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.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Affective startle modulation in anticipation and perception

Startle modulation was investigated as participants first anticipated and then viewed affective pictures in order to determine whether affective modulation of the startle reflex is similar in these different task contexts. During a 6-s anticipation period, a neutral light cue signaled whether the upcoming picture would portray snakes, erotica, or household objects; at the end of the anticipatory period, a picture in the signaled category was viewed for 6 s. Male participants highly fearful of snakes were recruited to maximize emotional arousal during anticipation and perception. Results indicated that the startle reflex was potentiated when anticipating either unpleasant (phobic) or pleasant (erotic) pictures, compared to neutral stimuli, whereas during perception, reflexes were potentiated when viewing unpleasant stimuli, and reduced when viewing pleasant pictures. The startle reflex is modulated by hedonic valence in picture perception, and by emotional arousal in a task context involving picture anticipation.     

Psychometric properties of startle and corrugator response in NPU,affective picture viewing,and resting state tasks

The current study provides a comprehensive evaluation of critical psychometric properties of commonly used psychophysiology laboratory tasks/measures within the NIMH RDoC. Participants (N = 128) completed the no‐shock, predictable shock, unpredictable shock (NPU) task, affective picture viewing task, and resting state task at two study visits separated by 1 week. We examined potentiation/modulation scores in NPU (predictable or unpredictable shock vs. no‐shock) and affective picture viewing tasks (pleasant or unpleasant vs. neutral pictures) for startle and corrugator responses with two commonly used quantification methods. We quantified startle potentiation/modulation scores with raw and standardized responses. We quantified corrugator potentiation/modulation in the time and frequency domains. We quantified general startle reactivity in the resting state task as the mean raw startle response during the task. For these three tasks, two measures, and two quantification methods, we evaluated effect size robustness and stability, internal consistency (i.e., split‐half reliability), and 1‐week temporal stability. The psychometric properties of startle potentiation in the NPU task were good, but concerns were noted for corrugator potentiation in this task. Some concerns also were noted for the psychometric properties of both startle and corrugator modulation in the affective picture viewing task, in particular, for pleasant picture modulation. Psychometric properties of general startle reactivity in the resting state task were good. Some salient differences in the psychometric properties of the NPU and affective picture viewing tasks were observed within and across quantification methods.     

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.     

Modulation of event-related brain potentials during affective picture processing: a complement to startle reflex and skin conductance response?

The present study compared startle response, skin conductance response (SCR) and subjective variables (valence and arousal ratings, viewing time) assessed in an affective picture paradigm with simultaneously registered event-related brain potentials (ERPs) parameters such as P300 and positive slow waves (PSW). Pleasant, neutral and unpleasant pictures from the International Affective Picture System [Lang, P.J., Bradley, M.M., Cuthbert, B.N., 1999. International Affective Picture System (IAPS): Instruction manual and affective ratings. Technical Report A-4, Center for Research in Psychophysiology. University of Florida, Gainesville, Florida] were presented for 8 s, and startle probes were delivered during picture presentation. Startle response was modulated by picture valence, and SCR by picture arousal. ERP positivity was greater for pleasant and unpleasant than for neutral pictures for the P300 amplitude and the positive slow wave (PSW). ERPs showed characteristic differences and a distinct time course for pictures of different valence categories and may deliver useful information not contained in startle response or SCR measures. The simultaneous registration of startle responses and ERPs in the affective picture paradigm seems valuable.     

Impact of motivational salience on affect modulated startle at early and late probe times.

This study systematically manipulated both picture content and noise probe time in order to evaluate the effects of motivational salience (as distinguished from affective valence) on both early and late modulation of the startle response. Specifically, modulation was compared for erotic versus action/adventure scenes, and for direct threat versus victim scenes, at early (300 and 800 ms) and late (3500 ms) probe times -- all relative to neutral. Blink inhibition was observed at all probe times during presentation of erotic pictures, and blink potentiation was evident at all times during presentation of direct threat pictures. Patterns of blink modulation were less consistent for action and victim picture contents. These findings are consistent with the hypothesis that under conditions of high motivational salience, affective startle modulation indexes the activation of appetitive-approach and defensive motivational states, even at early stages of picture processing.     

Addressing the specificity of affective startle modulation: fear versus disgust.

Previous research is equivocal regarding whether startle reflex eyeblink is specifically sensitive to the emotion of fear, or is more generally influenced by the valence dimension of emotion. Thirty-five undergraduate participants (17 male) viewed 60 pictures from five affective categories (pleasant, neutral, fear, disgust-blood, and disgust-other). Bilateral eyeblink EMG responses to acoustic startle probes were assessed during 2/3 of picture presentations, and 1/3 of intertrial intervals. Left corrugator and levator EMG responses to no-startle pictures were also examined. Supporting an emotional valence model of affective startle modulation, the startle potentiation effect was not specific to fear materials. In fact, women, but not men, had larger startles during disgust compared with fear pictures. Both corrugator and levator EMG responses were greater to disgust than fear stimuli. These findings limit the generalizability of corrugator as an index of all negative emotions and extend evidence for the specificity of levator to disgust.     

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.     

Impact of motivational salience on affect modulated startle at early and late probe times

This study systematically manipulated both picture content and noise probe time in order to evaluate the effects of motivational salience (as distinguished from affective valence) on both early and late modulation of the startle response. Specifically, modulation was compared for erotic versus action/adventure scenes, and for direct threat versus victim scenes, at early (300 and 800 ms) and late (3500 ms) probe times — all relative to neutral. Blink inhibition was observed at all probe times during presentation of erotic pictures, and blink potentiation was evident at all times during presentation of direct threat pictures. Patterns of blink modulation were less consistent for action and victim picture contents. These findings are consistent with the hypothesis that under conditions of high motivational salience, affective startle modulation indexes the activation of appetitive-approach and defensive motivational states, even at early stages of picture processing.     

Lateralization of the startle reflex circuit in humans: an examination with monaural probes following unilateral temporal lobe resection

Startle reflex (SR) modulation elicited by monaural probes during affective picture viewing was investigated in patients following left temporal lobectomy (LTL; n = 8) and right temporal lobectomy (RTL; n = 10) and in controls (n = 18). All patients had undergone anteromedial temporal lobe (ATL) resection. LTL participants exhibited attenuated overall SR magnitude. Affective SR modulation in controls was significant for left ear probes, at both eyes, but not for right ear probes. RTL but not LTL participants displayed significant startle attenuation during pleasant picture viewing. Results suggest that monaural startle probes primarily activate structures in the ipsilateral ATL and that the ATLs are interconnected, with the left ATL more critical in perceiving arousing properties of affective stimuli, necessary for affective SR modulation.     

Startle modulation by affective faces

Startle reflex modulation by affective pictures is a well-established effect in human emotion research. However, much less is known about startle modulation by affective faces, despite the growing evidence that facial expressions robustly activate emotion-related brain circuits. In this study, acoustic startle probes were administered to 37 young adult participants (20 women) during the viewing of slides from the Pictures of Facial Affect set including neutral, happy, angry, and fearful faces. The effect of expression valence (happy, neutral, and negative) on startle magnitude was highly significant (p < .001). Startle reflex was strongly potentiated by negative expressions (fearful and angry), however, no attenuation by happy faces was observed. A significant valence by gender interaction suggests stronger startle potentiation effects in females. These results demonstrate that affective facial expressions can produce significant modulation of the startle reflex.     

Fear appears fast: temporal course of startle reflex potentiation in animal fearful subjects

The temporal course of startle reflex modulation and autonomic response patterns to fear-relevant and fear-irrelevant pictures in subjects with high and low levels of animal fear was investigated. Thirty-eight high-fear and 48 low-fear volunteers viewed photos of snakes and spiders and pictures of neutral and pleasant content. The slides were presented for 6 s or for only 150 ms, depending on the group. Acoustic startle probes were presented at five different times after slide onset. Relative potentiation of the startle responses started 300 ms after onset of snake/spider pictures in fearful subjects. This fear-potentiated startle effect was maintained for the later probe times and was identical in the 150-ms condition. Fear-relevant pictures also prompted a sympathetically dominated autonomic response profile in fearful persons. These data support the idea that fear can be activated very rapidly, requiring only minimal stimulus input.     

Affective modulation and prepulse inhibition of startle among undergraduates high and low in behavioral inhibition and approach

Valence modulation and prepulse inhibition of startle were examined among 80 undergraduates scoring in the upper and lower quartiles of self-report measures of behavioral inhibition (BIS) and behavioral approach (BAS). Participants viewed a series of pleasant, neutral, and unpleasant pictures. Acoustic startle probes (102 dB) were presented during most pictures and during intertrial intervals, and a prepulse (120-ms SOA) preceded half of the probes. Valence modulation on no-prepulse trials was greater among high-BAS than low-BAS participants. Consistent with theory regarding behavioral approach, post hoc tests demonstrated robust inhibition during pleasant versus neutral pictures among high-BAS participants, but not low-BAS participants. Valence modulation was reliable among high-BIS but not low-BIS participants, but the group difference was not significant. Contrary to our prediction, prepulse inhibition tended to be greater among high-BAS than low-BAS participants. The present data call attention to the role of individual differences in pleasant affective experience in startle modification.