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Two studies examined emotional responding to food cues. In experiment 1, normal college students were assigned to 0-, 6- or 24-h of food deprivation prior to presentations of standard emotional and food-related pictures. Food deprivation had no impact on responses elicited by standard emotional pictures. However, subjective and psychophysiological reactions to food pictures were affected significantly by deprivation. Importantly, food-deprived subjects viewing food pictures showed an enhanced startle reflex and increased heart rate. Experiment 2 replicated the food deprivation effects from experiment 1, and examined participants reporting either a habitual pattern of restrained (anorexia-like) or binge (bulimia-like) eating. Food-deprived and binge eater groups showed startle potentiation to food cues, and rated these stimuli as more pleasant, relative to restrained eaters and control subjects. The results are interpreted from the perspective that startle modulation reflects activation of defensive or appetitive motivation. Implications of the data for understanding eating disorders are considered.

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.

The aim of this paper is to explicate what is special about emotional information processing, emphasizing the neural foundations that underlie the experience and expression of fear. A functional, anatomical model of defense behavior in animals is presented and applications are described in cognitive and physiological studies of human affect. It is proposed that unpleasant emotions depend on the activation of an evolutionarily primitive subcortical circuit, including the amygdala and the neural structures to which it projects. This motivational system mediates specific autonomic (e.g., heart rate change) and somatic reflexes (e.g., startle change) that originally promoted survival in dangerous conditions. These same response patterns are illustrated in humans, as they process objective, memorial, and media stimuli. Furthermore, it is shown how variations in the neural circuit and its outputs may separately characterize cue-specific fear (as in specific phobia) and more generalized anxiety. Finally, again emphasizing links between the animal and human data, we focus on special, attentional features of emotional processing: The automaticity of fear reactions, hyper-reactivity to minimal threat-cues, and evidence that the physiological responses in fear may be independent of slower, language-based appraisal processes.

Emotionally arousing picture stimuli evoked scalp-recorded event-related potentials. A late, slow positive voltage change was observed, which was significantly larger for affective than neutral stimuli. This positive shift began 200-300 ms after picture onset, reached its maximum amplitude approximately 1 s after picture onset, and was sustained for most of a 6-s picture presentation period. The positive increase was not related to local probability of content type, but was accentuated for pictures that prompted increased autonomic responses and reports of greater affective arousal (e.g. erotic or violent content). These results suggest that the late positive wave indicates a selective processing of emotional stimuli, reflecting the activation of motivational systems in the brain.

Functional activity in the visual cortex was assessed using functional magnetic resonance imaging technology while participants viewed a series of pleasant, neutral, or unpleasant pictures. Coronal images at four different locations in the occipital cortex were acquired during each of eight 12-s picture presentation periods (on) and 12-s interpicture interval (off). The extent of functional activation was larger in the right than the left hemisphere and larger in the occipital than in the occipitoparietal regions during processing of all picture contents compared with the interpicture intervals. More importantly, functional activity was significantly greater in all sampled brain regions when processing emotional (pleasant or unpleasant) pictures than when processing neutral stimuli. In Experiment 2, a hypothesis that these differences were an artifact of differential eye movements was ruled out. Whereas both emotional and neutral pictures produced activity centered on the calcarine fissure (Area 17), only emotional pictures also produced sizable clusters bilaterally in the occipital gyrus, in the right fusiform gyrus, and in the right inferior and superior parietal lobules.

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.

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 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 hypothesis that motivational priming underlies affective reflex modulation.

Previous research with both animal and human subjects has shown that startle reflex magnitude is potentiated in an aversive stimulus context, relative to responses elicited in a neutral or appetitive context. In the present experiment, the same pleasant, unpleasant, and neutral picture stimuli were repeatedly presented to human subjects. Startle reflex habituation was assessed in each stimulus context and was compared with the habituation patterns of heart rate, electrodermal, and facial corrugator muscle responses. All systems showed initial differentiation among affective picture contents and general habituation over trials. The startle reflex alone, however, continued to differentiate among pleasant, neutral, and unpleasant pictures throughout the presentation series. These results suggest that (a) the startle probe reflex is relatively uninfluenced by stimulus novelty, (b) the startle modulatory circuit (identified with amygdala-reticular connections in animals) varies systematically with affective valence, and (c) the modulatory influence is less subject to habituation than is the obligatory startle pathway or responses in other somatic and autonomic systems.

Startle-elicited blinks were measured during presentation of affective slides to test hypotheses concerning emotional responding in psychopaths. Subjects were 54 incarcerated sexual offenders divided into nonpsychopathic, psychopathic, and mixed groups based on file and interview data. Consistent with findings for normal college students, nonpsychopaths and mixed subjects showed a significant linear relationship between slide valence and startle magnitude, with startle responses largest during unpleasant slides and smallest during pleasant slides. This effect was absent in psychopaths. Group differences in startle modulation were related to affective features of psychopathy, but not to antisocial behavior per se. Psychopathy had no effect on autonomic or self-report responses to slides. These results suggest an abnormality in the processing of emotional stimuli by psychopaths that manifests itself independently of affective report.

Incidental memory performance for pictures that varied along the affective dimensions of pleasantness and arousal was assessed. For both an immediate and delayed (1 year later) free-recall task, only the arousal dimension had a stable effect on memory performance: Pictures rated as highly arousing were remembered better than low-arousal stimuli. This effect was corroborated in a speeded recognition test, in which high-arousal materials encoded earlier in the experiment produced faster reaction times than their low-arousal counterparts. Pleasantness affected reaction time decisions only for pictures not encoded earlier. These results suggest that whereas both the dimensions of pleasantness and arousal are processed at initial encoding, long-term memory performance is mainly affected by arousal.

Two important questions bearing on personality processes and individual differences are how do facial expressiveness and sympathetic activation vary as a function of the intensity of an emotional stimulus, and what is the functional mechanism underlying facial expressiveness and sympathetic activation in emotion? A formulation is proposed that is based on 2 propositions: (a) All strong emotions result in some degree of activation of the organism (i.e., principle of stimulus dynamism) and (b) there are individual differences in the gain (amplification) operating on the facial expressive and sympathetic response channels (i.e., principle of individual response uniqueness). This formulation organizes much of the existing data on internalizers and externalizers and yields novel predictions regarding the subpopulation labeled as generalizers.

Alternative interpretations of startle probe modulation by a pictorial foreground were tested: Either reflex amplitude varies as a function of modality-determined attention allocation, or, regardless of probe modality, reflex amplitude varies with the emotional valence of the foreground content. Thirty-six subjects viewed a series of 54 slides, divided into two 27-slide blocks. Each block consisted of nine exemplars of three independently rated emotional content categories--pleasant, neutral, and unpleasant. Startle probes, half visual (flashgun) and half acoustic (white noise), were presented unpredictably during and between slide presentations. Eyeblink reflexes, corrugator and orbicularis oculi muscle tension, heart rate, and skin conductance were recorded during a 6-s slide interval. Subjects subsequently rated the slides for emotional valence and arousal, and interest value. Free-viewing times were also recorded. Analysis of reflex response and all ancillary measures supported the hypothesis that the primary determinant of startle modulation was the emotional valence of foreground content.

This theoretical model of emotion is based on research using the startle-probe methodology. It explains inconsistencies in probe studies of attention and fear conditioning and provides a new approach to emotional perception, imagery, and memory. Emotions are organized biphasically, as appetitive or aversive (defensive). Reflexes with the same valence as an ongoing emotional state are augmented; mismatched reflexes are inhibited. Thus, the startle response (an aversive reflex) is enhanced during a fear state and is diminished in a pleasant emotional context. This affect-startle effect is not determined by general arousal, simple attention, or probe modality. The effect is found when affects are prompted by pictures or memory images, changes appropriately with aversive conditioning, and may be dependent on right-hemisphere processing. Implications for clinical, neurophysiological, and basic research in emotion are outlined.

Blink reflexes to acoustic probes, heart rate, and subjective reports were studied during affective memory imagery. Thirty-six undergraduates memorized 6 pairs of neutral and fearful sentences. After learning each pair, they relaxed and listened to a series of uniform tones, one every 6 s. A change in tone pitch (higher or lower) cued recall of one of the two sentences. At the first cue tone, groups (n = 12) were under different instructions: (a) ignore the sentence and relax, (b) silently articulate the sentence, and (c) imagine the sentence content as a personal experience. At the second cue tone, all subjects performed the imagery task. Startle probes (50-ms, 95-dB white noise) were presented unpredictably during relaxation and recall trials. Probe blink reflexes were larger and cardiac rate faster at fear sentence recall than at neutral sentence recall or relaxation. For probe reflexes, this effect was greater for imagery than for nonsemantic recall tasks.

A series of four experiments assessed the effects of instructions to lower heart rate on heart rate change and general arousal reduction. Various conditions of biofeedback, cognitive load, incentive, knowledge of results, and the experimenter-subject relationship were tested. Experiment 1 compared physiological responses to the delivery of direct organ feedback (i.e., heart rate) with responses to electromyographic biofeedback from the frontalis muscle area and with responses to a nonfeedback tracking task. The results suggest that neither heart rate nor muscle tension feedback is an especially powerful method for achieving sustained reductions in heart rate. Furthermore, although some specificity of physiologic pattern is apparent, biofeedback is no more effective in lowering general activation level than simple instructions to relax accompanied by a general knowledge of results. The second experiment was designed to assess the role of cognitive load in arousal reduction. Heart rate biofeedback was compared with a procedure involving minimal external information processing--the secular meditation exercise of Wallace and Benson. The results indicated a clear superiority for the meditation strategy in effecting reductions in cardiac rate and lowering activation. However, in a third experiment, meditation subjects lowered heart rate much less than observed in the previous study, and this time the reduction did not exceed that achieved by feedback subjects. Subsequent analysis suggested that the quality of the subject-experimenter relationship (active-supportive vs. formal-distant) was a significant variable in accounting for outcome differences. The above hypothesis was supported by a fourth experiment. Under conditions of high subject-experimenter involvement, the superior meditation performance of Experiment 2 was reproduced; under low-involvement conditions the Experiment 3 result of no difference between training groups was obtained. The findings suggest that the effectiveness of any method for achieving relaxation (or physiological control) rests on a complex interaction between informational and motivational imperatives of the stimulus context and definable aspects of the interpersonal exchange between subject and experimenter. This research raises serious questions about the effectiveness of the usual biofeedback paradigm as an aid to arousal reduction and the cost efficiency of its applications in the clinical situation. Furthermore, these results demonstrate the great power in relaxation experiments of psychosocial and other moderator variables, and signal the practical difficulty of their control when these variables appear to be as potent in changing physiology as the primary training methods.

An abstract of the article "Emotion, Motivation, and the Brain: Reflex Foundations in Animal and Human Research," by Peter J. Lang and Michael Davis is presented.