Startle modulation during emotional anticipation and perception

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.     

Prediction and perception: Defensive startle modulation

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.     

Genetic and environmental influences on emotion-modulated startle reflex: a twin study

Emotion-modulated startle reflex is an important indicator of traitlike differences in affective processing implicated in the biological basis of personality and psychopathology. This study examined heritability of startle modulation by affective pictures in 66 pairs of monozygotic and 57 pairs of dizygotic female twins. Consistent with previous studies, startle magnitude was significantly influenced by emotional valence of the picture (positive < neutral < negative). Absolute response magnitude showed high heritability in all three valence conditions (59-61%); however, there were no significant genetic influences on the amount of startle modulation. Thus, our data do not support the hypothesis that emotion-modulated startle can serve as an indicator of genetically transmitted individual differences in affective processing.     

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.     

Reciprocal modulation of eye-blink and pinna-flexion components of startle during reward anticipation

Because expectancies play a central role in current theories of dopaminergic neuron function, it is important to develop measures of reward anticipation processes. In the present study, reflexogenic bursts of white noise were presented to 39 healthy young adults as they awaited rewards and punishments in a gambling-like task. The rewards were small pieces of chocolate; the punishments, segments of bitter-tasting banana peel. Consistent with prior research on affective valence, postauricular reflexes were larger prior to rewards than punishments, whereas the reverse was true for acoustic blink reflexes. We theorized that potentiation of the postauricular reflex prior to consuming appetizing food is related to the priming of ear-retraction musculature during nursing in our remote ancestors.     

Startle potentiation in aversive anticipation: evidence for state but not trait effects

The present study was undertaken to determine whether aversiveness contributes to startle potentiation in anticipation of affective pictures above and beyond the effects of emotional arousal. Further, participants high in trait anxious apprehension, which is characterized by worry about the future, were expected to show especially pronounced anticipatory startle responses. Startle blink reflex was measured during warning stimuli that predicted the valence of ensuing aversive/unpleasant, pleasant, or neutral pictures. Startle magnitude was larger in anticipation of aversive than of pleasant pictures and smallest in anticipation of neutral pictures. Enhanced startle potentiation was not found in anxious apprehension subjects. These data suggest that the aversive nature of stimuli contribute to the potentiation of startle above and beyond the effects of emotional arousal, which may be a universal phenomenon not modulated by individual differences.     

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.     

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.     

Influence of individual differences in the Behavioral Inhibition System and stimulus content (fear versus blood-disgust) on affective startle reflex modulation

Inconsistencies among affective startle reflex modulation studies may be due to differences in the startle potentiation produced by the specific content of the images used, to individual differences in sensitivity to negative stimuli, or to the interaction of both factors. To explore this interaction, 52 undergraduates obtaining extreme scores on a self-report measure of the Behavioral Inhibition System (BIS) participated in an affective startle reflex modulation paradigm. A significant interaction between BIS group (high versus low) and image content emerged from the MANOVA. Comparing startle magnitude between fear and pleasant images, low BIS participants did not seem to show startle potentiation, whereas high BIS participants did. Both groups displayed potentiated startle during blood-disgust images. The present results suggest the importance of considering personality variables and their interaction with image content in the affective startle modulation paradigm.     

Low baseline startle and deficient affective startle modulation in remitted bipolar disorder patients and their unaffected siblings

We examined whether startle abnormalities are present in bipolar disorder (BD) patients and their unaffected siblings. Twenty‐one remitted patients with BD, 19 unaffected siblings, and 42 controls were presented with 18 pleasant, 18 unpleasant, and 18 neutral pictures. Acoustic probes (104 dB) were presented during 12 of 18 pictures in each affective category at 300, 3000, and 4500 ms after picture onset, so that there were 4 pictures per valence per probe onset type. Baseline startle was assessed during blank screens and was found reduced in patients and sibling groups. We found startle inhibition with the 300 probes and a linear increase in amplitude with valence with the late probes in controls; these effects were absent in patients and their siblings. Low startle and blunted startle reactivity may represent trait deficits in remitted BD patients and their relatives, possibly associated with attentional deficits and adaptive down‐regulation of emotion.     

Affect-modulated startle reflex and dopamine D4 receptor gene variation

The affect-modulated acoustic startle response (ASR) might be a promising indicator for emotional reactivity as an endophenotype (an intermediate level between genetics and phenotypes), which we expected to be associated with the DRD4 polymorphism. Therefore, the affect-modulated ASR was examined in 114 healthy volunteers, 74 lacking the DRD4 7R allele (7R-absent group) and 41 with at least one DRD4 7R allele (7R group). Results revealed the well-known affect–modulated ASR in the 7R-absent group. The 7R group, however, was characterized by a blunted affect-modulated ASR, especially by a reduced startle potentiation toward unpleasant pictures. Associations between the exploratory assessed 5-HTT, COMT, and DAT polymorphisms and affect-modulated ASR were not found. Results speak for the importance of the DRD4 polymorphism in modulating emotional responses and also for the usefulness of the affect-modulated ASR as an endophenotype.     

Intranasal oxytocin attenuates the human acoustic startle response independent of emotional modulation

Oxytocin promotes social affiliation in humans. However, the mechanisms underlying this phenomenon require further elucidation. The present study investigated the influence of intranasal oxytocin on basic emotional processing in men and women, using an emotion‐modulated startle response paradigm. Eighty‐four participants self‐administered 24 IU of intranasal oxytocin or saline and completed an assessment of the acoustic startle reflex, using electromyography (EMG), with varying emotional foregrounds. Oxytocin had no impact on the affective modulation of the startle eye blink response, but significantly diminished the acoustic startle reflex irrespective of the emotional foreground. The results suggest that oxytocin facilitates prosocial behavior, in part, by attenuating basic physiological arousal. The dampening effect of oxytocin on EMG startle could possibly be used as an inexpensive marker of oxytocin's effect on limbic brain circuits.     

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.     

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.     

Temporal stability of the emotion-modulated startle response

In the present study, we examined the stability of one measure of emotion, the emotion-modulated acoustic startle response, in an undergraduate sample. Using the acoustic startle paradigm on two different occasions, we measured stability of affective modulation of the startle response during and following the presentation of pictures selected to be of positive, negative, or neutral emotional valence. The two assessments were separated by 4 weeks. Two groups of subjects were compared: one group that viewed the same pictures at each assessment and a second group that viewed different pictures at the second assessment. We found that viewing different pictures at two assessments separated by 4 weeks yielded moderate stability of the emotion modulation of startle magnitude, whereas subjects who viewed the same pictures at both assessments showed poor stability. Furthermore, this difference was due to the stability of responses to high versus low arousal pictures, not to differences in valence.     

Affect, attention, or anticipatory arousal? Human blink startle modulation in forward and backward affective conditioning

Affect modulates the blink startle reflex in the picture-viewing paradigm, however, the process responsible for reflex modulation during conditional stimuli (CSs) that have acquired valence through affective conditioning remains unclear. In Experiment 1, neutral shapes (CSs) and valenced or neutral pictures (USs) were paired in a forward (CS-->US) manner. Pleasantness ratings supported affective learning of positive and negative valence. Post-acquisition, blink reflexes were larger during the pleasant and unpleasant CSs than during the neutral CS. Rather than affect, attention or anticipatory arousal were suggested as sources of startle modulation. Experiment 2 confirmed that affective learning in the picture-picture paradigm was not affected by whether the CS preceded the US. Pleasantness ratings and affective priming revealed similar extents of affective learning following forward, backward or simultaneous pairings of CSs and USs. Experiment 3 utilized a backward conditioning procedure (US-->CS) to minimize effects of US anticipation. Again, blink reflexes were larger during CSs paired with valenced USs regardless of US valence implicating attention rather than anticipatory arousal or affect as the process modulating startle in this paradigm.     

Emotional qualities of odors and their influence on the startle reflex in humans

Recent human and animal research suggests that the startle reflex might serve as a psychophysiological indicator of the emotional valence of foreground stimulation. The present experiment was designed to evaluate the emotional effects of positive and negative odorant stimuli. We examined the effects of continuous hydrogen sulphide (H2S) and vanillin stimulation on the magnitude of the acoustic startle reflex (measured at the M. orbicularis oculi) and on ratings of subjective valence in 16 healthy subjects. In accordance with the view that odors have emotional qualities, we found that H2S, a presumed negative foreground stimulus, significantly enhanced the startle-reflex amplitude relative to neutral air stimulation, whereas vanillin, a positive foreground stimulus, tended to reduce the reflex amplitude compared with neutral air stimulation. Both odorant stimuli were rated as equally intense by the subjects, and heart rate and electrodermal activity were not affected differentially by the two odorants.     

Startle reflex modulation, affective ratings and autonomic reactivity in incarcerated Spanish psychopaths

Startle probe modulation during affective picture viewing was assessed in a Spanish prison population. As for North American inmates, psychopaths failed to display normal blink potentiation during unpleasant slides even though their evaluative judgments and autonomic reaction to affective stimuli paralleled those of other inmate and noninmate participants. The results suggest that diminished defense activation characterizes psychopaths despite cultural differences.     

Early- and late-onset startle modulation in unipolar depression

In two experimental sessions, we assessed early- and late-onset acoustic startle eyeblink modulation and subjective ratings of emotional pictures by nondepressed participants and by unipolar depressed participants. Depressed participants were assessed before and after treatment with the antidepressant medication Bupropion SR. Both depressed and nondepressed participants exhibited arousal-dependent startle modulation to early probes occurring 300 ms after picture onset. Nondepressed participants demonstrated the expected valence-dependent startle modulation to late probes (3,500-4,500 ms post-onset). In contrast, the late-probe blink magnitudes of depressed patients were unrelated to picture valence. This pattern of group differences was not moderated by treatment. There were no between-group differences in self-report ratings to pictures. These results suggest that depression may be characterized by anomalous responses to affective stimuli and that startle modulation can be a more sensitive index of affective response deficits linked to depression than self-report measures.     

Affective modulation of eyeblink startle with reward and threat

An emotion-modulated acoustic startle paradigm for inducing positive and negative affect was used to address pregoal and postgoal affect. Participants played a computerized lottery task in which they chose digits that could match a subsequently displayed, random set of numbers. In the positive conditions, matches led to monetary rewards. In the negative condition, matches led to an aversive noise blast. In three experiments, we found eyeblink startle magnitude was potentiated just prior to feedback concerning reward outcome, suppressed following the feedback that a monetary reward was won, and potentiated when threatened with an aversive noise. When presented with a 0%, 45%, 90%, or 100% chance of winning, higher probabilities suppressed startle response after feedback whereas the 45% trials did not. These data indicate that postgoal positive affect (winning reward) reliably suppressed the startle response whereas pregoal positive affect did not.