Pictures as prepulse: Attention and emotion in startle modification

The effects of an emotional stimulus prepulse on probe startle response were examined here. Pleasant, neutral, and unpleasant pictures were viewed for 6 s, and an acoustic startle probe was presented either 300, 800, 1,300, or 3,800 ms after slide onset, or 300 or 3,800 ms after slide offset. Blink magnitude and onset latency demonstrated (a) an early (prepulse) inhibition effect in which reflexes elicited immediately after slide onset were smaller than reflexes elicited later in the viewing interval, and (b) affective modulation, in which unpleasant stimuli prompted larger reflexes than pleasant. Interactive effects of probe time and picture valence indicated attention/arousal effects early and pleasantness effects late in the picture interval. Effects of both attention and emotion can be simultaneously measured using this startle-probe paradigm, encouraging its use in both basic and clinical contexts.     

The selective processing of briefly presented affective pictures: an ERP analysis

Recent event-related potential (ERP) studies revealed the selective processing of affective pictures. The present study explored whether the same phenomenon can be observed when pictures are presented only briefly. Toward this end, pleasant, neutral, and unpleasant pictures from the International Affective Pictures Series were presented for 120 ms while event related potentials were measured by dense sensor arrays. As observed for longer picture presentations, brief affective pictures were selectively processed. Specifically, pleasant and unpleasant pictures were associated with an early endogenous negative shift over temporo-occipital sensors compared to neutral images. In addition, affective pictures elicited enlarged late positive potentials over centro-parietal sensor sites relative to neutral images. These data suggest that a quick glimpse of emotionally relevant stimuli appears sufficient to tune the brain for selective perceptual processing.     

The varying time courses of attentional and affective modulation of the startle eyeblink reflex

In previous research on modulation of the startle eyeblink reflex, emotional effects have been demonstrated only at late probe positions, whereas attentional effects have been found at both early and late positions but only when the prepulses were affectively neutral. In Experiment 1, participants viewed emotionally valenced pictures and were instructed to attend to the duration of half of the slides. Affective modulation of the startle eyeblink occurred at long lead intervals, but attentional modulation also occurred late. In Experiment 2, participants viewed the same slides used in Experiment 1 but were instructed to attend to the duration of only the positive or the negative slides. Affective modulation occurred at both early and late probe positions, whereas attentional effects occurred only following slide offset. Early (250 ms) affective modification of startle eyeblink has not been previously reported. These results suggest that the time courses of emotional and attentional modulation of startle are variable and can occur at both early and late startle probe positions, depending on task requirements.     

Resting heart rate variability and the startle reflex to briefly presented affective pictures

We have previously shown that persons with low HRV showed potentiated startle responses to neutral stimuli. In the present study we replicated our prior findings and extended them to examine the effects of HRV on the startle magnitude to pictures that were presented outside of conscious awareness. A total of 85 male and female students were stratified via median split on their resting HRV. They were presented pictures for 6 s or for 30 ms. Results indicated that the high HRV group showed the context appropriate startle magnitude increase to unpleasant foreground. The low HRV group showed startle magnitude increase from pleasant to neutral pictures but no difference between the neutral and unpleasant pictures. This pattern of results was similar for the 30 ms and the 6 s conditions. These results suggest that having high HRV may allow persons to more efficiently process emotional stimuli and to better recognize threat and safety signals.     

A multi-process account of startle modulation during affective perception

Modulation of the startle reflex by sensory, attentional, and emotional processes was explored by presenting acoustic startle probes at various delays following picture onset. Within 500 ms of onset, blinks were first greatly facilitated and then inhibited, indicating prepulse facilitation and prepulse inhibition that did not vary with affect. Attention allocation to the picture continued across the viewing interval and was most pronounced for emotional pictures, as determined by attenuation of the P3 component to the startle probe. Startle potentiation for unpleasant pictures occurred later in the viewing interval and was strongest for highly arousing pictures. Taken together, the startle reflex during picture viewing is modulated by sequential and sometimes concurrent processes of prepulse facilitation, prepulse inhibition, attentional inhibition, and affective modulation, with reflex magnitude reflecting the net effect of multiple processes.     

Independence of valence modulation and prepulse inhibition of startle

This study sought to determine whether prepulse inhibition and valence modulation of startle are independent, both within and across individuals. Acoustic probes (105 dB) were delivered as 68 undergraduate s viewed pleasant, neutral, and unpleasant pictures. Weak acoustic stimuli (8 dB above background) preceded half of the probes by 120 ms. As expected, startles were larger during unpleasant than during pleasant pictures, and smaller on prepulse than no-prepulse trials. In general, valence modulation and prepulse inhibition of startle were unrelated. That is, prepulse inhibition was consistent across affective states, valence modulation did not differ between no-prepulse and prepulse trials, and valence modulation and prepulse inhibition effects were uncorrelated across individuals. Analysis of raw and percent modification scores generally led to similar conclusions. It is concluded that valence modulation and prepulse inhibition are independent startle modulatory phenomena, although this conclusion is tempered by a finding of poor internal consistency reliability for valence modulation.     

Affective reactions to pictures of ingroup and outgroup members

A pervasive form of social categorization among humans is between us and them. In this study, we assessed emotional reactions when people viewed pictures depicting members of the same or different ethnic group. African American and European American participants viewed a series of pleasant and unpleasant pictures portraying either ingroup or outgroup members, while physiological, behavioral, and evaluative judgments were measured. Two hypotheses were assessed. The outgroup antipathy hypothesis predicts that people will respond to outgroup pictures with more negative affect than to ingroup pictures. In contrast, the ingroup empathy hypothesis predicts that people will show exaggerated (pleasant and unpleasant) affective responses to pictures of ingroup members, due to group identification or personal relevance. The data provided no support for the antipathy hypothesis, whereas facial EMG, skin conductance, rating, and viewing time data lent support to the ingroup empathy hypothesis, in which greater pleasure and displeasure were apparent when viewing ingroup pictures.     

Affective imagery and the startle response: probing mechanisms of modulation during pleasant scenes,personal experiences,and discrete negative emotions

Two experiments addressed the following issues concerning modulation of the acoustic startle reflex during emotional imagery: (1) Is startle inhibited or potentiated during imagery of pleasurable events? (2) Does startle modulation differ for personal versus standard imagery scenes? (3) Is startle modulated differently during anger versus fear? For standard scenes, startle was greater during aversive than pleasant imagery, with both exceeding neutral. Blink potentiation was greater during imagery of personal pleasant than nonpersonal pleasant scenes. Startle potentiation did not differ for anger versus fear material, but differences were found in self‐report, corrugator EMG, and HR response. These results suggest that in addition to the emotional valence of imagined material, startle reactivity is influenced by the degree of engagement or active disengagement from the sensory environment. The findings also indicate that fear and anger are differentiable in terms of affective report, cardiac mobilization, and expressive behavior, but not at the primary motivational level at which reflex priming occurs.     

Emotional specificity of startle potentiation during the early stages of picture viewing

Potentiation of the startle blink reflex shortly after picture onset has only been reported in a sample of high animal-fear participants during fear-evoking stimuli. The experiment tested the emotional specificity of startle reflex modification at early (300 ms) and late (2-5 s) stages of picture viewing in an unselected sample (n=55). Participants viewed two negative picture categories, threat and disgust contents, in addition to neutral and positive pictures. Blink potentiation occurred for threat contents at both probe times, relative to neutral responses. Disgust contents only potentiated blinks relative to positive content responses. These results are inconsistent with a basic, prepulse interpretation of early startle modification and suggest that affective modification can be observed shortly after picture onset, depending on the specific emotional content of the picture.     

Affective reactions to acoustic stimuli

Emotional reactions to naturally occurring sounds (e.g., screams, erotica, bombs, etc.) were investigated in two studies. In Experiment 1, subjects rated the pleasure and arousal elicited when listening to each of 60 sounds, followed by an incidental free recall task. The shape of the two-dimensional affective space defined by the mean ratings for each sound was similar to that previously obtained for pictures, and, like memory for pictures, free recall was highest for emotionally arousing stimuli. In Experiment 2, autonomic and facial electromyographic (EMG) activity were recorded while a new group of subjects listened to the same set of sounds; the startle reflex was measured using visual probes. Listening to unpleasant sounds resulted in larger startle reflexes, more corrugator EMG activity, and larger heart rate deceleration compared with listening to pleasant sounds. Electrodermal reactions were larger for emotionally arousing than for neutral materials. Taken together, the data suggest that acoustic cues activate the appetitive and defensive motivational circuits underlying emotional expression in ways similar to pictures.     

Lack of startle blink potentiation to mutilation pictures irrespective of fearfulness

Previous research has shown that in healthy individuals blood-related stimuli elicit a distinctive autonomic response pattern and heightened processing as compared with other unpleasant and arousing visual stimuli. In addition, growing evidence suggests that information processing of disorder-related stimuli is also different in blood phobia as compared with other specific phobias. In the present study, the magnitude of the startle eyeblink reflex elicited during the viewing of mutilation, human attack, erotica and neutral pictures was recorded in 22 blood phobics and 25 healthy controls. Startle eyeblink responses were measured at 300, 1500, 3500 and 4500-ms time intervals after picture onset in order to assess the attentional/affective modulation and its temporal course. Reliable startle inhibition to erotic pictures and startle potentiation to human attack scenes were found relative to neutral pictures. However, while both groups rated mutilations as the most unpleasant and arousing content, no blink facilitation relative to neutral contents was found at either early or late probe times. Crucially, such effect occurred independently of fear levels, as no difference between phobics and controls was found in the size of the startle blinks elicited throughout the viewing of blood pictures.     

Probing affective pictures: Attended startle and tone probes

Reflexive eyeblinks to a startle probe vary with the pleasantness of affective pictures, whereas the corresponding P300 varies with emotional arousal. The impact of attention to the probe on these effects was examined by varying task and probe type. Probes were either nonstartling tones or startling noises presented during affective picture viewing. Half the participants performed a task requiring attention to the probes; the other participants were told to ignore the probes. Blinks to the startle probe varied with picture pleasantness for both task and nontask conditions. In contrast, P300 magnitudes for both startle and tone probes were reduced during emotionally arousing pictures, irrespective of pleasantness, in task and nontask conditions. Further, attending to the startle probe prompted an augmentation of N100 during unpleasant pictures. The data suggest that affective modulation of probe responses reflects obligatory processes in picture perception.     

The pupil's response to affective pictures: Role of image duration,habituation, and viewing mode

The pupil has been shown to be sensitive to the emotional content of stimuli. We examined this phenomenon by comparing fearful and neutral images carefully matched in the domains of luminance, image contrast, image color, and complexity of content. The pupil was more dilated after viewing affective pictures, and this effect was (a) shown to be independent of the presentation time of the images (from 100–3,000 ms), (b) not diminished by repeated presentations of the images, and (c) not affected by actively naming the emotion of the stimuli in comparison to passive viewing. Our results show that the emotional modulation of the pupil is present over a range of variables that typically vary from study to study (image duration, number of trials, free viewing vs. task), and encourages the use of pupillometry as a measure of emotional processing in populations where alternative techniques may not be appropriate.     

Emotional reactions in children: verbal, physiological, and behavioral responses to affective pictures

Many studies have shown a consistent pattern in adults' responses to affective pictures and there is growing evidence of gender differences, as well. Little is known, though, about children's verbal, behavioral, and physiological responses to affective pictures. Two experiments investigated children's responses to pictures. In Experiment 1, children, adolescents, and adults viewed pictures varying in affective content and rated them for pleasure, arousal, and dominance. Results indicated that children and adolescents rated the pictures similarly to adults. In Experiment 2, physiological responses, self-report, and viewing time were measured while children viewed affective pictures. As with adults, children's responses reflected the affective content of the pictures. Gender differences in affective evaluations, corrugator activity, skin conductance, startle modulation, and viewing time indicated that girls were generally more reactive to unpleasant materials.     

Influence of a monetary incentive upon attentional modification of short-lead prepulse inhibition and long-lead prepulse facilitation of acoustic startle

Short-lead prepulse inhibition and long-lead prepulse facilitation of startle are greater during attended than ignored prestimuli. The present work examined whether this attentional modification is influenced by monetary incentive. Participants (43 college students) were randomly assigned to receive a small performance-based monetary incentive or were instructed to try their best. The task was to judge the duration of tones of one of two pitches during a series of 48 tones. Prepulse inhibition of startle eyeblink EMG was assessed at 60, 120, and 240 ms, and prepulse facilitation was assessed at 4,500 ms following tone onset. Short-lead percent prepulse inhibition was greater during attended than ignored prestimuli only at 120 ms among paid participants. Long-lead prepulse facilitation was greater for attended than ignored tones, but this effect did not vary with incentive condition. This study demonstrates that attentional modification of short-lead prepulse inhibition is sensitive to a monetary incentive and provides a basis for further examination of motivational effects on early attentional processing.     

Motivated attention: incentive effects on attentional modification of prepulse inhibition

Prepulse inhibition (PPI) of startle is greater for attended compared to ignored prestimuli, and, consistent with theories of motivated attention, initial evidence suggests that this effect is greater among participants given performance-based incentives. The present study examined a within-subjects incentive manipulation. Participants (n=41) completed two blocks of a tone discrimination task. During the incentive block, participants received trialwise feedback with small monetary incentives for task performance. Startle eyeblink EMG responses to auditory probes were assessed at 60-, 120-, and 180-ms tone-probe stimulus onset asynchronies (SOAs). As predicted, PPI was enhanced during attended compared to ignored prestimuli only at the 120-ms SOA in the incentive condition. There was no evidence of attentional modification in the no-incentive condition. These data suggest that attentional modification of PPI is sensitive to within-subjects manipulations of incentive, providing a useful tool for testing models of motivated attention in psychopathology and psychopharmacology.     

Affective modulation of tactile startle

Two studies were conducted to investigate affective modulation of startle responses to unilateral tactile probes and to determine whether such modulation is lateralized. Right-handed undergraduates received airpuffs to the left or right temple while viewing pleasant, neutral, and unpleasant pictures. Side of probe presentation was varied between the two trial blocks of the experiment in Study 1 (n= 48) but varied randomly within trial blocks in Study 2 (n= 48). Primary results were consistent across studies. Replicating and extending the findings for acoustic probes, eyeblink responses to tactile probes were larger during unpleasant than during pleasant pictures. However, affective modulation of startle did not differ reliably between the two sides of probe presentation (sensory laterality) or the two sides of the response (motor lateralily) in either study or in a combined analysis.     

To blink or not to blink: inhibition and facilitation of reflex blinks

A reflex blink typically inhibits subsequent blinks. In this study, we investigated whether the nature and time course of this inhibition vary when different combinations of blink-evoking stimuli are used. We used the paired stimulus paradigm, in which two blink-evoking stimuli — a conditioning stimulus followed by a test stimulus — are presented with a variety of interstimulus intervals, to examine the interactions between blinks evoked by trigeminal and acoustic stimuli in rats and humans. In addition, we studied the effect of a blink-evoking trigeminal stimulus on subsequent gaze-evoked blinks in humans. The results revealed that long-lasting inhibition occurred when the conditioning and test stimuli were within the same modality. A shorter period of inhibition followed by facilitation occurred when the stimuli were in different modalities. The data demonstrate that a blink-evoking stimulus initiates a lengthy period of inhibition in its own sensory pathway and a shorter period of inhibition in the reticular formation and/or in blink motoneurons. In addition, the results show that the blink-evoking stimulus also initiates a facilitatory process. Thus, the magnitude of a blink reflects a balance between inhibitory and facilitatory processes.     

Emotional event-related potentials are reduced if negative pictures presented at fixation are unattended

Viewing of emotional pictures elicits two event-related potentials (ERPs) to emotional versus neutral pictures: an early posterior negativity (EPN) and a late positive potential (LPP). Because it is unresolved whether these indexes of emotional processing are reduced to task-irrelevant pictures at fixation, negative and neutral pictures from the International Affective Picture Set (IAPS) were shown at fixation together with 6 letters that surrounded the pictures. In separate tasks, participants were instructed to attend either the pictures or the letters. When the pictures were task relevant, results showed an EPN and LPP. In contrast, when the pictures were task irrelevant, the EPN was eliminated and the LPP reduced. Performance was high in both tasks (hit rates>87%), but somewhat better when the pictures were relevant. However, analyses showed no relationship between this performance difference and the differences in EPN and LPP between tasks. These results suggest that emotional processing of strong, negative pictures is sensitive to manipulations of attention even if the pictures are shown at fixation.     

Selective Attention Effects on the Reflex Blink

Past work has shown that facilitation of reflex blinking accompanies cardiac deceleration when the attention of subjects is directed to a reflex-eliciting stimulus. The present studies showed that when warning stimuli directed attention instead to weak (tactile) stimuli presented simultaneously with reflex-eliciting (acoustic) stimuli, cardiac deceleration was still present but reflex magnitude was unchanged or inhibited. However, latency to reflex onset remained facilitated, i.e., latency and magnitude changes were discordant. The findings were interpreted as evidence for two independent processes: a process capable of selectively enhancing or attenuating sensory input and a non-selective process presumably facilitating motor pathways.