Emotionally arousing stimuli compete for attention with left hemispace

Rapid interaction of the emotional and attentional networks is critical for adaptive behavior. Here, we examined the effects of emotional stimulation on hemifield attention allocation using event-related potential and behavioral measures. Participants performed a visual-discrimination task on nonemotional targets presented randomly in the left or right hemifield. A brief task-irrelevant emotional (pleasant or unpleasant; 150-ms duration) or neutral picture was presented centrally 350 ms before the next target (150-ms duration). Unpleasant stimuli interfered with the left visual field attention capacity, slowing behavioral responses to attended left field stimuli. In keeping with the behavioral data, event-related potential responses to nonemotional attended left field stimuli were reduced over the right parietal regions when preceded by an unpleasant event. The results provide electrophysiological and behavioral evidence that unpleasant, emotionally arousing stimuli interfere with the right hemisphere-dependent attention capacity.     

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.     

Cerebral blood flow changes associated with attribution of emotional valence to pleasant, unpleasant, and neutral visual stimuli in a PET study of normal subjects.

OBJECTIVE: To assist in the development of a model for the psychopathology of emotions, the present study sought to identify the neural circuits associated with the evaluation of visual stimuli for emotional valence. METHOD: Seventeen healthy individuals were shown three sets of emotionally laden pictures carrying pleasant, unpleasant, and neutral content. While subjects evaluated the picture set for emotional valence, regional cerebral blood flow was measured with the use of [15O] water positron emission tomography. Subjective ratings of the emotional valence of the picture sets were recorded. Data were analyzed by comparing the images acquired during the neutral condition with the unpleasant and pleasant image sets and the unpleasant and pleasant conditions with each other. RESULTS: Processing of pleasant stimuli was associated with increased blood flow in the dorsal-lateral, orbital, and medial frontal cortex relative to the unpleasant condition and in the cingulate, precuneus, and visual cortex relative to the neutral condition. Evaluation of unpleasant stimuli activated the amygdala, visual cortex, and cerebellum relative to the pleasant condition and the nucleus accumbens, precuneus, and visual cortex relative to the neutral condition. CONCLUSIONS: Observing and assigning emotional value to unpleasant stimuli produced activations in subcortical limbic regions, whereas evaluation of pleasant stimuli produced activations in cortical limbic areas. These findings are consistent with the notion of a subcortical and archaic danger recognition system and a system detecting pleasantness in events and situations that is phylogenetically younger, involving primarily the prefrontal cortex.     

TMS to V1 spares discrimination of emotive relative to neutral body postures

This study used TMS to examine the role played by striate cortex (V1) in processing the emotional content of visual stimuli. Participants learned to discriminate two sets of body posture images. For half of each set, the posture's emotional significance (threat versus pleasant) provided a redundant cue for the discrimination; the other half were emotionally neutral. An image was briefly presented at a lateral location in the visual field where a TMS pulse produced a phosphene, or at a control location in the opposite hemifield. A TMS pulse 70–140 ms after stimulus presentation at the phosphene location impaired discrimination of neutral stimuli with little effect on discrimination of emotional stimuli; the two classes of stimuli were equally discriminable when presented at the control location. The results are consistent with the proposal that emotionally salient patterns, such as social threat, can be discriminated independently of the geniculo-striate pathway.     

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.     

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.     

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.     

Brain activity during expectancy of emotional stimuli: an fMRI study

We studied the neural activation associated with the expectancy of emotional stimuli using whole brain fMRI. Fifteen healthy subjects underwent fMRI scanning during which they performed a warned reaction task using emotional pictures carrying pleasant, unpleasant, or neutral content. The task involved an expected or unexpected condition. Data were analyzed by comparing the images acquired under the different conditions. In the expected condition, compared with the unexpected condition, significant activation was observed in the medial, inferior and dorsolateral prefrontal cortex. Whereas the expectancy of pleasant stimuli produced activation in the left dorsolateral and left medial prefrontal cortex as well as in the right cerebellum, the expectancy of unpleasant stimuli produced activation in the right inferior and right medial prefrontal cortex, the right amygdala, the left anterior cingulate cortex, and bilaterally in the visual cortex. These results suggest that the expectancy of emotional stimuli is mediated by the prefrontal area including the medial, inferior, and dorsolateral prefrontal cortex. Furthermore, our data suggest that left frontal activation is associated with the expectancy of pleasant stimuli and that right frontal activation is associated with the expectancy of unpleasant stimuli. Finally, our findings suggest that the amygdala and anterior cingulate cortex may play an important role in the expectancy of unpleasant stimuli and that the input of this negative information is modulated by these specific brain areas.     

His or mine? The time course of self-other discrimination in emotion processing

This electroencephalography (EEG) study investigated at which temporal processing stages self-other discrimination in emotion processing occurs. EEG was recorded in 23 healthy participants during silent reading of unpleasant, pleasant, and neutral pronoun-noun and article--noun expressions that were related to the participants themselves, related to an unknown third person, or had no self-other reference at all. Self- and other-related pronoun--noun pairs elicited larger cortical negativity relative to the processing of article--noun pairs at left posterior electrodes as early as 200 ms after stimulus onset. In the same time windows (from 200 ms to 300 ms and 300 ms to 400 ms) the emotionality of the words enhanced event-related brain potential (ERP) amplitudes at parieto-occipital electrodes. From 350 ms onwards, processing of self-related unpleasant words elicited larger frontal negativity compared to unpleasant words that were related to the other or that had no reference at all. In addition, processing of pleasant words vs. neutral or unpleasant words elicited larger positive amplitudes over parietal electrodes from 450 ms after stimulus onset, in particular when words were self-related. Our findings demonstrate that for verbal emotional stimuli, self--other discrimination first occurs at higher-order, cortical processing stages. This is consistent with the view that categorization of information according to certain stimulus aspects (self--other reference, emotionality) occurs before its meaning is integrated.     

Functional neurological symptoms modulate processing of emotionally salient stimuli

ObjectiveDysfunctional emotion processing has been discussed as a contributing factor to functional neurological symptoms (FNS) in the context of conversion disorder, and refers to blunted recognition and the expression of one's own feelings. However, the emotion processing components characteristic for FNS and/or relevant for conversion remain to be specified. With this goal, the present study targeted the initial, automatic discrimination of emotionally salient stimuli.MethodsThe magnetoencephalogram (MEG) was monitored in 21 patients with functional weakness and/or sensory disturbance subtypes of FNS and 21 healthy comparison participants (HC) while they passively watched 600 emotionally arousing, pleasant, unpleasant or neutral stimuli in a rapid serial visual presentation (RSVP) design. Neuromagnetic activity was analyzed 110–330 ms following picture onset in source space for prior defined posterior and central regions of interest.ResultsAs early as 110 ms and across presentation interval, posterior neural activity modulation by picture category was similar in both groups, despite smaller initial (110–150 ms) overall and posterior power in patients with FNS. The initial activity modulation by picture category was also evident in the left sensorimotor area in patients with FNS, but not significant in HC.ConclusionsSimilar activity modulation by emotional picture category in patients with FNS and HC suggests that the fast, automatic detection of emotional salience is unchanged in patients with FNS, but involves an emotion-processing network spanning posterior and sensorimotor areas.     

Amygdala activation during reading of emotional adjectives--an advantage for pleasant content

This event-related functional magnetic resonance imaging (fMRI)study investigated brain activity elicited by emotional adjectivesduring silent reading without specific processing instructions.Fifteen healthy volunteers were asked to read a set of randomlypresented high-arousing emotional (pleasant and unpleasant)and low-arousing neutral adjectives. Silent reading of emotionalin contrast to neutral adjectives evoked enhanced activationsin visual, limbic and prefrontal brain regions. In particular,reading pleasant adjectives produced a more robust activationpattern in the left amygdala and the left extrastriate visualcortex than did reading unpleasant or neutral adjectives. Moreover,extrastriate visual cortex and amygdala activity were significantlycorrelated during reading of pleasant adjectives. Furthermore,pleasant adjectives were better remembered than unpleasant andneutral adjectives in a surprise free recall test conductedafter scanning. Thus, visual processing was biased towards pleasantwords and involved the amygdala, underscoring recent theoreticalviews of a general role of the human amygdala in relevance detectionfor both pleasant and unpleasant stimuli. Results indicate preferentialprocessing of pleasant information in healthy young adults andcan be accounted for within the framework of appraisal theory.     

Do brain responses to emotional images and cigarette cues differ? An fMRI study in smokers

Chronic smoking is thought to cause changes in brain reward systems that result in overvaluation of cigarette-related stimuli and undervaluation of natural rewards. We tested the hypotheses that, in smokers, brain circuits involved in emotional processing: (i) would be more active during exposure to cigarette-related than neutral pictures; and (ii) would be less active to pleasant compared with cigarette-related pictures, suggesting a devaluation of intrinsically pleasant stimuli. We obtained whole-brain blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging data from 35 smokers during the presentation of pleasant (erotica and romance), unpleasant (mutilations and sad), neutral, and cigarette-related pictures. Whole-brain analyses showed significantly larger BOLD responses during presentation of cigarette-related pictures relative to neutral ones within the secondary visual areas, the cingulate gyrus, the frontal gyrus, the dorsal striatum, and the left insula. BOLD responses to erotic pictures exceeded responses to cigarette-related pictures in all clusters except the insula. Within the left insula we observed larger BOLD responses to cigarette-related pictures than to all other picture categories. By including intrinsically pleasant and unpleasant pictures in addition to neutral ones, we were able to conclude that the presentation of cigarette-related pictures activates brain areas supporting emotional processes, but we did not find evidence of overall reduced activation of the brain reward systems in the presence of intrinsically pleasant stimuli.     

“Emotions Guide Us”: Behavioral and MEG correlates

Affectively salient stimuli are capable of capturing attentional resources which allow the brain to change the current course of action in order to respond to potentially advantageous or threatening stimuli. Here, we investigated the behavioral and cerebral impact of peripherally presented affective stimuli on the subsequent processing of foveal information. To this end, we recorded whole-head magnetoencephalograms from 12 participants while they made speeded responses to the direction of left- or right-oriented arrows that were presented foveally at fixation. Each arrow was preceded by a peripherally presented pair of pictures, one emotional (unpleasant or pleasant), and one neutral. Paired pictures were presented at 12° of eccentricity to the left and right of a central fixation cross. We observed that the participants responded more quickly when the orientation of the arrow was congruent with the location of the previously presented emotional scene. Results show that non-predictive emotional information in peripheral vision interferes with subsequent responses to foveally presented targets. Importantly, this behavioral effect was correlated with an early (∼135 msec) increase of left fronto-central activity for the emotionally congruent combination, whose cerebral sources were notably located in the left orbitofrontal cortex. We therefore suggest that the prior spatial distribution of emotional salience, like physical salience, grabs attentional resources and modifies the performance in the center of the visual field. Thus, these data shed light on the neurobehavioral correlates of the emotional coding of visual space.     

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.     

Tagging cortical networks in emotion: A topographical analysis

Viewing emotional pictures is associated with heightened perception and attention, indexed by a relative increase in visual cortical activity. Visual cortical modulation by emotion is hypothesized to reflect re‐entrant connectivity originating in higher‐order cortical and/or limbic structures. The present study used dense‐array electroencephalography and individual brain anatomy to investigate functional coupling between the visual cortex and other cortical areas during affective picture viewing. Participants viewed pleasant, neutral, and unpleasant pictures that flickered at a rate of 10 Hz to evoke steady‐state visual evoked potentials (ssVEPs) in the EEG. The spectral power of ssVEPs was quantified using Fourier transform, and cortical sources were estimated using beamformer spatial filters based on individual structural magnetic resonance images. In addition to lower‐tier visual cortex, a network of occipito‐temporal and parietal (bilateral precuneus, inferior parietal lobules) structures showed enhanced ssVEP power when participants viewed emotional (either pleasant or unpleasant), compared to neutral pictures. Functional coupling during emotional processing was enhanced between the bilateral occipital poles and a network of temporal (left middle/inferior temporal gyrus), parietal (bilateral parietal lobules), and frontal (left middle/inferior frontal gyrus) structures. These results converge with findings from hemodynamic analyses of emotional picture viewing and suggest that viewing emotionally engaging stimuli is associated with the formation of functional links between visual cortex and the cortical regions underlying attention modulation and preparation for action. Hum Brain Mapp, 2012. © 2011 Wiley Periodicals, Inc.     

His or mine? The time course of self–other discrimination in emotion processing

This electroencephalography (EEG) study investigated at which temporal processing stages self–other discrimination in emotion processing occurs. EEG was recorded in 23 healthy participants during silent reading of unpleasant, pleasant, and neutral pronoun–noun and article–noun expressions that were related to the participants themselves, related to an unknown third person, or had no self–other reference at all. Self- and other-related pronoun–noun pairs elicited larger cortical negativity relative to the processing of article–noun pairs at left posterior electrodes as early as 200 ms after stimulus onset. In the same time windows (from 200 ms to 300 ms and 300 ms to 400 ms) the emotionality of the words enhanced event-related brain potential (ERP) amplitudes at parieto-occipital electrodes. From 350 ms onwards, processing of self-related unpleasant words elicited larger frontal negativity compared to unpleasant words that were related to the other or that had no reference at all. In addition, processing of pleasant words vs. neutral or unpleasant words elicited larger positive amplitudes over parietal electrodes from 450 ms after stimulus onset, in particular when words were self-related. Our findings demonstrate that for verbal emotional stimuli, self–other discrimination first occurs at higher-order, cortical processing stages. This is consistent with the view that categorization of information according to certain stimulus aspects (self–other reference, emotionality) occurs before its meaning is integrated.     

Behavioral and electrophysiological evidence of a right hemisphere bias for the influence of negative emotion on higher cognition

We examined how responses to aversive pictures affected performance and stimulus-locked event-related potentials (ERPs) recorded during a demanding cognitive task. Numeric Stroop stimuli were brief ly presented to either left or right visual hemifield (LVF and RVF, respectively) after a centrally presented aversive or neutral picture from the International Affective Picture System. Subjects indicated whether a quantity value from each Stroop stimulus matched the preceding Stroop stimulus while passively viewing the pictures. After aversive pictures, responses were more accurate for LVF Stroops and less accurate for RVF Stroops. Early-latency extrastriate attention-dependent visual ERPs were enhanced for LVF Stroops. The N2 ERP was enhanced for LVF Stroops over the right frontal and parietal scalp sites. Slow potentials (300-800 msec) recorded over the frontal and parietal regions showed enhanced picture related modulation and amplitude for LVF Stroops. These results suggest that emotional responses to aversive pictures selectively facilitated right hemisphere processing during higher cognitive task performance.     

Expecting unpleasant stimuli – An fMRI study

Expecting forthcoming events and preparing adequate responses are important cognitive functions that help the individual to deal with the environment. The emotional valence of an event is decisive for the resulting action. Revealing the underlying mechanisms may help to understand the dysfunctional information processing in depression and anxiety that are associated with negative expectation of the future. We were interested in selective brain activity during the expectation of unpleasant visual stimuli. Twelve healthy female subjects were biased to expect and then perceive emotionally unpleasant, pleasant or neutral stimuli during functional magnetic resonance imaging. Expecting unpleasant stimuli relative to expecting pleasant and neutral stimuli resulted in activation of mainly cingulate cortex, insula, prefrontal areas, thalamus, hypothalamus and striatum. While certain areas were also active during subsequent presentation of the emotional stimuli, distinct regions of the anterior cingulate gyrus and the thalamus were solely active during expectation of the unpleasant stimuli. The identified areas may reflect a network for internal adaptation and preparation processes in order to react adequately to expected unpleasant events. They are known as well to be altered in depression. Disorders of this network may be relevant for psychiatric disorders such as depression.     

Expecting unpleasant stimuli--an fMRI study

Expecting forthcoming events and preparing adequate responses are important cognitive functions that help the individual to deal with the environment. The emotional valence of an event is decisive for the resulting action. Revealing the underlying mechanisms may help to understand the dysfunctional information processing in depression and anxiety that are associated with negative expectation of the future. We were interested in selective brain activity during the expectation of unpleasant visual stimuli. Twelve healthy female subjects were biased to expect and then perceive emotionally unpleasant, pleasant or neutral stimuli during functional magnetic resonance imaging. Expecting unpleasant stimuli relative to expecting pleasant and neutral stimuli resulted in activation of mainly cingulate cortex, insula, prefrontal areas, thalamus, hypothalamus and striatum. While certain areas were also active during subsequent presentation of the emotional stimuli, distinct regions of the anterior cingulate gyrus and the thalamus were solely active during expectation of the unpleasant stimuli. The identified areas may reflect a network for internal adaptation and preparation processes in order to react adequately to expected unpleasant events. They are known as well to be altered in depression. Disorders of this network may be relevant for psychiatric disorders such as depression.     

Hemispheric asymmetries for the conscious and unconscious perception of emotional words

The current research examines the interactions between hemispheric asymmetries for visual perception and emotion. In a series of four experiments, participants completed tasks measuring both conscious and unconscious perception of linguistic stimuli. In these studies, stimulus-presentation parameters (brief exposure duration vs masking) and the emotional valence of the test stimuli (negative vs positive) were manipulated in order to create studies in which the visual and emotional asymmetries were congruent (favoured the same hemispheres) or were incongruent (favoured opposing hemispheres). The results demonstrated that negative emotional stimuli led to a right-hemisphere advantage for conscious perception only when stimuli were shown for brief exposures (17?ms). Positive emotional words did not elicit hemispheric asymmetries. The results are discussed in terms of their relevance for theories of emotional lateralisation.