Extent and time‐course of competition in visual cortex between emotionally arousing distractors and a concurrent task

Emotionally arousing cues automatically attract attentional resources, which may be at the cost of processing task‐related information. Of central importance is how the visual system resolves competition for processing resources among stimuli differing in motivational salience. Here, we assessed the extent and time‐course of competition between emotionally arousing distractors and task‐related stimuli in a frequency‐tagging paradigm. Steady‐state visual evoked potentials (ssVEPs) were evoked using random‐dot kinematograms that consisted of rapidly flickering (8.57 Hz) dots, superimposed upon emotional or neutral distractor pictures flickering at 12 Hz. The time‐varying amplitude of the ssVEP evoked by the motion detection task showed a significant reduction to the task‐relevant stream while emotionally arousing pictures were presented as distractors. Competition between emotionally arousing pictures and moving dots began 450 ms after picture onset and persisted for an additional 2600 ms. Competitive effects of the overlapping task and picture stream revealed cost effects for the motion detection task when unpleasant pictures were presented as distractors between 450 and 1650 ms after picture onset, where an increase in ssVEP amplitude to the flickering picture stimulus was at the cost of ssVEP amplitude to the flickering dot stimulus. Cost effects were generalized to all emotionally arousing contents between 1850 and 3050 ms after picture onset, where the greatest amount of competition was evident for conditions in which emotionally arousing pictures, compared to neutral, served as distractors. In sum, the processing capacity of the visual system as measured by ssVEPs is limited, resulting in prioritized processing of emotionally relevant cues.     

Transient interference of right hemispheric function due to automatic emotional processing

We examined the effects of emotional stimuli on right and left hemisphere detection performance in a hemifield visual discrimination task. A group of 18 healthy subjects were asked to discriminate between upright and inverted triangles (target). Targets were randomly presented in the left or right visual hemifield (150 ms target duration). A brief emotional picture (pleasant or unpleasant; 150 ms stimulus duration) or neutral picture selected from the International Affective Picture System was randomly presented either in the same (47%) or the opposite (47%) spatial location to the subsequent target. Emotional or neutral stimuli offset 150 ms prior to the subsequent target. Subjects were instructed to ignore the pictures and respond to the targets as quickly and accurately as possible. Independent of field of presentation, emotional stimuli prolonged reaction times (P < 0.01) to LVF targets, with unpleasant stimuli showing a greater effect than pleasant stimuli. The current study shows that brief emotional stimuli selectively impair right hemispheric visual discrimination capacity. The findings suggest automatic processing of emotional stimuli captures right hemispheric processing resources and transiently interferes with other right hemispheric functions.     

Slow biasing of processing resources in early visual cortex is preceded by emotional cue extraction in emotion–attention competition

In our previous studies on competition for attentional processing resources in early visual cortex between a foreground task and distracting emotional background images we found that emotional background images withdraw attentional resources from the foreground task after about 400 ms. Costs in behavioral data and a significant reduction of the steady state visual evoked potential (SSVEP) amplitude that was elicited by the foreground task lasted for several hundred milliseconds. We speculated that the differential effect in SSVEP amplitudes is preceded by the extraction of the emotional cue. Event related potential (ERP) studies to emotional and neutral complex images identified an early posterior negativity (EPN) as a robust neural signature of emotional cue extraction. The late positive potential (LPP) was related to in‐depth processing of the emotional image. We extracted ERPs that were evoked by the onset of background images concurrently with the SSVEP that was elicited by the foreground task. Emotional compared to neutral background pictures evoked a more negative EPN at about 190 ms and a more positive LPP at about 700 ms after image onset. SSVEP amplitudes became significantly smaller with emotional background images after about 400 ms lasting for several hundred ms. Interestingly, we found no significant correlations between the three components, indicating that they act independently. Source localizations resulted in nonoverlapping cortical generators. Results suggest a cascade of perceptual processes: Extraction of the emotional cue preceded biasing of attentional resources away from the foreground task towards the emotional image for an evaluation of the picture content. Hum Brain Mapp 35:1477–1490, 2014. © 2013 Wiley Periodicals, Inc.     

Stimulus novelty and emotion perception: the near absence of habituation in the visual cortex

In rapid serial visual presentation of pictures, an early event-related brain potential component shows enlarged negativity over occipital regions for emotional pictures compared with neutral pictures. The present study examined whether the processing of emotional target pictures varies as a function of stimulus repetition. Accordingly, pictures of erotica, neutral contents, and mutilations were repeatedly presented (90 times) while the electroencephalogram was recorded with a 129 dense sensor array. As in previous studies, emotional pictures were associated with a larger posterior negativity than neutral pictures. Furthermore, differential emotion processing did not vary as a function of stimulus repetition and was similarly expressed across blocks of picture presentation. These findings suggest the near absence of habituation in differential emotion processing during perceptual processing.     

Intact motivated attention in schizophrenia: evidence from event-related potentials

Emotionally significant stimuli typically capture attention (called motivated attention) even when they are irrelevant to tasks where attention is directed. Previous studies indicate that several components of emotional processing are intact in schizophrenia when subjects are instructed to attend to emotionally-evocative stimuli. However, few studies have examined whether emotional stimuli capture attention to a normal degree in people with schizophrenia when attention is directed elsewhere. The current event-related potential study examined motivated attention to task-irrelevant emotional stimuli in 35 stabilized outpatients and 26 healthy controls with a modified visual P300 oddball detection task. Participants viewed images of rare target and commonly occurring standard letter stimuli, as well as intermixed emotional (unpleasant, pleasant, neutral) pictures. Subjects were instructed to count the number of rare targets; the emotional valence of the picture stimuli was, therefore, task-irrelevant. We separately evaluated the Early Posterior Negativity (EPN) and Late Positive Potential (LPP) to emotional pictures and the P300 to target stimuli. Patients and controls showed similar patterns of EPN and LPP amplitude to the emotional stimuli, such that the EPN and LPP were larger for both pleasant and unpleasant versus neutral pictures. Although patients performed worse than controls on the target counting task, both groups showed comparable P300 differentiation between target versus non-target stimuli. Emotional stimuli captured attentional resources in people with schizophrenia even when the emotional stimuli were task-irrelevant, suggesting intact motivated attention at the level of early electrophysiological responding.     

Orbitofrontal cortex biases attention to emotional events

We examined the role of orbitofrontal (OF) cortex in regulating emotion-attention interaction and the balance between involuntary and voluntary attention allocation. We studied patients with OF lesion applying reaction time (RT) and event-related potential (ERP) measures in a lateralized visual discrimination task with novel task-irrelevant affective pictures (unpleasant, pleasant, or neutral) preceding a neutral target. This allowed for comparing the effects of automatic attention allocation to emotional versus neutral stimuli on subsequent voluntary attention allocation to target stimuli. N2-P3a and N2-P3b ERP components served as measures of involuntary and voluntary attention allocation correspondingly. Enhanced N2-P3a amplitudes to emotional distractors and reduced N2-P3b amplitudes to targets preceded by emotional distractors were observed in healthy subjects, suggesting automatic emotional orienting interfered with subsequent voluntary orienting. OF patients showed an opposite pattern with tendency towards reduced N2-P3a responses to emotional distractors, suggesting impaired automatic orienting to emotional stimuli due to orbitofrontal damage. Enhanced N2-P3b responses to targets preceded by any affective distractor were observed in OF patients, suggesting bias towards voluntary target-related attention allocation due to orbitofrontal lesion. Behavioral evidence indicated that left visual field (LVF) attention performance was modulated by emotional stimuli. Specifically, OF patients responded faster to LVF targets subsequent to pleasant emotional distractors. We suggest that damage to the orbitofrontal circuitry leads to dysbalance between voluntary and involuntary attention allocation in the context of affective distractors with predisposition to posterior target-related processing over frontal novelty and affect-related processing. Furthermore, we suggest that orbitofrontal influence on emotion-attention interaction is valence and hemisphere dependent.     

Fleeting images: rapid affect discrimination in the visual cortex

Converging electrophysiological and hemodynamic findings indicate sensory processing of emotional pictures is preferred to that of neutral pictures. Whereas neuroimaging studies of emotional picture perception have employed stimulus durations lasting several seconds, recent electrocortical investigations report early visual cortical discrimination between emotionally arousing and neutral picture processing. Here, we use a hybrid picture presentation paradigm covering a range of rapid presentation rates (0.75-6 Hz), while visual system activity is recorded with functional magnetic resonance imaging. Results demonstrate widespread sensitivity to emotional arousal in the secondary and inferior temporal visual cortex. Furthermore, activity in the lateral inferior occipital and medial inferior temporal visual cortex revealed equivalent emotion-sensitive activation across all presentation rates. Results further support the notion that attention and perceptual processing are in part directed by underlying motivational systems.     

Oxytocin increases recognition of masked emotional faces

The neuropeptide oxytocin has been shown to improve many aspects of social cognitive functioning, including facial emotion recognition, and to promote social approach behaviour. In the present study, we investigated the modulatory effects of oxytocin on the recognition of briefly presented facial expressions. In order to diversify the degree of visual awareness for the facial stimuli, presentation duration was systematically varied. Fifty-six participants were administered intranasal oxytocin or a placebo in a double-blind, randomized, between-subjects design. Participants viewed angry and happy target faces or neutral distractors for 18, 35, or 53 ms subsequently masked by neutral faces. Participants had to indicate the presence or absence of the briefly presented target face. Discrimination indices (d') showed that oxytocin generally enhanced detection accuracy of emotional stimuli. This effect was more pronounced for the recognition of happy faces. We provide evidence that a single dose of intranasally administered oxytocin enhances detection of briefly presented emotional stimuli. The possible role of stimulus valence and recognition difficulty is discussed.     

Spatiotemporal dynamics of affective picture processing revealed by intracranial high‐gamma modulations

Our comprehension of the neural mechanisms underlying emotional information processing has largely benefited from noninvasive electrophysiological and functional neuroimaging techniques in recent years. However, the spatiotemporal dynamics of the neural events occurring during emotional processing remain imprecise due to the limited combination of spatial and temporal resolution provided by these techniques. This study examines the modulations of high‐frequency activity of intracranial electroencephalography recordings associated with affective picture valence, in epileptic patients awaiting neurosurgery. Recordings were obtained from subdural grids and depth electrodes in eight patients while they viewed a series of unpleasant, pleasant and neutral pictures from the International Affective Picture System. Broadband high‐gamma (70–150 Hz) power was computed for separate 100‐ms time windows and compared according to ratings of emotional valence. Compared to emotionally neutral or pleasant pictures, unpleasant stimuli were associated with an early and long‐lasting (≈200–1,000 ms) bilateral increase in high‐gamma activity in visual areas of the occipital and temporal lobes, together with a late and transient (≈500–800 ms) decrease found bilaterally in the lateral prefrontal cortex (PFC). Pleasant pictures were associated with increased gamma activity in the occipital cortex, compared to the emotionally neutral stimuli. Consistent with previous studies, our results provide direct evidence of emotion‐related modulations in the visual ventral pathway during picture processing. Results in the lateral PFC also shed light on the neural mechanisms underlying its role in negative emotions processing. This study demonstrates the utility of intracranial high‐gamma modulations to study emotional process with a high spatiotemporal precision. Hum Brain Mapp, 36:16–28, 2015.. © 2014 Wiley Periodicals, Inc.     

Attentional load affects automatic emotional processing: evidence from event-related potentials

One open question on the relation between attention and emotion concerns the automatic processing of emotional visual stimuli outside the focus of attention. This study examined to what extent the emotional processing at unattended locations is modulated by the processing load at attended locations. Event-related potentials were measured to task-irrelevant unpleasant and neutral pictures briefly presented at peripheral locations while participants performed a visual central task varying in load (low and high load). Unpleasant pictures elicited larger amplitudes of N1-P2 at parietoccipital and occipital sites than that of neutral pictures. This effect was only significant in the low-load condition. Data suggest that brain responses to affective value of task-irrelevant peripheral pictures are modulated by attentional load at fixation.     

Rapid discrimination of visual scene content in the human brain

The rapid evaluation of complex visual environments is critical for an organism's adaptation and survival. Previous studies have shown that emotionally significant visual scenes, both pleasant and unpleasant, elicit a larger late positive wave in the event-related brain potential (ERP) than emotionally neutral pictures. The purpose of the present study was to examine whether neuroelectric responses elicited by complex pictures discriminate between specific, biologically relevant contents of the visual scene and to determine how early in the picture processing this discrimination occurs. Subjects (n = 264) viewed 55 color slides differing in both scene content and emotional significance. No categorical judgments or responses were required. Consistent with previous studies, we found that emotionally arousing pictures, regardless of their content, produce a larger late positive wave than neutral pictures. However, when pictures were further categorized by content, anterior ERP components in a time window between 200 and 600 ms following stimulus onset showed a high selectivity for pictures with erotic content compared to other pictures regardless of their emotional valence (pleasant, neutral, and unpleasant) or emotional arousal. The divergence of ERPs elicited by erotic and non-erotic contents started at 185 ms post-stimulus in the fronto-central midline region, with a later onset in parietal regions. This rapid, selective, and content-specific processing of erotic materials and its dissociation from other pictures (including emotionally positive pictures) suggests the existence of a specialized neural network for prioritized processing of a distinct category of biologically relevant stimuli with high adaptive and evolutionary significance.     

Effects of emotional arousal in the cerebral hemispheres: a study of oscillatory brain activity and event-related potentials.

OBJECTIVE: The present study aimed at examining the time course and topography of oscillatory brain activity and event-related potentials (ERPs) in response to laterally presented affective pictures. METHODS: Electroencephalography was recorded from 129 electrodes in 10 healthy university students during presentation of pictures from the international affective picture system. Frequency measures and ERPs were obtained for pleasant, neutral, and unpleasant pictures. RESULTS: In accordance with previous reports, a modulation of the late positive ERP wave at parietal recording sites was found as a function of emotional arousal. Early mid gamma band activity (GBA; 30-45 Hz) at 80 ms post-stimulus was enhanced in response to aversive stimuli only, whereas the higher GBA (46-65 Hz) at 500 ms showed an enhancement of arousing, compared to neutral pictures. ERP and late gamma effects showed a pronounced right-hemisphere preponderance, but differed in terms of topographical distribution. CONCLUSIONS: Late gamma activity may represent a correlate of widespread cortical networks processing different aspects of emotionally arousing visual objects. In contrast, differences between affective categories in early gamma activity might reflect fast detection of aversive stimulus features.     

Snake fearfulness is associated with sustained competitive biases to visual snake features: Hypervigilance without avoidance

The extent and time course of competition between a specific fear cue and task-related stimuli in early human visual cortex was investigated using electrophysiology. Steady-state visual evoked potentials (ssVEPs) were evoked using random-dot kinematograms that consisted of rapidly flickering (8.57 Hz) dots moving randomly, superimposed upon emotional or neutral distractor pictures. Participants were asked to detect intervals of coherently moving dots, ignoring the distractor pictures that varied in hedonic content. Women reporting high or low levels of snake fear were recruited from a large sample of healthy college students, and snake pictures served as fear-relevant distractors. The time-varying amplitude of the ssVEP evoked by the motion detection task showed significant reduction when viewing emotionally arousing, compared to neutral, distractors, replicating previous studies. For high-fear participants, snake distractors elicited a sustained attenuation of task evoked ssVEP amplitude, greater than the attenuation prompted by other unpleasant arousing content. These findings support a hypothesis that fear cues prompt sustained hypervigilance rather than perceptual avoidance.     

Cross-modal emotional attention: emotional voices modulate early stages of visual processing

Emotional attention, the boosting of the processing of emotionally relevant stimuli, has, up to now, mainly been investigated within a sensory modality, for instance, by using emotional pictures to modulate visual attention. In real-life environments, however, humans typically encounter simultaneous input to several different senses, such as vision and audition. As multiple signals entering different channels might originate from a common, emotionally relevant source, the prioritization of emotional stimuli should be able to operate across modalities. In this study, we explored cross-modal emotional attention. Spatially localized utterances with emotional and neutral prosody served as cues for a visually presented target in a cross-modal dot-probe task. Participants were faster to respond to targets that appeared at the spatial location of emotional compared to neutral prosody. Event-related brain potentials revealed emotional modulation of early visual target processing at the level of the P1 component, with neural sources in the striate visual cortex being more active for targets that appeared at the spatial location of emotional compared to neutral prosody. These effects were not found using synthesized control sounds matched for mean fundamental frequency and amplitude envelope. These results show that emotional attention can operate across sensory modalities by boosting early sensory stages of processing, thus facilitating the multimodal assessment of emotionally relevant stimuli in the environment.     

情绪词汇认知加工诱发电位时空模式的脑功能研究

目的应用多通道事件相关电位(event-related potential,ERP)分析技术,探讨正常青年受试者对情绪性词汇认知加工脑神经功能电活动机制。方法应用不同情绪效价的情绪刺激图片对31例健康志愿者进行情绪效价5个水平情绪脑电诱发电位的测试判别,观察受试者对5个情绪效价水平情绪词汇的情绪反应。结果 ERPs统计参数映像图(SPM)结果,对词汇情绪性加工的显著性差异最早可见于刺激后60~200ms;中后期(刺激后420~960ms)极负性和中性情绪反应显著性差异表现为右腹侧记忆-注意整合网络出现刺激负荷效应。极正性和中性情绪反应差异为中期(刺激后320~420ms)的左顶、颞、枕,晚期(刺激后680~1 000ms)见到右额中央负荷效应。极正性和极负性情绪反应差异于刺激后400~1 000ms见于双侧眶额皮层、腹内侧前额皮层、前扣带回皮层和躯体感觉皮层等脑区的广泛记忆-注意整合网络引起更强的激活。结论个体认知行为差异和社会环境背景影响不同情绪效价情绪性词汇的加工,不同情绪效价的情绪性词汇加工过程中存在显著性差异。正负性情绪词汇的加工是双侧半球广泛脑区的记忆-注意整合网络激活效应,并存在脑的偏侧化激活效应。     

Neural processing stages during object-substitution masking and their relationship to perceptual awareness

The extent of visual perceptual processing that occurs in the absence of awareness is as yet unclear. Here we examined event-related-potential (ERP) indices of visual and cognitive processes as awareness was manipulated through object-substitution masking (OSM), an awareness-disrupting effect that has been hypothesized to result from the disruption of reentrant signaling to low-level visual cortical areas. In OSM, a visual stimulus array is briefly presented that includes a parafoveal visual target denoted by a cue, typically consisting of several surrounding dots. When the offset of the target-surrounding cue dots is delayed relative to the rest of the array, a striking reduction in the perception of the target image surrounded by the dots is observed. Using faces and houses as the target stimuli, we found that successful OSM reduced or eliminated all the measured electrophysiological indices of visual processing stages after 130 ms post-stimulus. More specifically, when targets were missed within the masked condition (i.e., on trials with effective OSM that disrupted awareness), we observed fully intact early feed-forward processing up through the visual extrastriate P1 ERP component peaking at 100 ms, followed by reduced low-level activity over the occipital pole 130–170 ms post-stimulus, reduced ERP indices of lateralized shifts of attention toward the parafoveal target, reduced object-generic visual processing, abolished object-category-specific (face-specific) processing, and reduced late visual short-term-memory processing activity. The results provide a comprehensive electrophysiological account of the neurocognitive underpinnings of effective OSM of visual-object images, including evidence for central roles of early reentrant signal disruption and insufficient visual attentional deployment.     

Threat interferes with response inhibition

A potential threat, such as a spider, captures attention and engages executive functions to adjust ongoing behavior and avoid danger. We and many others have reported slowed responses to neutral targets in the context of emotional distractors. This behavioral slowing has been explained in the framework of attentional competition for limited resources with emotional stimuli prioritized. Alternatively, slowed performance could reflect the activation of avoidance/freezing-type motor behaviors associated with threat. Although the interaction of attention and emotion has been widely studied, little is known on the interaction between emotion and executive functions. We studied how threat-related stimuli (spiders) interact with executive performance and whether the interaction profile fits with a resource competition model or avoidance/freezing-type motor behaviors. Twenty-one young healthy individuals performed a Go-NoGo visual discrimination reaction time (RT) task engaging several executive functions with threat-related and emotionally neutral distractors. The threat-related distractors had no effect on the RT or the error rate in the Go trials. The NoGo error rate, reflecting failure in response inhibition, increased significantly because of threat-related distractors in contrast to neutral distractors, P less than 0.05. Thus, threat-related distractors temporarily impaired response inhibition. Threat-related distractors associated with increased commission errors and no effect on RT does not suggest engagement of avoidance/freezing-type motor behaviors. The results fit in the framework of the resource competition model. A potential threat calls for evaluation of affective significance as well as inhibition of undue emotional reactivity. We suggest that these functions tax executive resources and may render other executive functions, such as response inhibition, temporarily compromised when the demands for resources exceed availability.     

Affective picture perception: gender differences in visual cortex?

Activity in extrastriate visual cortex is greater when people view emotional relative to neutral pictures. Prior brain imaging and psychophysiological work has further suggested a bias for men to react more strongly to pleasant pictures, and for women to react more strongly to unpleasant pictures. Here we investigated visual cortical activity using fMRI in 28 men and women during picture viewing. Men and women showed reliably greater visual cortical reactivity during both pleasant and unpleasant pictures, relative to neutral, consistent with the view that the motivational relevance of visual stimuli directs attention and enhances elaborative perceptual processing. However, men did show greater extrastriate activity than women specifically during erotic picture perception, possibly reflecting a gender-specific visual mechanism for sexual selection.     

Emotion processing in Parkinson's disease: dissociation between early neuronal processing and explicit ratings.

OBJECTIVE: Patients suffering from Parkinson's disease (PD) have a diminished ability to discriminate facial expressions of emotion. We investigated early emotion discrimination deficits in PD by means of event-related potentials (ERPs). METHODS: Emotional pictures were presented to 14 PD patients and 14 healthy controls in a rapid serial visual presentation paradigm (three frames per second) while EEG was recorded. In addition, valence and arousal ratings were obtained for a representative subsample of 54 pictures. RESULTS: PD patients rated pictures of highly arousing content as less exciting than did healthy controls. Pictures of high compared to low emotional arousal were associated with a pronounced relative negative shift in the ERP waveform over parietal and occipital sites developing about 220 ms after picture onset. This early posterior negativity (EPN) did not differ between PD and control group. CONCLUSIONS: This dissociation of affective ratings and early ERP components supports the view that PD is associated with blunted emotional responses, but there is no evidence for a deteriorated early visual processing of emotional stimuli. SIGNIFICANCE: Frequently reported deficits in emotion discrimination are likely not due to deficits in early emotion processing.     

Emotionally negative stimuli can overcome attentional deficits in patients with visuo-spatial hemineglect

Left unilateral spatial neglect resulting from right brain damage is characterized by loss of awareness for stimuli in the contralesional side of space, despite intact visual pathways. We examined using fMRI whether patients with neglect are more likely to consciously detect in the neglected hemifield, emotionally negative complex scenes rather than visually similar neutral pictures and if so, what neural mechanisms mediate this effect. Photographs of emotional and neutral scenes taken from the IAPS were presented in a divided visual field paradigm. As expected, the detection rate for emotional stimuli presented in the neglected field was higher than for neutral ones. Successful detection of emotional scenes as opposed to neutral stimuli in the left visual field (LVF) produced activations in the parahippocampal and anterior cingulate areas in the right hemisphere. Detection of emotional stimuli presented in the intact right visual field (RVF) activated a distributed network of structures in the left hemisphere, including anterior and posterior cingulate cortex, insula, as well as visual striate and extrastriate areas. LVF-RVF contrasts for emotional stimuli revealed activations in right and left attention related prefrontal areas whereas RVF-LVF comparison showed activations in the posterior cingulate and extrastriate visual cortex in the left hemisphere. An additional analysis contrasting detected vs. undetected emotional LVF stimuli showed involvement of left anterior cingulate, right frontal and extrastriate areas. We hypothesize that beneficial role of emotion in overcoming neglect is achieved by activation of frontal and limbic lobe networks, which provide a privileged access of emotional stimuli to attention by top-down modulation of processing in the higher-order extrastriate visual areas. Our results point to the importance of top-down regulatory role of the frontal attentional systems, which might enhance visual activations and lead to greater salience of emotional stimuli for perceptual awareness.