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Meta-analytic and experimental studies investigating the neural basis of emotion often compare functional activation in different emotional induction contexts, assessing evidence for a "core affect" or "salience" network. Meta-analyses necessarily aggregate effects across diverse paradigms and different samples, which ignore potential neural differences specific to the method of affect induction. Data from repeated measures designs are few, reporting contradictory results with a small N. In the current study, functional brain activity is assessed in a large (N = 61) group of healthy participants during two common emotion inductions-scene perception and narrative imagery-to evaluate cross-paradigm consistency. Results indicate that limbic and paralimbic regions, together with visual and parietal cortex, are reliably engaged during emotional scene perception. For emotional imagery, in contrast, enhanced functional activity is found in several cerebellar regions, hippocampus, caudate, and dorsomedial prefrontal cortex, consistent with the conception that imagery is an action disposition. Taken together, the data suggest that a common emotion network is not engaged across paradigms, but that the specific neural regions activated during emotional processing can vary significantly with the context of the emotional induction., (© 2020 Society for Psychophysiological Research.)

Pupil diameter is enhanced in a variety of emotional contexts, including viewing pictures, listening to sounds, and during threat of shock. In this study, we investigated pupil diameter changes during emotional imagery. Participants imagined scenes describing pleasant, unpleasant, or neutral events while pupil diameter was continuously recorded. Second by second changes in pupil diameter were analyzed to determine whether, and when, modulation of the pupil as a function of hedonic content is found. Results indicated a significant effect of hedonic content beginning shortly after script onset, with enhanced pupil diameter when imagining emotional (pleasant or unpleasant), compared to neutral, scenes. Pupil diameter during imagery covaried with rated emotional arousal, consistent with an interpretation that changes in pupil diameter during emotional imagery reflect sympathetic nervous system activity. Because emotional imagery is a key element in clinical assessment and treatment, pupil diameter could prove a useful index of emotional engagement in a variety of clinically pertinent contexts., (© 2017 Society for Psychophysiological Research.)

During threat of shock, the startle reflex is potentiated, suggesting modulation by defensive mobilization. To determine whether startle potentiation is specific to aversive anticipation, startle reflexes were measured in the context of either aversive or appetitive anticipation in a between-subject study. Participants wore a device on the wrist that could deliver electrical shock (n = 49), or vibrotactile stimulation indicating monetary reward (n = 48). Cues signaling "threat" or "safe" periods were presented alone, or accompanied by presentation of affective and neutral pictures on half of the trials. Results indicated that the startle reflex was significantly potentiated when anticipating either shock or reward, compared to safe periods, both when no picture was presented, as well as during picture viewing. The difference between threat and safety in both reflex magnitude and skin conductance changes was larger for those anticipating shock, suggesting that the aversive context was more motivationally engaging. The pattern of reflex modulation as a function of picture valence varied under threat and safety, but was identical in the shock and reward groups, consistent with a hypothesis that anticipation of either aversive or appetitive events prompts heightened perceptual vigilance, potentiating the acoustic startle reflex., (© 2017 Society for Psychophysiological Research.)

A series of studies investigated the effects of hedonic content, brightness, and contrast on pupil diameter during free viewing of natural scenes, assessing the amplitude of the initial light reflex and subsequent sustained pupil diameter change. Hedonic picture content varied from highly arousing scenes of erotica and violence to scenes depicting nature, babies, loss, contamination, food, and more. Despite equivalent overall picture brightness and contrast, pupil diameter still varied as a function of the local brightness of central vision at fixation. Statistical (Experiment 1) and methodological (Experiment 2, 3) solutions produced complementary data indicating that scenes of erotica and violence reliably attenuate the amplitude of the initial light reflex and prompt enhanced late diameter pupil changes, compared to other scene contents. A principal components analysis supported the hypothesis that a single sympathetically mediated process enhances pupil dilation during picture viewing, modulating both initial constriction and late diameter changes. Rather than being a subtle index of "liking," pupil diameter is primarily sensitive to events that reliably elicit measurable sympathetic nervous system activity., (© 2017 Society for Psychophysiological Research.)

fMRI studies of reward find increased neural activity in ventral striatum and medial prefrontal cortex (mPFC), whereas other regions, including the dorsolateral prefrontal cortex (dlPFC), anterior cingulate cortex (ACC), and anterior insula, are activated when anticipating aversive exposure. Although these data suggest differential activation during anticipation of pleasant or of unpleasant exposure, they also arise in the context of different paradigms (e.g., preparation for reward vs. threat of shock) and participants. To determine overlapping and unique regions active during emotional anticipation, we compared neural activity during anticipation of pleasant or unpleasant exposure in the same participants. Cues signalled the upcoming presentation of erotic/romantic, violent, or everyday pictures while BOLD activity during the 9-s anticipatory period was measured using fMRI. Ventral striatum and a ventral mPFC subregion were activated when anticipating pleasant, but not unpleasant or neutral, pictures, whereas activation in other regions was enhanced when anticipating appetitive or aversive scenes., (Copyright © 2017 Elsevier B.V. All rights reserved.)

A set of studies are reviewed that investigate the effects of repetition during scene perception on event-related potentials, elucidating perceptual, memory and emotional processes. Repetition suppression was consistently found for the amplitude of early frontal N2 and posterior P2 components, which was greatly enhanced for massed, compared to distributed, repetition. Both repetition suppression and enhancement of the amplitude of a centro-parietal positive potential (LPP) were found in specific contexts. Suppression was reliably found following a massive number of repetitions of few items, whereas enhancement is found when repetitions are spaced; enhancement was apparent both during simple free viewing as well as on an explicit recognition test. Regardless of repetition, an enhanced LPP was always found for emotional, compared to neutral, scenes. Taken together, the data suggest that different effects of massed and distributed repetitions on specific ERP components index perceptual priming, habituation, and spontaneous episodic retrieval., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Hedonic bias during free viewing of novel emotional and neutral scenes was investigated in older adults and college students. A neurophysiological index of emotional picture processing-the amplitude of the centroparietal late positive potential (LPP)-was recorded from the scalp using a dense sensor array while participants (29 older adults; 21 college students) viewed emotionally engaging or mundane natural scenes that varied in specific content. Both students and older adults showed LPP enhancement when viewing affective, compared to neutral, scenes, and there was no difference in LPP amplitude between older individuals and college students when viewing neutral everyday scenes. However, compared to the college students, older individuals showed attenuated LPP amplitude when viewing emotional scenes, regardless of hedonic valence or specific content. Age related differences could be mediated by a reduction in reactive emotional arousal with age, possible mediated by repeated life exposure to emotional stimuli., Competing Interests: The authors have declared that no competing interests exist.

Previous research indicates that predictive cues can dampen subsequent defensive reactions. The present study investigated whether effects of cuing are specific to aversive stimuli, using modulation of the blink startle reflex as a measure of emotional reactivity. Participants viewed pictures depicting violence, romance/erotica, or mundane content. On half of all trials, a cue (color) predicted the content of the upcoming picture; on the remaining trials, scenes were presented without a cue. Acoustic startle probes were presented during picture viewing on trials with predictive cues and trials without a cue. Replicating previous studies, blink reflexes elicited when viewing violent pictures that had not been preceded by a cue were potentiated compared to uncued mundane scenes, and reflexes were attenuated when viewing scenes of erotica/romance that had not been cued. On the other hand, reflex potentiation when viewing scenes of violence (relative to mundane scenes) was eliminated when these pictures were preceded by a predictive cue, whereas scenes of romance prompted reliable reflex attenuation regardless of whether pictures were cued or not. Taken together, the data suggest that cuing elicits an anticipatory coping process that is specific to aversive stimuli., (© 2015 Society for Psychophysiological Research.)

Choice viewing behavior when looking at affective scenes was assessed to examine differences due to hedonic content and gender by monitoring eye movements in a selective looking paradigm. On each trial, participants viewed a pair of pictures that included a neutral picture together with an affective scene depicting either contamination, mutilation, threat, food, nude males, or nude females. The duration of time that gaze was directed to each picture in the pair was determined from eye fixations. Results indicated that viewing choices varied with both hedonic content and gender. Initially, gaze duration for both men and women was heightened when viewing all affective contents, but was subsequently followed by significant avoidance of scenes depicting contamination or nude males. Gender differences were most pronounced when viewing pictures of nude females, with men continuing to devote longer gaze time to pictures of nude females throughout viewing, whereas women avoided scenes of nude people, whether male or female, later in the viewing interval. For women, reported disgust of sexual activity was also inversely related to gaze duration for nude scenes. Taken together, selective looking as indexed by eye movements reveals differential perceptual intake as a function of specific content, gender, and individual differences., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Recent studies have suggested that pupil diameter, like the "old-new" ERP, may be a measure of memory. Because the amplitude of the old-new ERP is enhanced for items encoded in the context of repetitions that are distributed (spaced), compared to massed (contiguous), we investigated whether pupil diameter is similarly sensitive to repetition. Emotional and neutral pictures of natural scenes were viewed once or repeated with massed (contiguous) or distributed (spaced) repetition during incidental free viewing and then tested on an explicit recognition test. Although an old-new difference in pupil diameter was found during successful recognition, pupil diameter was not enhanced for distributed, compared to massed, repetitions during either recognition or initial free viewing. Moreover, whereas a significant old-new difference was found for erotic scenes that had been seen only once during encoding, this difference was absent when erotic scenes were repeated. Taken together, the data suggest that pupil diameter is not a straightforward index of prior occurrence for natural scenes., (© 2015 Society for Psychophysiological Research.)

Cues that signal the possibility of receiving an electric shock reliably induce defensive activation. To determine whether cues can also easily reverse defensive reactions, a threat reversal paradigm was developed in which a cue signaling threat of shock reversed its meaning across the course of the study. This allowed us to contrast defensive reactions to threat cues that became safe cues, with responses to cues that continued to signal threat or safety. Results showed that, when participants were instructed that a previously threatening cue now signaled safety, there was an immediate and complete attenuation of defensive reactions compared to threat cues that maintained their meaning. These findings highlight the role that language can play both in instantiating and attenuating defensive reactions, with implications for understanding emotion regulation, social communication, and clinical phenomena., (© 2014 Society for Psychophysiological Research.)

Emotionally arousing scenes readily capture visual attention, prompting amplified neural activity in sensory regions of the brain. The physical stimulus features and related information channels in the human visual system that contribute to this modulation, however, are not known. Here, we manipulated low-level physical parameters of complex scenes varying in hedonic valence and emotional arousal in order to target the relative contributions of luminance based versus chromatic visual channels to emotional perception. Stimulus-evoked brain electrical activity was measured during picture viewing and used to quantify neural responses sensitive to lower-tier visual cortical involvement (steady-state visual evoked potentials) as well as the late positive potential, reflecting a more distributed cortical event. Results showed that the enhancement for emotional content was stimulus-selective when examining the steady-state segments of the evoked visual potentials. Response amplification was present only for low spatial frequency, grayscale stimuli, and not for high spatial frequency, red/green stimuli. In contrast, the late positive potential was modulated by emotion regardless of the scene's physical properties. Our findings are discussed in relation to neurophysiologically plausible constraints operating at distinct stages of the cortical processing stream., (Copyright © 2015 Elsevier B.V. All rights reserved.)

The startle reflex is potentiated when anticipating emotional, compared to neutral, pictures. This study investigated the time course of reflex modulation during anticipation and the impact of informative cuing on picture perception. Colors were used to signal the thematic content of emotional and neutral scenes; blink response modulation was measured by presenting acoustic startle probes 3, 2, or 1 s before picture onset or 2 s after picture onset. During anticipation of neutral scenes, blink magnitude showed increasing attenuation as picture onset approached, consistent with a modality-directed vigilance account. Conversely, when anticipating emotional scenes, reflex magnitude did not change over time, and blinks elicited closest to picture onset were potentiated compared to neutral. During perception, the expected reflex potentiation for unpleasant pictures was not found, suggesting that cuing may dampen defensive activation., (Copyright © 2014 Society for Psychophysiological Research.)

An initial reflexive constriction of the pupil to stimulation-the light reflex-is primarily modulated by brightness, but is attenuated when participants are under threat of shock (i.e., fear-inhibited light reflex). The present study assessed whether the light reflex is similarly attenuated when viewing emotional pictures. Pupil diameter was recorded while participants viewed erotic, violent, and neutral scenes that were matched in brightness; scrambled versions identical in brightness were also presented as an additional control. Compared to viewing neutral scenes, the light reflex was reliably modulated by hedonic content, with significant attenuation both when viewing unpleasant as well as pleasant pictures. No differences in the light reflex were found among scrambled versions. Thus, emotional modulation of the initial light reflex is not confined to a context of fear and is not indicative of brightness differences when viewing pictures of natural scenes., (Copyright © 2014 Society for Psychophysiological Research.)

On a recognition test, stimuli originally encoded in the context of shock threat show an enhanced late parietal positivity during later recognition compared to stimuli encoded during safety, particularly for emotionally arousing stimuli. The present study investigated whether this ERP old/new effect is further influenced when a threat context is reinstated during the recognition test. ERPs were measured in a yes-no recognition test for words rated high or low in emotional arousal that were encoded and recognized in the context of cues that signaled threat of shock or safety. Correct recognition of words encoded under threat, irrespective of reinstatement, was associated with an enhanced old-new ERP difference (500-700ms; centro-parietal), and this difference was only reliable for emotionally arousing words. Taken together, the data suggest that information processed in a stressful context are associated with better recollection on later recognition, an effect that was not modulated by reinstating the stressful context at retrieval., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Repetition has long been known to facilitate memory performance, but its effects on event-related potentials (ERPs), measured as an index of recognition memory, are less well characterized. In Experiment 1, effects of both massed and distributed repetition on old-new ERPs were assessed during an immediate recognition test that followed incidental encoding of natural scenes that also varied in emotionality. Distributed repetition at encoding enhanced both memory performance and the amplitude of an old-new ERP difference over centro-parietal sensors. To assess whether these repetition effects reflect encoding or retrieval differences, the recognition task was replaced with passive viewing of old and new pictures in Experiment 2. In the absence of an explicit recognition task, ERPs were completely unaffected by repetition at encoding, and only emotional pictures prompted a modestly enhanced old-new difference. Taken together, the data suggest that repetition facilitates retrieval processes and that, in the absence of an explicit recognition task, differences in old-new ERPs are only apparent for affective cues.

During rapid serial visual presentation (RSVP), the perceptual system is confronted with a rapidly changing array of sensory information demanding resolution. At rapid rates of presentation, previous studies have found an early (e.g., 150-280 ms) negativity over occipital sensors that is enhanced when emotional, as compared with neutral, pictures are viewed, suggesting facilitated perception. In the present study, we explored how picture composition and the presence of people in the image affect perceptual processing of pictures of natural scenes. Using RSVP, pictures that differed in perceptual composition (figure-ground or scenes), content (presence of people or not), and emotional content (emotionally arousing or neutral) were presented in a continuous stream for 330 ms each with no intertrial interval. In both subject and picture analyses, all three variables affected the amplitude of occipital negativity, with the greatest enhancement for figure-ground compositions (as compared with scenes), irrespective of content and emotional arousal, supporting an interpretation that ease of perceptual processing is associated with enhanced occipital negativity. Viewing emotional pictures prompted enhanced negativity only for pictures that depicted people, suggesting that specific features of emotionally arousing images are associated with facilitated perceptual processing, rather than all emotional content.

When event-related potentials (ERP) are measured during a recognition task, items that have previously been presented typically elicit a larger late (400-800 ms) positive potential than new items. Recent data, however, suggest that emotional, but not neutral, pictures show ERP evidence of spontaneous retrieval when presented in a free-viewing task (Ferrari, Bradley, Codispoti, Karlsson, & Lang, 2012). In two experiments, we further investigated the brain dynamics of implicit and explicit retrieval. In Experiment 1, brain potentials were measured during a semantic categorization task, which did not explicitly probe episodic memory, but which, like a recognition task, required an active decision and a button press, and were compared to those elicited during recognition and free viewing. Explicit recognition prompted a late enhanced positivity for previously presented, compared with new, pictures regardless of hedonic content. In contrast, only emotional pictures showed an old-new difference when the task did not explicitly probe episodic memory, either when making an active categorization decision regarding picture content, or when simply viewing pictures. In Experiment 2, however, neutral pictures did prompt a significant old-new ERP difference during subsequent free viewing when emotionally arousing pictures were not included in the encoding set. These data suggest that spontaneous retrieval is heightened for salient cues, perhaps reflecting heightened attention and elaborative processing at encoding.

The anticipation of highly aversive events is associated with measurable defensive activation, and both animal and human research suggests that stress-inducing contexts can facilitate memory. Here, we investigated whether encoding stimuli in the context of anticipating an aversive shock affects recognition memory. Event-related potentials (ERPs) were measured during a recognition test for words that were encoded in a font color that signaled threat or safety. At encoding, cues signaling threat of shock, compared to safety, prompted enhanced P2 and P3 components. Correct recognition of words encoded in the context of threat, compared to safety, was associated with an enhanced old-new ERP difference (500-700 msec; centro-parietal), and this difference was most reliable for emotional words. Moreover, larger old-new ERP differences when recognizing emotional words encoded in a threatening context were associated with better recognition, compared to words encoded in safety. Taken together, the data indicate enhanced memory for stimuli encoded in a context in which an aversive event is merely anticipated, which could assist in understanding effects of anxiety and stress on memory processes., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

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., (Copyright © 2011 Wiley Periodicals, Inc.)

Pictures of emotional facial expressions or natural scenes are often used as cues in emotion research. We examined the extent to which these different stimuli engage emotion and attention, and whether the presence of social anxiety symptoms influences responding to facial cues. Sixty participants reporting high or low social anxiety viewed pictures of angry, neutral, and happy faces, as well as violent, neutral, and erotic scenes, while skin conductance and event-related potentials were recorded. Acoustic startle probes were presented throughout picture viewing, and blink magnitude, probe P3 and reaction time to the startle probe also were measured. Results indicated that viewing emotional scenes prompted strong reactions in autonomic, central, and reflex measures, whereas pictures of faces were generally weak elicitors of measurable emotional response. However, higher social anxiety was associated with modest electrodermal changes when viewing angry faces and mild startle potentiation when viewing either angry or smiling faces, compared to neutral. Taken together, pictures of facial expressions do not strongly engage fundamental affective reactions, but these cues appeared to be effective in distinguishing between high and low social anxiety participants, supporting their use in anxiety research., (Copyright © 2012 Elsevier B.V. All rights reserved.)

Eye movements were monitored during picture viewing, and effects of hedonic content, perceptual composition, and repetition on scanning assessed. In Experiment 1, emotional and neutral pictures that were figure-ground compositions or more complex scenes were presented for a 6-s free viewing period. Viewing emotional pictures or complex scenes prompted more fixations and broader scanning of the visual array, compared to neutral pictures or simple figure-ground compositions. Effects of emotion and composition were independent, supporting the hypothesis that these oculomotor indices reflect enhanced information seeking. Experiment 2 tested an orienting hypothesis by repeatedly presenting the same pictures. Although repetition altered specific scan patterns, emotional, compared to neutral, picture viewing continued to prompt oculomotor differences, suggesting that motivationally relevant cues enhance information seeking in appetitive and defensive contexts., (Copyright © 2011 Society for Psychophysiological Research.)

Parkinson's disease (PD) is a neurodegenerative disease that affects motor, cognitive, and emotional functioning. Previous studies reported reduced skin conductance responses in PD patients, compared to healthy older adults when viewing emotionally arousing pictures. Attenuated skin conductance changes in PD may reflect peripheral autonomic dysfunction (e.g., reduced nerve endings at the sweat gland) or, alternatively, a more central emotional deficit. The aim of the current study was to investigate a second measure of sympathetic arousal-change in pupil dilation. Eye movements, a motor-based correlate of emotional processing, were also assessed. Results indicated that pupil dilation was significantly greater when viewing emotional, compared to neutral pictures for both PD patients and controls. On the other hand, PD patients made fewer fixations with shorter scan paths, particularly when viewing pleasant pictures. These results suggest that PD patients show normal sympathetic arousal to affective stimuli (indexed by pupil diameter), but differences in motor correlates of emotion (eye movements)., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Effects of massed repetition on the modulation of the late positive potential elicited during affective picture viewing were investigated in two experiments. Despite a difference in the number of repetitions across studies (from 5 to 30), results were quite similar: The late positive potential continued to be enhanced when participants viewed emotional, compared to neutral, pictures. On the other hand, massed repetition did prompt a reduction in the late positive potential that was most pronounced for emotional pictures. Startle probe P3 amplitude generally increased with repetition, suggesting diminished attention allocation to repeated pictures. The blink reflex, however, continued to be modulated by hedonic valence, despite massive massed repetition. Taken together, the data suggest that the amplitude of the late positive potential during picture viewing reflects both motivational significance and attention allocation., (Copyright © 2010 Society for Psychophysiological Research.)

Viewing a series of aversive pictures prompts emotional reactivity reflecting sustained defensive engagement. The present study examined the effects of a concurrent visual task on autonomic, somatic, electrocortical, and facial components of this defensive state. Results indicated that emotional activation was largely preserved despite continuous visual distraction, although evidence of attenuation was observed in startle reflex and electrocortical measures. Concurrent task-specific reactivity was also apparent, suggesting that motivational circuits can be simultaneously activated by stimuli with intrinsic survival significance and instructed task significance and that these processes interact differently across the separate components of defensive engagement., (Copyright © 2010 Society for Psychophysiological Research.)

Research on emotional perception and learning indicates appetitive cues engage nucleus accumbens (NAc) and medial prefrontal cortex (mPFC), whereas amygdala activity is modulated by the emotional intensity of appetitive and aversive cues. This study sought to determine patterns of functional activation and connectivity among these regions during narrative emotional imagery. Using event-related fMRI, we investigate activation of these structures when participants vividly imagine pleasant, neutral, and unpleasant scenes. Results indicate that pleasant imagery selectively activates NAc and mPFC, whereas amygdala activation was enhanced during both pleasant and unpleasant imagery. NAc and mPFC activity were each correlated with the rated pleasure of the imagined scenes, while amygdala activity was correlated with rated emotional arousal. Functional connectivity of NAc and mPFC was evident throughout imagery, regardless of hedonic content, while correlated activation of the amygdala with NAc and mPFC was specific to imagining pleasant scenes. These findings provide strong evidence that pleasurable text-driven imagery engages a core appetitive circuit, including NAc, mPFC, and the amygdala., (Hum Brain Mapp, 2010. © 2010 Wiley-Liss, Inc.)

Dense array event-related potentials (ERPs) and memory performance were assessed following rapid serial visual presentation (RSVP) of emotional and neutral pictures. Despite the extremely brief presentation, emotionally arousing pictures prompted an enhanced negative voltage over occipital sensors, compared to neutral pictures, replicating previous encoding effects. Emotionally arousing pictures were also remembered better in a subsequent recognition test, with higher hit rates and better discrimination performance. ERPs measured during the recognition test showed both an early (250-350 ms) frontally distributed difference between hits and correct rejections, and a later (400-500 ms), more centrally distributed difference, consistent with effects of recognition on ERPs typically found using slower presentation rates. The data are consistent with the hypothesis that features of affective pictures pop out during rapid serial visual presentation, prompting better memory performance.

Psychophysiological and neuroscience studies of emotional processing undertaken by investigators at the University of Florida Laboratory of the Center for the Study of Emotion and Attention (CSEA) are reviewed, with a focus on reflex reactions, neural structures and functional circuits that mediate emotional expression. The theoretical view shared among the investigators is that expressed emotions are founded on motivational circuits in the brain that developed early in evolutionary history to ensure the survival of individuals and their progeny. These circuits react to appetitive and aversive environmental and memorial cues, mediating appetitive and defensive reflexes that tune sensory systems and mobilize the organism for action and underly negative and positive affects. The research reviewed here assesses the reflex physiology of emotion, both autonomic and somatic, studying affects evoked in picture perception, memory imagery, and in the context of tangible reward and punishment, and using the electroencephalograph (EEG) and functional magnetic resonance imaging (fMRI), explores the brain's motivational circuits that determine human emotion., (Copyright © 2009 Elsevier B.V. All rights reserved.)

Emotion influences the perception of respiratory sensations, although the specific mechanism underlying this modulation is not yet clear. We examined the impact of viewing pleasant, neutral, and unpleasant affective pictures on the respiratory-related evoked potential (RREP) elicited by a short inspiratory occlusion in healthy volunteers. Reduced P3 amplitude of the RREP was found for respiratory probes presented when viewing pleasant or unpleasant series, when compared to those presented during the neutral series. Earlier RREP components, such as Nf, P1, N1, and P2, showed no modulation by emotion. The results suggest that emotion impacts the perception of respiratory sensations by reducing the attentional resources available for processing afferent respiratory sensory signals.

Background: Posttraumatic stress disorder (PTSD) is characterized as a disorder of exaggerated defensive physiological arousal. The novel aim of the present research was to investigate within PTSD a potential dose-response relationship between past trauma recurrence and current comorbidity and intensity of physiological reactions to imagery of trauma and other aversive scenarios., Methods: A community sample of principal PTSD (n = 49; 22 single-trauma exposed, 27 multiple-trauma exposed) and control (n = 76; 46 never-trauma exposed, 30 trauma exposed) participants imagined threatening and neutral events while acoustic startle probes were presented and the eye-blink response (orbicularis occuli) was recorded. Changes in heart rate, skin conductance level, and facial expressivity were also indexed., Results: Overall, PTSD patients exceeded control participants in startle reflex, autonomic responding, and facial expressivity during idiographic trauma imagery and, though less pronounced, showed heightened reactivity to standard anger, panic, and physical danger imagery. Concerning subgroups, control participants with and without trauma exposure showed isomorphic patterns. Within PTSD, only the single-trauma patients evinced robust startle and autonomic responses, exceeding both control participants and multiple-trauma PTSD. Despite greater reported arousal, the multiple-trauma relative to single-trauma PTSD group showed blunted defensive reactivity associated with more chronic and severe PTSD, greater mood and anxiety disorder comorbidity, and more pervasive dimensional dysphoria (e.g., depression, trait anxiety)., Conclusions: Whereas PTSD patients generally show marked physiological arousal during aversive imagery, concordant with self-reported distress, the most symptomatic patients with histories of severe, cumulative traumatization show discordant physiological hyporeactivity, perhaps attributable to sustained high stress and an egregious, persistent negative affectivity that ultimately compromises defensive responding., (2010 Society of Biological Psychiatry. All rights reserved.)

Models of visual emotional perception suggest a reentrant organization of the ventral visual system with the amygdala. Using focused functional magnetic resonance imaging in humans with a sampling rate of 100 ms, here we determine the relative timing of emotional discrimination in amygdala and ventral visual cortical structures during emotional perception. Results show that amygdala and inferotemporal visual cortex differentiate emotional from nonemotional scenes approximately 1 s before extrastriate occipital cortex, whereas primary occipital cortex shows consistent activity across all scenes. This pattern of discrimination is consistent with a reentrant organization of emotional perception in visual processing, in which transaction between rostral ventral visual cortex and amygdala originates the identification of emotional relevance.

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

The foundations of orienting and attention are hypothesized to stem from activation of defensive and appetitive motivational systems that evolved to protect and sustain the life of the individual. Motivational activation initiates a cascade of perceptual and motor processes that facilitate the selection of appropriate behavior. Among these are detection of significance, indexed by a late centro-parietal positivity in the event-related potential, enhanced perceptual processing, indexed by a initial cardiac deceleration, and preparation for action, indexed by electrodermal changes. Data exploring the role of stimulus novelty and significance in orienting are presented that indicate different components of the orienting response habituate at different rates. Taken together, it is suggested that orienting is mediated by activation of fundamental motivational systems that have evolved to support survival.

Visual attention can be voluntarily oriented to detect target stimuli in order to facilitate goal-directed behaviors. Other visual stimuli capture attention because of motivational significance. The aim of the present study was to investigate the relationship between directed and motivated attention using event-related potentials. Affectively engaging pictures were presented either as target stimuli or as nontargets in a categorization task. Results indicated that both task relevance and emotional significance modulated the late positive potential (LPP) over centro-parietal sensors. Effects of directed and motivated attention on the LPP were additive, with the largest centro-parietal positivity found for emotional pictures that were targets of directed attention, and the least for neutral pictures that were nontargets. Taken together, the data provide new information regarding the relationship between motivated and directed attention, and suggest that the LPP reflects the operation of attentional neural circuits that are utilized by both top-down and bottom-up processes.

Pupil diameter was monitored during picture viewing to assess effects of hedonic valence and emotional arousal on pupillary responses. Autonomic activity (heart rate and skin conductance) was concurrently measured to determine whether pupillary changes are mediated by parasympathetic or sympathetic activation. Following an initial light reflex, pupillary changes were larger when viewing emotionally arousing pictures, regardless of whether these were pleasant or unpleasant. Pupillary changes during picture viewing covaried with skin conductance change, supporting the interpretation that sympathetic nervous system activity modulates these changes in the context of affective picture viewing. Taken together, the data provide strong support for the hypothesis that the pupil's response during affective picture viewing reflects emotional arousal associated with increased sympathetic activity.

Electrophysiological studies of human visual perception typically involve averaging across trials distributed over time during an experimental session. Using an oscillatory presentation, in which affective or neutral pictures were presented for 6 s, flickering on and off at a rate of 10 Hz, the present study examined single trials of steady-state visual evoked potentials. Moving window averaging and subsequent Fourier analysis at the stimulation frequency yielded spectral amplitude measures of electrocortical activity. Cronbach's alpha reached values >.79, across electrodes. Single-trial electrocortical activation was significantly related to the size of the skin conductance response recorded during affective picture viewing. These results suggest that individual trials of steady-state potentials may yield reliable indices of electrocortical activity in visual cortex and that amplitude modulation of these indices varies with emotional engagement.

Event-related potentials (ERP) were measured when pleasant, neutral or unpleasant pictures were presented in the context of similarly valenced stimuli, and compared to ERPs elicited when the same pictures were viewed in an intermixed context. An early ERP component (150-300 ms) measured over occipital and fronto-central sensors was specific to viewing pleasant pictures and was not affected by presentation context. Replicating previous studies, emotional pictures prompted a larger late positive potential (LPP, 400-700 ms) and a larger positive slow wave (1-6 s) over centro-parietal sensors that also did not differ by presentation context. On the other hand, ERPs elicited when viewing neutral pictures varied as a function of context, eliciting somewhat larger LPPs when presented in blocks, and prompting smaller slow waves over occipital sensors. Taken together, the data indicate that emotional pictures prompt increased attention and orienting that is unaffected by its context of presentation, whereas neutral pictures are more vulnerable to context manipulations.

Recent human functional imaging studies have linked the processing of pleasant visual stimuli to activity in mesolimbic reward structures. However, whether the activation is driven specifically by the pleasantness of the stimulus, or by its salience, is unresolved. Here we find in two studies that free viewing of pleasant images of erotic and romantic couples prompts clear, reliable increases in nucleus accumbens (NAc) and medial prefrontal cortex (mPFC) activity, whereas equally arousing (salient) unpleasant images, and neutral pictures, do not. These data suggest that in visual perception, the human NAc and mPFC are specifically reactive to pleasant, rewarding stimuli and are not engaged by unpleasant stimuli, despite high stimulus salience.

Event-related potentials (ERPs) were measured while participants viewed affectively arousing and neutral pictures depicting either simple figure-ground compositions or more complex scenes to assess the timing and topography of perceptual and emotional modulation. Emotional pictures elicited a larger late positive potential than neutral pictures in a 400-700-ms window over centro-parietal sensors both for pictures with simple figure-ground composition and for more complex scenes. Picture composition affected ERPs beginning earlier (around 150 ms), with simple figure-ground compositions eliciting less positivity over posterior sensors and less negativity over frontal sensors. Emotionality had little effect on modulation of these early ERPs. These data suggest that the late centro-parietal positive potential primarily reflects motivational relevance, and that earlier posterior (and anterior) components reflect, at least in part, differences in a picture's perceptual organization.

Dense-array electrocortical and functional hemodynamic measures of human brain activity were collected to assess the relationship between 2 established neural measures of emotional reactivity. Recorded in parallel sessions, the slow-wave late positive potential (LPP) and visual cortical blood oxygen level-dependent (BOLD) signals were both modulated by the rated intensity of picture arousal. The amplitude of the LPP correlated significantly with BOLD intensity in lateral occipital, inferotemporal, and parietal visual areas across picture contents. Estimated strength of modeled regional sources did not correlate significantly with regional BOLD intensity. These data suggest that the enhanced positive slow wave seen over posterior sites during emotional picture processing represents activity in a circuit of visual cortical structures, reflecting a perceptual sensitivity to the motivational relevance of visual scenes.

It has been proposed that narrative emotional imagery activates an associative network of stimulus, semantic, and response (procedural) information. In previous research, predicted response components have been demonstrated through psychophysiological methods in peripheral nervous system. Here we investigate central nervous system concomitants of pleasant, neutral, and unpleasant narrative imagery with functional magnetic resonance imaging. Subjects were presented with brief narrative scripts over headphones, and then imagined themselves engaged in the described events. During script perception, auditory association cortex showed enhanced activation during affectively arousing (pleasant and unpleasant), relative to neutral imagery. Structures involved in language processing (left middle frontal gyrus) and spatial navigation (retrosplenium) were also active during script presentation. At the onset of narrative imagery, supplementary motor area, lateral cerebellum, and left inferior frontal gyrus were initiated, showing enhanced signal change during affectively arousing (pleasant and unpleasant), relative to neutral scripts. These data are consistent with a bioinformational model of emotion that considers response mobilization as the measurable output of narrative imagery.

Affectively arousing visual stimuli have been suggested to automatically attract attentional resources in order to optimize sensory processing. The present study crosses the factors of spatial selective attention and affective content, and examines the relationship between instructed (spatial) and automatic attention to affective stimuli. In addition to response times and error rate, electroencephalographic data from 129 electrodes were recorded during a covert spatial attention task. This task required silent counting of random-dot targets embedded in a 10 Hz flicker of colored pictures presented to both hemifields. Steady-state visual evoked potentials (ssVEPs) were obtained to determine amplitude and phase of electrocortical responses to pictures. An increase of ssVEP amplitude was observed as an additive function of spatial attention and emotional content. Statistical parametric mapping of this effect indicated occipito-temporal and parietal cortex activation contralateral to the attended visual hemifield in ssVEP amplitude modulation. This difference was most pronounced during selection of the left visual hemifield, at right temporal electrodes. In line with this finding, phase information revealed accelerated processing of aversive arousing, compared to affectively neutral pictures. The data suggest that affective stimulus properties modulate the spatiotemporal process along the ventral stream, encompassing amplitude amplification and timing changes of posterior and temporal cortex.

Much research demonstrates that emotional stimuli prompt increased amygdala and visual cortical activation. Here we measure functional activity in the visual cortex and amygdala with fMRI while selected fearful and control participants view a range of neutral, emotionally arousing, and fear-relevant pictures. BOLD signal in the amygdala and inferotemporal visual cortex closely covaried during emotional picture viewing, increasing systematically with rated picture arousal. Furthermore, fearful individuals reacting to specific fear cues show parallel, heightened activation in these two structures compared with non-fearful controls. The findings suggest an individually-sensitive, positive linear relationship between the arousing quality of visual stimuli and activation in amygdala and ventral visual cortex, supporting the hypothesized functional connectivity described in the animal model.

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., (Copyright 2004 Lippincott Williams and Wilkins)

Background: There is growing interest in the role of disgust in the pathogenesis of obsessive-compulsive disorder (OCD)., Methods: Eight OCD subjects with contamination preoccupations and eight gender- and age-matched healthy volunteers viewed pictures from the International Affective Picture System during functional magnetic resonance imaging scans., Results: A different distribution of brain activations was found during disgust-inducing visual stimulation in several areas, most notably the insula, compared with neutral stimulation in both OCD subjects and healthy volunteers. Furthermore, whereas activation during the threat-inducing task in OCD subjects showed a pattern similar to that in healthy volunteers, the pattern of activation during the disgust-inducing task was significantly different, including greater increases in the right insula, parahippocampal region, and inferior frontal sites., Conclusions: This pilot study supports the relevance of disgust in the neurocircuitry of OCD with contamination-preoccupation symptoms; future studies looking at non-OCD individuals with high disgust ratings, non-contamination-preoccupied OCD individuals, and individuals with other anxiety disorders are needed.

Allocation of processing resources to emotional picture stimuli was examined using steady-state visual evoked brain potentials (ssVEPs). Participants viewed a set of 60 colored affective pictures from the International Affective Picture System, presented in a flickering mode at 10 Hz in order to elicit ssVEPs. Phase and amplitude of the 10-Hz ssVEP were examined for six picture categories: threat and mutilation (unpleasant), families and erotica (pleasant), and household objects and persons (neutral). Self-reported affective arousal and hedonic valence of the picture stimuli were assessed by means of subjective ratings. Viewing affectively arousing (unpleasant and pleasant) pictures was associated with enhanced ssVEP amplitude at parieto-occipital recording sites, as compared with neutral stimuli. Phase information suggested increased coactivation of right occipitotemporal and frontotemporal sources during processing of affectively arousing stimuli. These findings are consistent with reentrant modulation of early visual processing by distributed networks including subcortical and neocortical structures according to a stimulus's motivational relevance.

Hemodynamic and electrophysiological studies indicate differential brain response to emotionally arousing, compared to neutral, pictures. The time course and source distribution of electrocortical potentials in response to emotional stimuli, using a high-density electrode (129-sensor) array were examined here. Event-related potentials (ERPs) were recorded while participants viewed pleasant, neutral, and unpleasant pictures. ERP voltages were examined in six time intervals, roughly corresponding to P1, N1, early P3, late P3 and a slow wave window. Differential activity was found for emotional, compared to neutral, pictures at both of the P3 intervals, as well as enhancement of later posterior positivity. Source space projection was performed using a minimum norm procedure that estimates the source currents generating the extracranially measured electrical gradient. Sources of slow wave modulation were located in occipital and posterior parietal cortex, with a right-hemispheric dominance.

Emotional reactions are organized by underlying motivational states--defensive and appetitive--that have evolved to promote the survival of individuals and species. Affective responses were measured while participants viewed pictures with varied emotional and neutral content. Consistent with the motivational hypothesis, reports of the strongest emotional arousal, largest skin conductance responses, most pronounced cardiac deceleration, and greatest modulation of the startle reflex occurred when participants viewed pictures depicting threat, violent death, and erotica. Moreover, reflex modulation and conductance change varied with arousal, whereas facial patterns were content specific. The findings suggest that affective responses serve different functions-mobilization for action, attention, and social communication-and reflect the motivational system that is engaged, its intensity of activation, and the specific emotional context.

Adhering to the view that emotional reactivity is organized in part by underlying motivational states--defensive and appetitive--we investigated sex differences in motivational activation. Men's and women's affective reactions were measured while participants viewed pictures with varied emotional and neutral content. As expected, highly arousing contents of threat, mutilation, and erotica prompted the largest affective reactions in both men and women. Nonetheless, women showed a broad disposition to respond with greater defensive reactivity to aversive pictures, regardless of specific content, whereas increased appetitive activation was apparent for men only when viewing erotica. Biological and sociocultural factors in shaping sex differences in emotional reactivity are considered as possible mediators of sex differences in emotional response.