 1932-1949

The human brain has evolved to process motivationally relevant information in an optimized manner. The perceptual benefit for emotionally arousing material, termed motivated attention, is indexed by electrocortical amplification at various levels of stimulus analysis. An outstanding issue, particularly on a neuronal level, refers to whether and how perceptual enhancement for arousing signals translates into modified processing of information presented in temporal or spatial proximity to the affective cue. The present studies aimed to examine facilitation and interference effects of task-irrelevant emotional pictures on subsequent word identification. In the context of forced-choice lexical decision tasks, pictures varying in hedonic valence and emotional arousal preceded word/pseudoword targets. Across measures and experiments, high-arousing compared to low-arousing pictures were associated with impaired processing of word targets. Arousing pleasant and unpleasant pictures prolonged word reaction times irrespective of stimulus-onset asynchrony (80 msec, 200 msec, 440 msec) and salient semantic category differences (e.g., erotica vs. mutilation pictures). On a neuronal level, interference was reflected in reduced N1 responses (204–264 msec) to both target types. Paralleling behavioral effects, suppression of the late positivity (404–704 msec) was more pronounced for word compared to pseudoword targets. Regional source modeling indicated that early reduction effects originated from inhibited cortical activity in posterior areas of the left inferior temporal cortex associated with orthographic processing. Modeling of later reduction effects argues for interference in distributed semantic networks comprising left anterior temporal and parietal sources. Thus, affective processing interferes with subsequent lexico-semantic analysis along the ventral stream.

ARTICLE UPDATE - Unilateral Amygdala Lesions Hamper Attentional Orienting Triggered by Gaze Direction

Tomoko Akiyama, Motoichiro Kato, Taro Muramatsu, Satoshi Umeda, Fumie Saito and Haruo Kashima

Cerebral Cortex, in press

The newly discovered deficit in a bilateral amygdala-damaged case, of not being able to allocate attention to the critical feature of a face (Adolphs R, Gosselin F, Buchanan TW, Tranel D, Schyns P, Damasio AR. 2005. A mechanism for impaired fear recognition after amygdala damage. Nature. 433:68--72.), has opened a new window into the function of the amygdala. This case implies that the amygdala might be essential in detecting potentially relevant social stimuli, and directing attention accordingly. In this study, we have sought to test this implication by investigating the behavioral performance of 5 unilateral amygdala-damaged subjects on spatial cueing tasks. The tasks employed central gaze and arrow direction as cues to trigger attentional orienting in peripheral target detection. Although age-matched normal controls demonstrated a significant congruency effect such that targets presented congruently to cue direction elicited faster detection, amygdala subjects demonstrated no such congruency effect for gaze cues in the face of a significant congruency effect for arrow cues. The results suggest that the social valence of a stimulus is critical for amygdala involvement in visual processing. The results also support the implicated role of the amygdala in detecting and analyzing relevant social stimuli, and orienting attention accordingly.

ARTICLE UPDATE - Anxiety and orienting of gaze to angry and fearful faces.

Mogg K, Garner M, Bradley BP.

Biological Psychology, in press

Neuroscience research indicates that individual differences in anxiety may be attributable to a neural system for threat-processing, involving the amygdala, which modulates attentional vigilance, and which is more sensitive to fearful than angry faces. Complementary cognitive studies indicate that high-anxious individuals show enhanced visuospatial orienting towards angry faces, but it is unclear whether fearful faces elicit a similar attentional bias. This study compared biases in initial orienting of gaze to fearful and angry faces, which varied in emotional intensity, in high- and low-anxious individuals. Gaze was monitored whilst participants viewed a series of face-pairs. Results showed that fearful and angry faces elicited similar attentional biases. High-anxious individuals were more likely to direct gaze at intense negative facial expressions, than low-anxious individuals, whereas the groups did not differ in orienting to mild negative expressions. Implications of the findings for research into the neural and cognitive bases of emotion processing are discussed.

ARTICLE UPDATE - Neural Biases to Covert and Overt Signals of Fear: Dissociation by Trait Anxiety and Depression

Leanne M. Williams, Andrew H. Kemp, Kim Felmingham, Belinda J. Liddell, Donna M. Palmer and Richard A. Bryant

Journal of Cognitive Neuroscience, 19, 1595-1608

Although biases toward signals of fear may be an evolutionary adaptation necessary for survival, heightened biases may be maladaptive and associated with anxiety or depression. In this study, event-related potentials (ERPs) were used to examine the time course of neural responses to facial fear stimuli (versus neutral) presented overtly (for 500 msec with conscious attention) and covertly (for 10 msec with immediate masking to preclude conscious awareness) in 257 nonclinical subjects. We also examined the impact of trait anxiety and depression, assessed using psychometric ratings, on the time course of ERPs. In the total subject group, controlled biases to overtly processed fear were reflected in an enhancement of ERPs associated with structural encoding (120–220 msec) and sustained evaluation persisting from 250 msec and beyond, following a temporo-occipital to frontal topography. By contrast, covert fear processing elicited automatic biases, reflected in an enhancement of ERPs prior to structural encoding (80–180 msec) and again in the period associated with automatic orienting and emotion encoding (230–330 msec), which followed the reverse frontal to temporo-occipital topography. Higher levels of trait anxiety (in the clinical range) were distinguished by a heightened bias to covert fear (speeding of early ERPs), compared to higher depression which was associated with an opposing bias to overt fear (slowing of later ERPs). Anxiety also heightened early responses to covert fear, and depression to overt fear, with subsequent deficits in emotion encoding in each case. These findings are consistent with neural biases to signals of fear which operate automatically and during controlled processing, feasibly supported by parallel networks. Heightened automatic biases in anxiety may contribute to a cycle of hypervigilance and anxious thoughts, whereas depression may represent a "burnt out" emotional state in which evaluation of fear stimuli is prolonged only when conscious attention is allocated.

ARTICLE UPDATE - The shape of threat: Simple geometric forms evoke rapid and sustained capture of attention.

Larson, Christine L.; Aronoff, Joel; Stearns, Jeffrey J.

Emotion, 7, 526-534

Previous work has indicated that simple geometric shapes underlying facial expressions are capable of conveying emotional meaning. Specifically, a series of studies found that a simple shape, a downward-pointing "V," which is similar to the geometric configuration of the face in angry expressions, is perceived as threatening. A parallel line of research has determined that threatening stimuli more readily capture attention. In five experiments, the authors sought to determine whether this preferential processing was also present for the simple geometric form of a downward-pointing "V." Using a visual search paradigm, across these experiments the authors found that, when embedded in a field of other shapes, downward-pointing V's were detected faster and, in some cases, more accurately than identical shapes pointing upward. These findings indicate that the meaning of threat can be conveyed rapidly with minimal stimulus detail. In addition, in some cases, during trials of homogeneous fields of stimuli, fields of downward-pointing V's led to slower response times, suggesting that this shape's ability to capture attention may also extend to difficulty in disengaging attention as well.

ARTICLE UPDATE - Here is looking at you: Emotional faces predominate in binocular rivalry.

Alpers, Georg W.; Gerdes, Antje B. M.

Emotion, 7, 495-506

Two incompatible pictures compete for perceptual dominance when they are presented to one eye each. This so-called binocular rivalry results in an alternation of dominant and suppressed percepts. In accordance with current theories of emotion processing, the authors' previous research has suggested that emotionally arousing pictures predominate in this perceptual process. Three experiments were run with pictures of emotional facial expressions that are known to induce emotions while being well controlled in terms of physical characteristics. In Experiment 1, photographs of emotional and neutral facial expressions were presented of the same actor to minimize physical differences. In Experiment 2, schematic emotional expressions were presented to further eliminate low-level differences. In Experiment 3, a probe-detection task was conducted to control for possible response-biases. Together, these data clearly demonstrate that emotional facial expressions predominate over neutral expressions; they are more often the first percept and they are perceived for longer durations. This is not caused by physical stimulus properties or by response-biases. This novel approach supports that emotionally significant visual stimuli are preferentially perceived.