The effects of contextual threat and anxiety on affective startle modulation

The startle reflex is attenuated and potentiated when participants are viewing pleasant and unpleasant images, respectively. Research demonstrates that threatening contexts also potentiate startle, but it remains unclear how a threatening context might impact startle modulation to emotional images, especially as a function of trait anxiety. The present study measured startle reactivity while 43 participants viewed pleasant, unpleasant, and neutral images across conditions of threat-of-shock and safety (i.e., no shock). Compared to neutral images, startle was potentiated during unpleasant images and attenuated during pleasant images. Threat-of-shock potentiated startle during all picture types, suggesting that threat-of-shock broadly sensitized the defensive system but did not change affective modulation of startle. Lastly, higher levels of trait anxiety were associated with less startle potentiation during unpleasant images across both conditions—a finding in line with previous research demonstrating deficient threat mobilization in response to unpleasant stimuli among highly anxious individuals.     

Temporal course of emotional startle modulation in schizophrenia patients.

The temporal course of startle reflex modulation to emotional pictures was investigated in 49 relatively asymptomatic schizophrenia outpatients and 46 normal controls. Participants viewed pleasant, neutral and unpleasant pictures, which were presented for 6 s and acoustic startle probes were delivered at five different times after picture onset. Replicating previous findings schizophrenia patients showed the same affective modulation of the startle reflex as healthy controls when probes were presented later during the picture-viewing period (3800 ms). For the early probe times (300 and 800 ms), affective pictures proved to be the effective prepulse stimuli resulting in a clear Prepulse inhibition (PPI) effect in both groups. In contrast to previous findings, the PPI of startle response was not stronger during processing emotional stimuli relative to neutral stimuli. For control subjects, blink reflexes were larger for unpleasant pictures compared with neutral and pleasant pictures, representing an early activation of motivational systems. Schizophrenia patients on the other hand, did not show this early affective modulation of the acoustic startle reflex.     

Emotion and motivated behavior: postural adjustments to affective picture viewing

Thirty-six participants (18 female, 18 male) viewed affective pictures to investigate the coupling between emotional reactions and motivated behavior. Framed within the biphasic theory of emotion, the three systems approach was employed by collecting measures of subjective report, expressive physiology, and motivated behavior. Postural adjustments associated with viewing affective pictures were measured. Results indicated sex-differences for postural responses to unpleasant pictures; an effect not found for pleasant and neutral picture contents. Females exhibited increased postural movement in the posterior direction, and males exhibited increased movement in the anterior direction, for unpleasant pictures. Subjective report of valence and arousal using the self-assessment manikin (SAM), and the startle eye-blink reflex were collected during a separate session, which replicated previous picture-viewing research. Specifically, participants rated pleasant pictures higher in valence and exhibited smaller startle responses compared to unpleasant pictures. Females also reported lower valence ratings compared to males across all picture contents. These findings extend our knowledge of motivated engagement with affective stimuli and indicate that postural responses may provide insight into sex-related differences in withdrawal behavior.     

Modulation of spinal reflexes: Arousal, pleasure, action

The human startle reflex is reliably modulated by the affective valence of foreground pictures, with larger reflexes elicited when viewing unpleasant relative to pleasant scenes. If this modulation is due to priming of the defensive startle reflex by an aversive foreground, a different pattern should occur for a reflex that is not inherently defensive in nature. In the current study, affective modulation was investigated using the spinal tendinous (T) reflex, which is well documented as sensitive to differences in arousal and is involved in actions that are both appetitively and defensively motivated. As such, T reflexes elicited during unpleasant pictures were not expected to be augmented relative to those elicited in the context of pleasant pictures. Results showed that T reflexes were facilitated during processing of arousing stimuli – either pleasant or unpleasant relative to low-arousal neutral materials. These effects of emotional stimuli on T-reflex amplitude are consistent with the hypothesis that motivational priming underlies affective reflex modulation.     

Probe P3 and blinks: Two measures of affective startle modulation

Two concurrent measures of the evoked startlé response, the elicited blink reflex and the event-related potential, were measured while individuals viewed pictures that varied in pleasure and arousal. Replicating previous findings, the blink response was modulated by picture pleasantness, with larger reflexes elicited in the context of viewing unpleasant versus pleasant pictures. However, the probe P3 was primarily modulated by picture arousal, with smaller P3 responses elicited when viewing affective (pleasant or unpleasant) than when viewing neutral pictures. Both modulatory effects were sustained for probes presented in a subsequent picture imagery period. These data suggest that two measurable responses to the same startle probe are differentially modified by emotional context, with blink magnitude varying with pleasure and probe P3 varying with stimulus arousal.     

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.     

The psychophysiology of social anxiety: Emotional modulation of the startle reflex during socially-relevant and -irrelevant pictures

The present study examined affective processes of social anxiety (SA) through emotional modulation of the startle reflex. Eighty-four high and low trait socially anxious undergraduates viewed socially-relevant and socially-irrelevant pleasant, neutral, and unpleasant pictures, and acoustic startle probes were presented during pictures and the inter-trial interval. Startle was potentiated during unpleasant compared to pleasant stimuli, but this valence modulation did not reliably vary between groups or socially-relevant and -irrelevant stimuli. However, when participants were categorized based on public-speaking fears rather than general SA symptoms, the high fear group demonstrated reliable valence modulation, whereas the low fear group did not. These findings are interpreted within the context of the broader literature suggesting that the specificity of fear in SA may influence psychophysiological reactivity.     

Neural response to emotional pictures is unaffected by concurrent task difficulty: an event-related potential study

The late positive potential (LPP) is an event-related potential that is enhanced when viewing arousing (pleasant and unpleasant) pictures compared to neutral pictures. The affective modulation of the LPP is believed to reflect the increased attention to, and perceptual processing of, emotional stimuli. The present study examined whether concurrent task difficulty (performing mathematics) would modulate the LPP while participants viewed emotionally arousing stimuli. Results indicated that the LPP was larger following pleasant and unpleasant stimuli than it was following neutral stimuli; moreover, the magnitude of this increase was not influenced by concurrent task difficulty. This finding suggests that the affective modulation of neural activity during picture viewing is relatively automatic and is insusceptible to competing task demands. Results are further discussed in terms of the LPP's role in motivated attention and implications for research on emotion regulation.     

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.     

Probing picture perception: Activation and emotion

When acoustic startle probes are presented during picture viewing, the blink reflex is augmented for unpleasant foreground stimuli and reduced during pleasant stimuli. The present experiment assessed the hypothesis that this affect-Startle effect increases as pictures are judged to be more arousing. Eyeblinks elicited by startle probes of three different intensities were recorded while subjects viewed pictures varying in both pleasure (pleasant, neutral, and unpleasant) and arousal (low, moderate, and high). Both blink potentiation during unpleasant content and blink diminution during pleasant content were clearly strongest for picture contents high in arousal. The effect was present for all probe intensities tested.     

Startle modulation before, during and after exposure to emotional stimuli.

Although affective modulation of the startle reflex is a highly replicable effect, the majority of studies have administered startle probes during exposure to affective stimuli. To examine more comprehensively the temporal course of startle potentiation, we assessed blink modulation before, during and immediately after exposure to positive, negative and neutral pictures. During each trial, cues about the affective content of pictures were presented, after which acoustic startle probes were delivered either before picture onset, during picture onset or immediately after picture offset. As expected, we observed a linear relation between picture valence and startle amplitude during picture viewing. Surprisingly, startle amplitude was larger while anticipating pleasant and unpleasant pictures relative to neutral pictures. No significant effects were observed during the offset phase. These results indicate that startle modulation is conditional upon temporal factors linked to stimulus onset and offset.     

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.     

Postauricular reflexes elicited by soft acoustic clicks and loud noise probes: Reliability,prepulse facilitation,and sensitivity to picture contents

The startle blink reflex is facilitated during early picture viewing, then inhibited by attention during pleasant and aversive pictures compared to neutral pictures, and finally potentiated during aversive pictures specifically. However, it is unclear whether the postauricular reflex, which is elicited by the same loud acoustic probe as the startle blink reflex but enhanced by appetitive instead of defensive emotion, has the same pattern and time course of emotional modulation. We examined this issue in a sample of 90 undergraduates using serially presented soft acoustic clicks that elicited postauricular (but not startle blink) reflexes in addition to standard startle probes. Postauricular reflexes elicited by both clicks and probes correlated during food and nurturant contents, during which they were potentiated compared to neutral pictures, suggesting clicks effectively elicit emotionally modulated postauricular reflexes. The postauricular reflex was initially facilitated during the first 500 ms of picture processing but was larger during pleasant than neutral pictures throughout picture processing, with larger effect sizes during the latter half of picture processing. Across reflexes and eliciting stimuli, measures of emotional modulation had higher coefficient alphas than magnitudes during specific picture contents within each valence, indicating that only emotional modulation measures assess higher‐order appetitive or defensive processing.     

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.     

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.     

Emotional modulation of the post-auricular reflex

A large literature now exists on emotional modulation of the startle blink reflex. The current study examined affective modulation of the post-auricular reflex, which can be measured in relation to the same noise probe used to evoke the startle reflex. We recorded the post-auricular reflex during viewing of pictures that varied systematically in emotional valence, content, and intensity. A significant linear valence modulation effect was found, with pleasant pictures potentiating and aversive pictures inhibiting the post-auricular reflex in comparison with neutral pictures. This modulatory effect did not vary as a function of picture content, but it was most robust for highly intense emotional pictures. Implications for the assessment of basic emotional action tendencies are discussed.     

The impact of emotion on the perception of graded magnitudes of respiratory resistive loads

Emotional state can modulate the perception of respiratory loads but the range of respiratory load magnitudes affected by emotional state is unknown. We hypothesized that viewing pleasant, neutral and unpleasant affective pictures would modulate the perception of respiratory loads of different load magnitudes. Twenty-four healthy adults participated in the study. Five inspiratory resistive loads of increasing magnitude (5, 10, 15, 20, 45 cmH₂O/L/s) were repeatedly presented for one inspiration while participants viewed pleasant, neutral and unpleasant affective picture series. Participants rated how difficult it was to breathe against the load immediately after each presentation. Only at the lowest load, magnitude estimation ratings were greater when subjects viewed the unpleasant series compared to the neutral and pleasant series. These results suggest that negative emotional state increases the sense of respiratory effort for single presentations of a low magnitude resistive load but high magnitude loads are not further modulated by emotional state.     

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.     

Affective modulation of nociception at spinal and supraspinal levels

This study was designed to examine the effect of emotion on the nociceptive flexion reflex and pain ratings. To do so, 28 participants viewed pictures varying in emotional valence (unpleasant, neutral, pleasant) and electric stimulations were delivered during and in between pictures. Biceps femoris EMG resulting from the stimulations was used to quantify the nociceptive flexion reflex (spinal nociception), and pain ratings to the stimulations were used as an evaluative measure of supraspinal nociception. Manipulation checks suggested that pictures effectively manipulated emotion. Moreover, nociceptive flexion reflex magnitudes and pain ratings were modulated in a parallel manner. Specifically, viewing unpleasant pictures enhanced the nociceptive flexion reflex and pain, whereas viewing pleasant pictures inhibited the reflex and pain. Analyses suggested that emotional valence, but not arousal, mediated the effects of pictures.     

Modulation of the initial light reflex during affective picture viewing

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.