Why do we leak lubricant from the eyes to solicit comfort from others? Why do we bare our teeth and crinkle our faces to express non-aggression? The defensive mimic theory proposes that a broad range of human emotional expressions evolved originally as exaggerated, temporally extended mimics of the fast, defensive reflexes that normally protect the body surface. Defensive reflexes are so important to survival that they cannot be safely suppressed; yet they also broadcast information about an animal's internal state, information that can potentially be exploited by other animals. Once others can observe and exploit an animal's defensive reflexes, it may be advantageous to the animal to run interference by creating mimic defensive actions, thereby manipulating the behaviour of others. Through this interaction over millions of years, many human emotional expressions may have evolved. Here, human social signals including smiling, laughing and crying, are compared component-by-component with the known, well-studied features of primate defensive reflexes. It is suggested that the defensive mimic theory can adequately account for the physical form of not all, but a large range of, human emotional expression.

The focus of the present study was on further exploring anticipatory responses to emotional stimuli by measuring the eyeblink startle reflex in a variation of the picture-picture affective learning procedure. Participants (113 undergraduate women) were not explicitly instructed before the experiment began. Instead, they had to learn the specific relations between cues (geometrical shapes) and emotional pictures based on pairings during the first part of the task. Plausible contingency learning effects were tested afterwards, in a parallel sequence of trials including auditory probes during cues and pictures processing during the second part of the task. Results did show the typical affective startle modulation pattern during perception, linear F(1, 200) = 52.67, p < .0001, but unexpected inhibition for both pleasant and unpleasant, compared to neutral cues, during anticipation, quadratic F(1, 200) = 7.07, p < .009. All patterns of startle modulation were independent of cue-picture contingency awareness (all interactions Fs < 1). Skin conductance changes showed the predictable quadratic trend either during picture perception or anticipatory periods (greater activity for emotional vs. neutral; overall quadratic F(1, 224) = 7.04, p < .01), only for participants fully aware of the cue-picture contingency, quadratic F(1, 158) = 5.86, p < .02. Overall, our results during anticipation (cues processing) seem to suggest that more resources were allocated to highly arousing pictures that engage attention. Differences between the present results and prior research may be attributed to procedural variations in the sample, cues, or instructions. Future studies should also explore in more detail the role of the contingency awareness during anticipation.

Previous research on the components of the emotional response employing factor analytic studies has yielded a two-factor structure (Lang, Greenwald, Bradley, & Hamm, 1993; Cuthbert, Schupp, Bradley, Birbaumer, & Lang, 2000). However, the startle blink reflex, a widely employed measure of the emotional response, has not been considered to date. We decided to include two parameters of the startle reflex (magnitude and latency) in order to explore further how this response fits into the two-factor model of emotion. We recorded the acoustic startle blink response, skin conductance response, heart rate, free viewing time, and picture valence and arousal ratings of 45 subjects while viewing 54 pictures from the International Affective Picture System (IAPS; 18 unpleasant, 18 neutral, and 18 pleasant). Factorizations of all measures gave a two-factor solution (valence and arousal) that accounted for 70% of the variance. Although some measurements, including heart rate change, did not behave as predicted, our results reinforce the two-dimension model of the emotion, and show that startle fits into the model.

The study of cardiac defense has a long tradition in psychological research both within the cognitive approach—linked to Pavlov, Sokolov, and Graham's work on sensory reflexes—and within the motivational one—linked to the work of Cannon and subsequent researchers on the concepts of activation and stress. These two approaches have been difficult to reconcile in the past. We summarize a series of studies on cardiac defense from a different perspective, which allows integration of the traditional approaches. This new perspective emphasizes a sequential process interpretation of the cardiac defense response. Results of descriptive and parametric studies, as well as those of studies examining the physiological and psychological mechanisms underlying the response, show a complex response pattern with both accelerative and decelerative components, with both sympathetic and parasympathetic influences, and with both attentional and emotional significance. The implications of this new look at cardiac defense are discussed in relation to defensive reactions in natural settings, the brain mechanisms controlling such reactions, and their effects on health and illness.

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

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.

In adults, short-lead prepulse inhibition and long-lead prepulse facilitation of startle are greater during attended than ignored prestimuli. To examine these phenomena in children, fourteen 9- to 12-year-old boys completed a tone discrimination task in two sessions separated by 1 week. During each tone series, participants attended to one pitch and ignored the other. Startle probes (102 dB) were presented 120, 240, 2,000 or 4,500 ms following the onset of two-thirds of tones, and during one-third of intertrial intervals. Eyeblink EMG startle was recorded. Percent prepulse inhibition at 120 ms was greater for attended than ignored stimuli in Session 1 but not Session 2. Long-lead prepulse facilitation was marginally greater for attended than ignored tones and did not vary across sessions. Test–retest reliability was moderate during attended prestimuli but was modest during ignored prestimuli. Reliability of attentional modification was poorest at 120 ms and strongest at 4,500 ms. Overall, this study extended prior work in adults and provided a basis for further study of controlled attentional modification of startle in children.

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.

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.

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

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 undergraduates 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.

We investigated emotional disturbances in 36 schizophrenia patients, 48 of their first-degree relatives, and 56 controls to determine if abnormal affective startle modulation could be associated with genetic risk for schizophrenia. Both patients and relatives had a pattern of startle modulation indistinguishable from controls, with potentiated startle amplitude while viewing negative valence slides and attenuation while viewing positive slides. Patients with flat affect did not differ from those without in startle modulation or slide ratings. The patients and their relatives had lower pleasantness ratings of positive slides and the patients had higher pleasantness ratings of the negative slides than controls. The startle paradigm may not be useful for identifying individuals with a genetic liability for schizophrenia. The results suggest that low-level defensive and appetitive behaviors are unaffected in schizophrenia.

The present study examined the effect of self-focused attention on the startle reflex and heart rate and assessed the assumption that socially anxious individuals become self-focused in evaluative situations. Twenty-five high and 30 low socially anxious men performed a digit recall task under evaluative or nonevaluative instructions. Half of the trials were performed under self-focused conditions and half under non-self-focused conditions. Self-focus led to larger startle responses among socially anxious individuals and had no effect on heart rate. Self-focus combined with evaluation led to poorer recall performance among all subjects. Results provide evidence against a directional change in attention during self-focused attention and favor the view that self-focus increases access to self-relevant thought content and decreases available processing resources.