Startle Eyeblink Modulation Research Articles

Reduced emotional responsiveness in individuals with marginal elevation in blood pressure within the normal range: Evidence from altered affect-modulated startle response

Reduced responsiveness to emotional stimuli (‘emotional dampening’) has been observed in normotensives with elevated blood pressure (BP) and hypertensives but it is not known whether this is due to aberrant responding to emotional information at the involuntary level and whether it is also associated with minimal elevations in BP in the normal range. In this study, we examined emotional dampening using the affect-modulated startle paradigm given its proven sensitivity to motivational states of approach and withdrawal, typically independent of conscious intentional control. Acoustically elicited startle eye-blink modulation was measured using electromyography of the orbicularis oculi muscle beneath the left eye in 59 healthy individuals while they viewed pleasant, unpleasant and neutral standardized pictures. The expected startle attenuation to pleasant pictures, and startle potentiation to unpleasant pictures, relative to neutral pictures, was found in people in the comparison (N = 29) but not elevated BP (N = 30) group. This finding was further supported by significant moderating effect (assessed using ANCOVA and sub-sample analysis) of BP on valence-startle amplitude relationship. The comparison BP group also showed slower latencies to response onset for pleasant stimuli compared to neutral and unpleasant, with no effect of valence in the elevated BP group. However, BP did not moderate the valence-onset latency relationship. Our findings indicate that previously reported emotional dampening associated with elevated BP extends to reduced involuntary emotional reactivity and to individuals with even minimal BP elevations (i.e. higher but still within the normal range). Future research needs to confirm these findings in hypertensive individuals, preferably using within-subjects designs.

Flashing a smile: Startle eyeblink modulation by masked affective faces.

Affective faces are important stimuli with relevance to healthy and abnormal social and affective information processing. The aim of this study was to investigate the effect of brief presentations of affective faces on attention and emotional state across the time course of stimulus processing, as indexed by startle eyeblink response modulation. Healthy adults were presented with happy, neutral, and disgusted male and female faces that were backward masked by neutral faces. Startle responses were elicited at 300, 800, and 3,500 ms following stimulus presentation to probe early and late startle eyeblink modulation, indicative of attention allocation and emotional state, respectively. Results revealed that, at 300 ms, both face expression and face gender modulated startle eyeblink response, suggesting that more attention was allocated to masked happy compared to disgusted female faces, and masked disgusted compared to neutral male faces. There were no effects of either face expression or face gender on startle modulation at 800 ms. At 3,500 ms, target face expression did not modulate startle, but male faces elicited larger startle responses than female faces, indicative of a more negative emotional state. These findings provide a systematic investigation of attention and emotion modulation by brief affective faces across the time course of stimulus processing.

Enhanced startle reflexivity during presentation of visual nurture cues in young adults who experienced parental divorce in early childhood

Adverse childhood experiences (ACE) may influence stress and affective processing in adulthood. Animal and human studies show enhanced startle reflexivity in adult participants with ACE. This study examined the impact of one of the most common ACE, parental divorce, on startle reflexivity in adulthood.Affective modulation of acoustically-elicited startle eye blink was assessed in a group of 23 young adults with self-reported history of parental divorce, compared to an age- and sex-matched control group (n=18). Foreground pictures were either aversive (e.g. mutilation and injury), standard appetitive (e.g. erotic, recreational sport), or nurture pictures (e.g. related to early life, parental care), intermixed with neutral pictures (e.g. household objects), and organized in three valence blocks delivered in a balanced, pseudo-randomized sequence. During picture viewing startle eye blinks were elicited by binaural white noise bursts (50ms, 105 dB) via headphones and recorded at the left orbicularis oculi muscle via EMG.A significant interaction of group×picture valence (p=0.01) was observed. Contrast with controls revealed blunted startle responsiveness of the ACE group during presentation of aversive pictures, but enhanced startle during presentation of nurture-related pictures. No group differences were found during presentation of standard appetitive pictures. ACE participants rated nurture pictures as more arousing (p=0.02) than did control participants.Results suggest that divorce in childhood led to altered affective context information processing in early adulthood. When exposed to unpleasant (vs. neutral) pictures participants with ACE showed less startle potentiation than controls. Nurture context, however, potentiated startle in ACE participants, suggesting visual cuing to activate protective behavioral responses.

Respiratory modulation of startle eye blink: a new approach to assess afferent signals from the respiratory system.

Current approaches to assess interoception of respiratory functions cannot differentiate between the physiological basis of interoception, i.e. visceral-afferent signal processing, and the psychological process of attention focusing. Furthermore, they typically involve invasive procedures, e.g. induction of respiratory occlusions or the inhalation of CO2-enriched air. The aim of this study was to test the capacity of startle methodology to reflect respiratory-related afferent signal processing, independent of invasive procedures. Forty-two healthy participants were tested in a spontaneous breathing and in a 0.25 Hz paced breathing condition. Acoustic startle noises of 105 dB(A) intensity (50 ms white noise) were presented with identical trial frequency at peak and on-going inspiration and expiration, based on a new pattern detection method, involving the online processing of the respiratory belt signal. The results show the highest startle magnitudes during on-going expiration compared with any other measurement points during the respiratory cycle, independent of whether breathing was spontaneous or paced. Afferent signals from slow adapting phasic pulmonary stretch receptors may be responsible for this effect. This study is the first to demonstrate startle modulation by respiration. These results offer the potential to apply startle methodology in the non-invasive testing of interoception-related aspects in respiratory psychophysiology.This article is part of the themed issue 'Interoception beyond homeostasis: affect, cognition and mental health'.

Evidence that Illness-Compatible Cues Are Rewarding in Women Recovered from Anorexia Nervosa: A Study of the Effects of Dopamine Depletion on Eye-Blink Startle Responses.

In anorexia nervosa (AN), motivational salience is attributed to illness-compatible cues (e.g., underweight and active female bodies) and this is hypothesised to involve dopaminergic reward circuitry. We investigated the effects of reducing dopamine (DA) transmission on the motivational processing of AN-compatible cues in women recovered from AN (AN REC, n = 17) and healthy controls (HC, n = 15). This involved the acute phenylalanine and tyrosine depletion (APTD) procedure and a startle eye-blink modulation (SEM) task. In a balanced amino acid state, AN REC showed an increased appetitive response (decreased startle potentiation) to illness-compatible cues (underweight and active female body pictures (relative to neutral and non-active cues, respectively)). The HC had an aversive response (increased startle potentiation) to the same illness-compatible stimuli (relative to neutral cues). Importantly, these effects, which may be taken to resemble symptoms observed in the acute stage of illness and healthy behaviour respectively, were not present when DA was depleted. Thus, AN REC implicitly appraised underweight and exercise cues as more rewarding than did HC and the process may, in part, be DA-dependent. It is proposed that the positive motivational salience attributed to cues of emaciation and physical activity is, in part, mediated by dopaminergic reward processes and this contributes to illness pathology. These observations are consistent with the proposal that, in AN, aberrant reward-based learning contributes to the development of habituation of AN-compatible behaviours.

Startle eye-blink modulation by facial self-resemblance and current mood

Although salient stimuli are known to modulate startle eye-blink responses, and one's own face is considered of particular salience, effects of facial self-resemblance on startle responsiveness have not been systematically investigated. For the present study, pictures from the FACES database (rated as neutral) were digitally morphed to resemble the participants' (N=37) faces to varying degrees (25–50–75%). Perceptually matched geometrical shapes served as a control condition. At SOAs of either 300ms or 3000ms after picture onset, startle responses were elicited by white noise (50ms, 105dB), and recorded at the orbicularis oculi via EMG. Prior to the experiment, self-reported mood was assessed by means of the PANAS. Relative to non-face stimuli, the presentation of faces reduced startle magnitude at short, but not long, lead intervals. Furthermore, for probes presented at a SOA of 300ms, a linear decrease in startle magnitude with higher levels of self-resemblance was observed, presumably reflecting higher salience of the self-face. The startle modulating effect of self-resembling faces during longer lead intervals was moderated by the participants' current mood: negative affect predicted stronger patterns of attenuation, which might be interpreted as an increase in self-focus resulting from more negative mood.

Self-Reported Trait Mindfulness and Affective Reactivity: A Motivational Approach Using Multiple Psychophysiological Measures

As a form of attention, mindfulness is qualitatively receptive and non-reactive, and is thought to facilitate adaptive emotional responding. One suggested mechanism is that mindfulness facilitates disengagement from an affective stimulus and thereby decreases affective reactivity. However, mindfulness has been conceptualized as a state, intervention, and trait. Because evidence is mixed as to whether self-reported trait mindfulness decreases affective reactivity, we used a multi-method approach to study the relationship between individual differences in self-reported trait mindfulness and electrocortical, electrodermal, electromyographic, and self-reported responses to emotional pictures. Specifically, while participants (N = 51) passively viewed pleasant, neutral, and unpleasant IAPS pictures, we recorded high-density (128 channels) electrocortical, electrodermal, and electromyographic data to the pictures as well as to acoustic startle probes presented during the pictures. Afterwards, participants rated their subjective valence and arousal while viewing the pictures again. If trait mindfulness spontaneously reduces general emotional reactivity, then for individuals reporting high rather than low mindfulness, response differences between emotional and neutral pictures would show relatively decreased early posterior negativity (EPN) and late positive potential (LPP) amplitudes, decreased skin conductance responses, and decreased subjective ratings for valence and arousal. High mindfulness would also be associated with decreased emotional modulation of startle eyeblink and P3 amplitudes. Although results showed clear effects of emotion on the dependent measures, in general, mindfulness did not moderate these effects. For most measures, effect sizes were small with rather narrow confidence intervals. These data do not support the hypothesis that individual differences in self-reported trait mindfulness are related to spontaneous emotional responses during picture viewing.

Startle Eyeblink Modulation as a Measure of Environmental Concern

Event Abstract Back to Event Startle Eyeblink Modulation as a Measure of Environmental Concern Royce Willis1* and Stephen Provost1 1 Southern Cross University, Australia Aims: Fear appeals seek to elicit behaviour designed to reduce negative affective associated with certain images and the thoughts which they engender. They are widely employed in efforts to encourage environmentally sustainable behaviour. The success of fear appeal depends upon whether the images employed elicit negative affect in the target group. Subjective reporting may not provide accurate information on which to base intervention decisions for a variety of reasons, including social desirability. The startle eye-blink modulation paradigm is founded on evidence showing that startles are inhibited by pleasant stimuli and facilitated by unpleasant stimuli, enabling the measurement of attitudes in a number of social domains. The present study was designed to provide initial evidence for its utility in assessing stimuli relevant to environmentally sustainable action. Method: Nineteen participants completed an environmental concern questionnaire. Participants then viewed environmentally positive, environmentally negative, and neutral images (5 seconds in duration each) prior to startle elicitation by 50 milliseconds of white noise at 95 decibels, presented 3.5 seconds after image onset. Results: Positive environmental images resulted in suppression of eyeblink startle responses, relative to neutral pictures, as anticipated. However, negative environmental images also resulted in smaller eyeblink startle responses. The size of this suppressive effect for both types of images correlated with the environmental concern scale. Eyeblink startle elicited during neutral stimuli was negatively correlated with environmental concern scores. Conclusions: The influence of environmental images on eyeblink suppression is complex. Further research including unambiguously valenced pictorial cues is needed to determine the likely value of such images in fear appeals. The relation between environmental concern and eyeblink startle to neutral stimuli, though unexpected, is consistent with reports of similar effects obtained with respect to, for example, political conservatism. These data thus suggest possible links between environmental attitudes and other dimensions of social concern. Acknowledgements I would like to thank Prof. Lesley Christidis, Prof. Robert P. O’Shea, Urte Roeber, and Duncan Blair for their contributions. Keywords: startle eyeblink modulation, startle reflex modulation, environmental concern, Electromyographic (EMG), attitude measurement, physiological measurement Conference: ASP2013 - 23rd Annual meeting of the Australasian Society for Psychophysiology, Wollongong, Australia, 20 Nov - 22 Nov, 2013. Presentation Type: Oral Presentation Topic: Emotion Citation: Willis R and Provost S (2013). Startle Eyeblink Modulation as a Measure of Environmental Concern. Conference Abstract: ASP2013 - 23rd Annual meeting of the Australasian Society for Psychophysiology. doi: 10.3389/conf.fnhum.2013.213.00050 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 05 Nov 2013; Published Online: 05 Nov 2013. * Correspondence: Mr. Royce Willis, Southern Cross University, Coffs Harbour, Australia, vourne@hotmail.com Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Royce Willis Stephen Provost Google Royce Willis Stephen Provost Google Scholar Royce Willis Stephen Provost PubMed Royce Willis Stephen Provost Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

An Ensemble Method for Classifying Startle Eyeblink Modulation from High-Speed Video Records

Psychophysiological measurements of startle eyeblink can provide information about the state of an individual regarding sensory, attentional, cognitive, and affective processing, and thus reveal valences of interest for affective computing. However, eyeblink is usually measured using intrusive contact electromyographic (EMG) electrodes, accompanied by a laborious manual process of feature extraction. We introduce a new noninvasive automatic system using high-speed video recording of startle blinks in conjunction with data-driven feature selection and support vector machine (SVM) ensembles to classify startle eyeblinks. Using a prestimulus (prepulse) to produce robust modulation of acoustically elicited startle eyeblinks, we tracked the blinks using 250 frames per second video, and extracted different features from eyelid displacement and velocity signals. The SVMs were able to determine whether a trial had contained startle or prepulse+startle stimuli with an accuracy of up to 73 percent (five-fold cross validation). By fusing the decisions made on different feature sets, an ensemble of seven SVMs increased this rate to almost 79 percent. Since startle eyeblinks are robustly modulated by not only sensory events (such as the prepulse used in this study) but also affective and cognitive states, eyelid tracking using high-speed video, in conjunction with the introduced classification method, is an effective and user-friendly alternative to EMG for classification of startle blinks to infer users' affective-cognitive states.

Lateralization effects on the cardiac modulation of acoustic startle eye blink

Cardiac modulation of startle eye blink has been introduced as a methodology to reflect baro-afferent signal transmission. Recent studies showed that affective startle modulation is specific to left-ear presentation that may be due to hemispheric specificity in processing emotional-relevant stimuli, similar to the processing of visceral- and baro-afferent stimuli. To explore whether cardiac modulation of startle eye blink is lateralized as well, 37 healthy volunteers received 160 unilateral acoustic startle probes of 105 dB(A) intensity presented to both ears, one at a time. They were elicited 0, 100, 230, and 530 ms after the R-wave of the cardiac cycle. Startle response magnitude was significantly diminished at a latency of 230 ms, which may be due to the baro-afferent neural feedback at this temporal location, but only for left-ear presentation. This lateralization effect in the cardiac modulation of startle eye blink may reflect the previously described advantages of right-hemispheric brain structures in relaying viscero- and baro-afferent signal transmission.

The chronometry of affective startle modulation in unipolar depression.

Affective startle eyeblink modulation by unipolar depressed and nondepressed participants was assessed during the anticipation and viewing of emotional pictures. Anticipatory startle probes were presented at 2,000 ms and 750 ms before picture onset. Startle probes during picture viewing were presented at 300 ms and 3,500-4,500 ms after picture onset. Although nondepressed participants demonstrated the predicted quadratic and linear patterns of responding in the 2,000-ms anticipatory and 3,500-4,500-ms viewing conditions, respectively, depressed participants were not significantly responsive to differences among picture valence categories at these probe conditions. There were no between-groups differences in startle modulation at the other two probe intervals, in picture ratings, or in behavioral responses to pictures. There was also little evidence of hyperresponsivity to negatively valenced stimuli in the depressed group. These results indicate that depression-related affective hyporesponsivity extended to startle modulation but that the nature and magnitude of the differences between depressed and nondepressed individuals were conditional on the specific cognitive and motivational processes recruited at different points in time.

Attention and Emotional Responses to Sexual Stimuli and Their Relationship to Sexual Desire

Little is known about why individuals vary in their levels of sexual desire. Information processing models, like Barlow's (Journal of Consulting and Clinical Psychology 54:140-148, 1986) model of sexual functioning, suggest that individuals with higher sexual desire attend more and respond with more pleasant emotions to sexual cues than individuals with lower levels of sexual desire. In this study, 69 participants (36 women, 33 men) completed a dot detection task measuring attention capture by sexual stimuli and a startle eyeblink modulation task indexing the valence of emotional response to affective stimuli. Participants with high levels of sexual desire were slower to detect targets in the dot detection task that replaced sexual images but did not differ in startle eyeblink responses to sexual stimuli. The results suggest that the amount of attention captured by sexual stimuli is a stronger predictor of a person's sexual desire level than the valence of the emotional responses elicited by such stimuli.

Startling secrets: Startle eye blink modulation by concealed crime information

We used the startle eye blink paradigm to investigate the processes underlying physiological responding to concealed information. Autonomic responding was measured together with eye blink responses to startle probes presented during mock crime and control pictures. Based upon ‘orienting theory’, greater startle modulation to crime pictures in comparison to control pictures was expected. The electrodermal, heart rate and respiratory changes in Experiment 1 ( n = 29) showed enhanced orienting to the crime pictures, but we observed reduced rather than enhanced startle modulation. In Experiment 2 ( n = 91) and Experiment 3 ( n = 38), participants either were or were not instructed to inhibit physiological responding in order to test whether inhibition was an explanation for the pattern of startle blink responding. Reduced startle modulation was observed only when participants inhibited physiological responding, confirming the proposed role of inhibition. The data suggest that not only orienting, but also inhibition contributes to physiological responding to concealed information.

Deficient attentional modulation of startle eyeblink is associated with symptom severity in the schizophrenia spectrum

Patients with schizophrenia-spectrum disorders show deficient prepulse inhibition (PPI) of the startle eyeblink reflex which is thought to reflect an early stage of information processing called automatic sensorimotor gating. They also exhibit deficient attentional modulation of PPI and prepulse facilitation (PPF) of startle which is thought to reflect deficient early and later controlled attentional processing. This is the first study to assess attentional modulation of PPI and PPF in a 3-group schizophrenia-spectrum sample of age- and sex-matched unmedicated schizotypal personality disorder (SPD) and schizophrenia patients, and healthy controls. Participants performed a tone-length judgment task involving attended, ignored, and novel tone prepulses while the acoustic startle eyeblink reflex was measured. Healthy controls showed greater PPI and PPF during the attended prepulses compared with the ignored prepulses. In contrast, both the SPD and schizophrenia patient groups failed to show this pattern, indicating deficient early and later controlled attentional processing. These findings suggest abnormal attentional modulation of PPI and PPF may be a trait-like feature found in patients with schizophrenia-spectrum disorders. Among the schizophrenia-spectrum sample, more deficient PPI during the attended prepulses was associated with greater symptom severity as measured by the total 18-item Brief Psychiatric Rating Scale score.

Differential motivational responses to food and pleasurable cues in anorexia and bulimia nervosa: a startle reflex paradigm

Abnormal perceptions of food and shape underpin the cognitive behavioural model of eating disorders (EDs). The aim of this study was to investigate motivational processing of disorder-specific and standard emotional cues in anorexia (AN) and bulimia nervosa (BN) using startle eyeblink modulation (SEM). SEM was used because it is sensitive to motivational states of approach (appetitive response) and withdrawal (aversive response), which are independent of conscious intentional control. Acoustically elicited SEM and subjective anxiety ratings were measured in 30 female patients with an ED (n=15 AN, n=15 BN) and 30 female control subjects while they viewed ED-relevant stimuli (food, female bodies) and standardized emotional pictures. BN subjects showed an appetitive response (startle inhibition) to food relative to neutral cues that differed significantly from AN subjects. By contrast, self-reports indicated significantly increased anxiety related to food cues across both ED groups. To female body picture (relative to neutral) cues, no significant between-group differences were found for SEM. ED subjects unexpectedly showed an aversive response (startle potentiation) to positive cues, in contrast to controls, who showed the established startle attenuation. These preliminary results suggest that BN patients demonstrated an exaggerated appetitive response to food but not to standardized positive cues, whereas SEM in AN patients points to a generalized failure to activate the appetitive motivational system. Differences in motivational salience to food cues are in line with distinctions between AN and BN in eating behaviour and food consumption and support differential treatment requirements.

Startle eyeblink modulation: detecting changes in directed attentional allocation during early preattentive processing

Startle eyeblink modification was examined as a measure of allocation of attentional resources during active attention tasks in the early stage of information processing. Fifty-five participants were presented with a series of 250- and 40-ms tones of either high or low pitch which were followed by startle-eliciting stimuli at a lead interval of 120 ms. Attentional allocation was manipulated by instructing one group (Passive) to simply listen to the tones; the second group (Active 1) to count the number of low tones and the third group (Active 2) to count the long high-pitched tones and the short low-pitched tones. Startle eyeblink was significantly more inhibited for the Active 1 group than the Passive group (control) with no significant difference between the two directed attentional conditions (Active 1 and Active 2 groups). However, across the three attentional groups, the degree of startle eyeblink modulation appeared to reflect the degree of attention allocated to the task. The results support the utility of the startle probe in evaluating controlled attentional allocation during the early stages of information processing.

Effect of varying levels of mental workload on startle eyeblink modulation

Previous research using punctuate reaction time and counting tasks has found that the startle eyeblink reflex is sensitive to attentional demands. The present experiment explored whether startle eyeblink is also modulated during a complex continuous task and is sensitive to different levels of mental workload. Participants (N = 14) performed a visual horizontal tracking task either alone (single-task condition) or in combination with a visual gauge monitoring task (multiple-task condition) for three minutes. On some task trials, the startle eyeblink reflex was elicited by a noise burst. Results showed that startle eyeblink was attenuated during both tasks and that the attenuation was greater during the multiple-task condition than during the single-task condition. Subjective ratings, endogenous eyeblink rate, heart period, and heart period variability provided convergent validity of the workload manipulations. The findings suggest that the startle eyeblink is sensitive to the workload demands associated with a continuous visual task. The application of startle eyeblink modulation as a workload metric and the possibility that it may be diagnostic of workload demands in different stimulus modalities is discussed.

Assessing the Effects of Attention and Emotion on Startle Eyeblink Modulation

Abstract Two experiments were conducted to assess simultaneously the effects of attentional and emotional processes on startle eyeblink modulation. In each experiment, participants were presented with a pleasant and an unpleasant picture. Half the participants were asked to attend to the pleasant picture and to ignore the unpleasant picture, whereas the reverse was the case for the other participants. Startle probes were presented at 3500 and 4500 ms after stimulus onset in Experiment 1 and at 250, 750, and 4450 ms after stimulus onset and 950 ms after stimulus offset in Experiment 2. Attentional processing affected startle eyeblink modulation and electrodermal responses in both experiments. However, effects of picture valence on startle eyeblink modulation were found only in Experiment 2. The results confirm the utility of startle eyeblink modulation as an index of attentional and emotional processing. They also illustrate that procedural characteristics, such as the nature of the lead intervals and how attention and emotion are operationalized, can determine whether emotional or attentional processes will be reflected in startle eyeblink.

Attentional modulation of startle in psychosis-prone college students.

Attentional modulation of startle eyeblink was studied in college students putatively at risk for psychosis and in normal controls. At-risk subjects had extreme scores on scales for either anhedonia or perceptual aberration-magical thinking (per-mags). Subjects were presented with to-be-attended and to-be-ignored tones; white noise startle probes were presented at lead intervals of 60, 120, 240, or 2,000 ms following the onset of attended and ignored tones and during intertone intervals. Controls showed greater inhibition of startle blink at 120 ms and greater facilitation at 2,000 ms during to-be-attended than to-be-ignored tone, demonstrating attentional modulation of prepulse inhibition and facilitation. Both at-risk groups showed normal overall levels of early inhibition and late facilitation. However, per-mags failed to show attentional modulation of either inhibition at 120 ms or facilitation at 2,000 ms; anhedonics showed no modulation of inhibition and modulation of facilitation was delayed in development. The results for the per-mags are strikingly similar to those observed in schizophrenic patients and suggest that these deficits index a trait-linked vulnerability to disorders in the schizophrenic spectrum.