Motivational engagement in Parkinson's disease: Preparation for motivated action.

The current study investigated whether motivational dysfunction in Parkinson's patients is related to a deficit in preparing for motivated behavior. Based on previous studies, it was hypothesized that PD patients would show reduced preparation for action specifically when faced with threat (of loss) and that reduced action preparation would relate to self-report of apathy symptoms. The study measured an electrocortical correlate of preparation for action (CNV amplitude) in PD patients and healthy controls, as well as defensive and appetitive activation during emotional perception (LPP amplitude). The sample included 18 non-demented PD patients (tested on dopaminergic medications) and 15 healthy controls who responded as quickly as possible to cues signaling threat of loss or reward, in which the speed of the response determined the outcome. Results indicated that, whereas PD patients showed similar enhanced action preparation with the addition of incentives to controls, PD patients showed generally reduced action preparation, evidenced by reduced CNV amplitude overall. Results suggest that PD patients may have behavioral issues due to globally impaired action preparation but that this deficit is not emotion-specific, and movement preparation may be aided by incentive in PD patients.

The late positive potential, emotion and apathy in Parkinson's disease.

Parkinson's disease is associated with emotional changes including depression, apathy, and anxiety. The current study investigated emotional processing in non-demented individuals with Parkinson disease (PD) using an electrophysiological measure, the centro-parietal late positive potential (LPP). Non-demented patients with Parkinson's disease (n=17) and healthy control participants (n=16) viewed pleasant, neutral, and unpleasant pictures while EEG was recorded from a 64-channel geodesic net. The Parkinson patients did not differ from controls in terms of early electrophysiological components that index perceptual processing (occipital P100, N150, P250). Parkinson patients, however, showed reduced LPP amplitude specifically when viewing unpleasant, compared to pleasant, pictures as well as when compared to controls, consistent with previous studies suggesting a specific difference in aversive processing between PD patients and healthy controls. Importantly, LPP amplitude during unpleasant picture viewing was most attenuated for patients reporting high apathy. The data suggest that apathy in PD may be related to a deficit in defensive activation, and may be indexed cortically using event-related potentials.

Emotion and ocular responses in Parkinson's disease.

Parkinson's disease (PD) is a neurodegenerative disease that affects motor, cognitive, and emotional functioning. Previous studies reported reduced skin conductance responses in PD patients, compared to healthy older adults when viewing emotionally arousing pictures. Attenuated skin conductance changes in PD may reflect peripheral autonomic dysfunction (e.g., reduced nerve endings at the sweat gland) or, alternatively, a more central emotional deficit. The aim of the current study was to investigate a second measure of sympathetic arousal-change in pupil dilation. Eye movements, a motor-based correlate of emotional processing, were also assessed. Results indicated that pupil dilation was significantly greater when viewing emotional, compared to neutral pictures for both PD patients and controls. On the other hand, PD patients made fewer fixations with shorter scan paths, particularly when viewing pleasant pictures. These results suggest that PD patients show normal sympathetic arousal to affective stimuli (indexed by pupil diameter), but differences in motor correlates of emotion (eye movements).

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.

Food deprivation and emotional reactions to food cues: implications for eating disorders.

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.

Affective reactions to briefly presented pictures.

Affective reactions to briefly presented pictures were investigated to determine whether fleeting stimuli engage the motivational systems mediating emotional responses. Emotional and neutral pictures were presented for 500 ms; heart rate, skin conductance, corrugator EMG, and the evoked startle reflex were measured. The time course of reflex modulation was similar to that obtained with longer (6 s) presentations, suggesting that picture processing continues in the absence of a sensory stimulus. Affective reactions found with more sustained presentation were also obtained, with more corrugator EMG activity for unpleasant pictures, and greater skin conductance reactivity for emotional pictures. Heart rate modulation, however, appears to rely on the presence of a sensory stimulus. The data also suggest that brief presentations of unpleasant pictures may result in less defensive activation than sustained presentation.

Fleeting images: a new look at early emotion discrimination.

The visual brain quickly sorted stimuli for emotional impact despite high-speed presentation (3 or 5 per s) in a sustained, serial torrent of 700 complex pictures. Event-related potentials, recorded with a dense electrode array, showed selective discrimination of emotionally arousing stimuli from less affective content. Primary sources of this activation were over the occipital cortices, extending to right parietal cortex, suggesting a processing focus in the posterior visual system. Emotion discrimination was independent of formal pictorial properties (color, brightness. spatial frequency, and complexity). The data support the hypothesis of a very short-term conceptual memory store (M. C. Potter, 1999)-shown here to include a fleeting but reliable assessment of affective meaning.

Emotional reactions in children: verbal, physiological, and behavioral responses to affective pictures.

Many studies have shown a consistent pattern in adults' responses to affective pictures and there is growing evidence of gender differences, as well. Little is known, though, about children's verbal, behavioral, and physiological responses to affective pictures. Two experiments investigated children's responses to pictures. In Experiment 1, children, adolescents, and adults viewed pictures varying in affective content and rated them for pleasure, arousal, and dominance. Results indicated that children and adolescents rated the pictures similarly to adults. In Experiment 2, physiological responses, self-report, and viewing time were measured while children viewed affective pictures. As with adults, children's responses reflected the affective content of the pictures. Gender differences in affective evaluations, corrugator activity, skin conductance, startle modulation, and viewing time indicated that girls were generally more reactive to unpleasant materials.

Emotion and motivation I: defensive and appetitive reactions in picture processing.

Emotional reactions are organized by underlying motivational states--defensive and appetitive--that have evolved to promote the survival of individuals and species. Affective responses were measured while participants viewed pictures with varied emotional and neutral content. Consistent with the motivational hypothesis, reports of the strongest emotional arousal, largest skin conductance responses, most pronounced cardiac deceleration, and greatest modulation of the startle reflex occurred when participants viewed pictures depicting threat, violent death, and erotica. Moreover, reflex modulation and conductance change varied with arousal, whereas facial patterns were content specific. The findings suggest that affective responses serve different functions-mobilization for action, attention, and social communication-and reflect the motivational system that is engaged, its intensity of activation, and the specific emotional context.

Emotion and motivation II: sex differences in picture processing.

Adhering to the view that emotional reactivity is organized in part by underlying motivational states--defensive and appetitive--we investigated sex differences in motivational activation. Men's and women's affective reactions were measured while participants viewed pictures with varied emotional and neutral content. As expected, highly arousing contents of threat, mutilation, and erotica prompted the largest affective reactions in both men and women. Nonetheless, women showed a broad disposition to respond with greater defensive reactivity to aversive pictures, regardless of specific content, whereas increased appetitive activation was apparent for men only when viewing erotica. Biological and sociocultural factors in shaping sex differences in emotional reactivity are considered as possible mediators of sex differences in emotional response.

The psychopath as observer: emotion and attention in picture processing.

This study extended prior work showing abnormal affect-startle modulation in psychopaths. Male prisoners viewed specific categories of pleasant (erotic or thrilling) and unpleasant (victim or direct threat) slide pictures, along with neutral pictures. Acoustic startle probes were presented early (300 and 800 ms) and late (1,800, 3,000, and 4,500 ms) in the viewing interval. At later times, nonpsychopaths showed moderate and strong reflex potentiation for victim and threat scenes, respectively. For psychopaths, startle was inhibited during victim scenes and only weakly potentiated during threat. Psychopaths also showed more reliable blink inhibition across pleasant contents than nonpsychopaths and greater heart rate orienting to affective pictures overall. These results indicate a heightened aversion threshold in psychopaths. In addition, deficient reflex modulation at early times suggested a weakness in initial stimulus evaluation among psychopaths.

Affective reactions to acoustic stimuli.

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.

Affective picture processing: the late positive potential is modulated by motivational relevance.

Recent studies have shown that the late positive component of the event-related-potential (ERP) is enhanced for emotional pictures, presented in an oddball paradigm, evaluated as distant from an established affective context. In other research, with context-free, random presentation, affectively intense pictures (pleasant and unpleasant) prompted similar enhanced ERP late positivity (compared with the neutral picture response). In an effort to reconcile interpretations of the late positive potential (LPP), ERPs to randomly ordered pictures were assessed, but using the faster presentation rate, brief exposure (1.5 s), and distinct sequences of six pictures, as in studies using an oddball based on evaluative distance. Again, results showed larger LPPs to pleasant and unpleasant pictures, compared with neutral pictures. Furthermore, affective pictures of high arousal elicited larger LPPs than less affectively intense pictures. The data support the view that late positivity to affective pictures is modulated both by their intrinsic motivational significance and the evaluative context of picture presentation.

Brain potentials in affective picture processing: covariation with autonomic arousal and affective report.

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.

Emotion, motivation, and anxiety: brain mechanisms and psychophysiology.

The organization of response systems in emotion is founded on two basic motive systems, appetitive and defensive. The subcortical and deep cortical structures that determine primary motivated behavior are similar across mammalian species. Animal research has illuminated these neural systems and defined their reflex outputs. Although motivated behavior is more complex and varied in humans, the simpler underlying response patterns persist in affective expression. These basic phenomena are elucidated here in the context of affective perception. Thus, the research examines human beings watching uniquely human stimuli--primarily picture media (but also words and sounds) that prompt emotional arousal--showing how the underlying motivational structure is apparent in the organization of visceral and behavioral responses, in the priming of simple reflexes, and in the reentrant processing of these symbolic representations in the sensory cortex. Implications of the work for understanding pathological emotional states are discussed, emphasizing research on psychopathy and the anxiety disorders.

Startle potentiation: shock sensitization, aversive learning, and affective picture modulation.

This study investigated the size of, and relationship between, different modulatory effects of aversive stimulation on the acoustic startle reflex. This reflex is potentiated by shock exposure and associative shock conditioning (in animals and human volunteers) and unpleasant pictures (in human volunteers). In this study, dramatic sensitization of the probe-startle response was observed after shock exposure but not after a control task. Magnitude of sensitization was significantly larger than associative shock conditioning and picture modulation effects (also significant). Sensitization and conditioning scores showed modest, significant correlations with one another but not with picture modulation scores, consistent with animal data showing that partially overlapping brain mechanisms (i.e., amygdaloid-reticular projections) mediate these effects. The present results also indicate that sensitization of startle in human volunteers is a relatively more robust defensive response to aversive stimulation.

Emotional arousal and activation of the visual cortex: an fMRI analysis.

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.

Neuroanatomical correlates of pleasant and unpleasant emotion.

Substantial evidence suggests that a key distinction in the classification of human emotion is that between an appetitive motivational system association with positive or pleasant emotion and an aversive motivational system associated with negative or unpleasant emotion. To explore the neural substrates of these two systems, 12 healthy women viewed sets of pictures previously demonstrated to elicit pleasant, unpleasant and neutral emotion, while positron emission tomographic (PET) measurements of regional cerebral blood flow were obtained. Pleasant and unpleasant emotions were each distinguished from neutral emotion conditions by significantly increased cerebral blood flow in the vicinity of the medial prefrontal cortex (Brodmann's area 9), thalamus, hypothalamus and midbrain (P < 0.005). Unpleasant was distinguished from neutral or pleasant emotion by activation of the bilateral occipito-temporal cortex and cerebellum, and left parahippocampal gyrus, hippocampus and amygdala (P < 0.005). Pleasant was also distinguished from neutral but not unpleasant emotion by activation of the head of the left caudate nucleus (P < 0.005). These findings are consistent with those from other recent PET studies of human emotion and demonstrate that there are both common and unique components of the neural networks mediating pleasant and unpleasant emotion in healthy women.