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To test the effects of cortisol on affective experience, the authors orally administered a placebo, 20 mg cortisol, or 40 mg cortisol to 85 men. Participants' affective responses to negative and neutral stimuli were measured. Self-reported affective state was also assessed. Participants in the 40-mg group (showing extreme cortisol elevations within the physiological range) rated neutral stimuli as more highly arousing than did participants in the placebo and 20-mg groups. Furthermore, within the 20-mg group, individuals with higher cortisol elevations made higher arousal ratings of neutral stimuli. However, cortisol was unrelated to self-reported affective state. Thus, findings indicate that acute cortisol elevations cause heightened arousal in response to objectively nonarousing stimuli, in the absence of effects on mood.
An emotion-modulated acoustic startle paradigm for inducing positive and negative affect was used to address pregoal and postgoal affect. Participants played a computerized lottery task in which they chose digits that could match a subsequently displayed, random set of numbers. In the positive conditions, matches led to monetary rewards. In the negative condition, matches led to an aversive noise blast. In three experiments, we found eyeblink startle magnitude was potentiated just prior to feedback concerning reward outcome, suppressed following the feedback that a monetary reward was won, and potentiated when threatened with an aversive noise. When presented with a 0%, 45%, 90%, or 100% chance of winning, higher probabilities suppressed startle response after feedback whereas the 45% trials did not. These data indicate that postgoal positive affect (winning reward) reliably suppressed the startle response whereas pregoal positive affect did not.
Although the co-occurrence of negative affect and pain is well recognized, the mechanism underlying their association is unclear. To examine whether a common self-regulatory ability impacts the experience of both emotion and pain, we integrated neuroimaging, behavioral, and physiological measures obtained from three assessments separated by substantial temporal intervals. Our results demonstrated that individual differences in emotion regulation ability, as indexed by an objective measure of emotional state, corrugator electromyography, predicted self-reported success while regulating pain. In both emotion and pain paradigms, the amygdala reflected regulatory success. Notably, we found that greater emotion regulation success was associated with greater change of amygdalar activity following pain regulation. Furthermore, individual differences in degree of amygdalar change following emotion regulation were a strong predictor of pain regulation success, as well as of the degree of amygdalar engagement following pain regulation. These findings suggest that common individual differences in emotion and pain regulatory success are reflected in a neural structure known to contribute to appraisal processes.
This study compared the asymmetry of different features of brain electrical activity during the performance of a verbal task (word finding) and a spatial task (dot localization) that had been carefully matched on psychometric properties and accompanying motor activity. Nineteen right-handed subjects were tested. EEG was recorded from F3, F4, C3, C4, P3, and P4, referred to both CZ and computer-derived averaged-ears references, and Fourier transformed. Power in the delta, theta, alpha, and beta bands was computed. There were significant Task X Hemisphere effects in all bands for CZ-referenced data and for the alpha and beta bands for ears-referenced data. The effects were always either greater power suppression in the hemisphere putatively most engaged in task processing or greater power in the opposite hemisphere. Correlations between EEG and task performance indicated that CZ-referenced parietal alpha asymmetry accounted for the most variance in verbal task performance. Power within individual hemispheres or across hemispheres was unrelated to task performance. The findings indicate robust differences in asymmetrical brain physiology that are produced by well-matched verbal and spatial cognitive tasks.
Research on the neural substrates of emotion has found evidence for cortical asymmetries for aspects of emotion. A recent article by Nicholls et al. has used a new imaging method to interrogate facial movement in 3D to assess possible asymmetrical action during expressions of happiness and sadness. Greater left-sided movement, particularly during expressions of sadness was observed. These findings have implications for understanding hemispheric differences in emotion and lend support to the notion that aspects of emotion processing might be differentially localized in the two hemispheres.
Buddhist meditation practices have become a topic of widespread interest in both science and medicine. Traditional Buddhist formulations describe meditation as a state of relaxed alertness that must guard against both excessive hyperarousal (restlessness) and excessive hypoarousal (drowsiness, sleep). Modern applications of meditation have emphasized the hypoarousing and relaxing effects without as much emphasis on the arousing or alertness-promoting effects. In an attempt to counterbalance the plethora of data demonstrating the relaxing and hypoarousing effects of Buddhist meditation, this interdisciplinary review aims to provide evidence of meditation's arousing or wake-promoting effects by drawing both from Buddhist textual sources and from scientific studies, including subjective, behavioral, and neuroimaging studies during wakefulness, meditation, and sleep. Factors that may influence whether meditation increases or decreases arousal are discussed, with particular emphasis on dose, expertise, and contemplative trajectory. The course of meditative progress suggests a nonlinear multiphasic trajectory, such that early phases that are more effortful may produce more fatigue and sleep propensity, while later stages produce greater wakefulness as a result of neuroplastic changes and more efficient processing.
The heart rate, breathing rate, and skin resistance were recorded for 20 community home girls (Home group) and for 20 age-matched girls from a regular school (School group). The former group had a significantly higher rate of breathing and a more irregular breath pattern known to correlate with high fear and anxiety, than the School group. Skin resistance was significantly lower in the School group, which may suggest greater arousal, 28 girls of the Home group formed 14 pairs, matched for age and duration of stay in the home. Subjects of a pair were randomly assigned to either yoga or games groups. For the former emphasis was on relaxation and awareness, whereas for the latter increasing physical activity was emphasized. At the end of an hour daily for six months both groups showed a significant decrease in the resting heart rate relative to initial values (Wilcoxon paired-sample rest), and the yoga group showed a significant decrease in breath rate, which appeared more regular but no significant increase in the skin resistance. These results suggest that a yoga program which includes relaxation, awareness, and graded physical activity is a useful addition to the routine of community home children.
What is compassion? And how did it evolve? In this review, we integrate 3 evolutionary arguments that converge on the hypothesis that compassion evolved as a distinct affective experience whose primary function is to facilitate cooperation and protection of the weak and those who suffer. Our empirical review reveals compassion to have distinct appraisal processes attuned to undeserved suffering; distinct signaling behavior related to caregiving patterns of touch, posture, and vocalization; and a phenomenological experience and physiological response that orients the individual to social approach. This response profile of compassion differs from those of distress, sadness, and love, suggesting that compassion is indeed a distinct emotion. We conclude by considering how compassion shapes moral judgment and action, how it varies across different cultures, and how it may engage specific patterns of neural activation, as well as emerging directions of research.
Facial expression, EEG, and self-report of subjective emotional experience were recorded while subjects individually watched both pleasant and unpleasant films. Smiling in which the muscle that orbits the eye is active in addition to the muscle that pulls the lip corners up (the Duchenne smile) was compared with other smiling in which the muscle orbiting the eye was not active. As predicted, the Duchenne smile was related to enjoyment in terms of occurring more often during the pleasant than the unpleasant films, in measures of cerebral asymmetry, and in relation to subjective reports of positive emotions, and other smiling was not.
Individual variation in the experience and expression of pleasure may relate to differential patterns of lateral frontal activity. Brain electrical measures have been used to study the asymmetric involvement of lateral frontal cortex in positive emotion, but the excellent time resolution of these measures has not been used to capture second-by-second changes in ongoing emotion until now. The relationship between pleasure and second-by-second lateral frontal activity was examined with the use of hierarchical linear modeling in a sample of 128 children ages 6-10 years. Electroencephalographic activity was recorded during "pop-out toy," a standardized task that elicits pleasure. The task consisted of 3 epochs: an anticipation period sandwiched between 2 play periods. The amount of pleasure expressed during the task predicted the pattern of nonlinear change in lateral frontal activity. Children who expressed increasing amounts of pleasure during the task exhibited increasing left lateral frontal activity during the task, whereas children who expressed contentment exhibited increasing right/decreasing left activity. These findings indicate that task-dependent changes in pleasure relate to dynamic, nonlinear changes in lateral frontal activity as the task unfolds.
The authors address 2 questions about embarrassment. First, Is embarrassment a distinct emotion? The evidence indicates that the antecedents, experience, and display of embarrassment, and to a limited extent its autonomic physiology, are distinct from shame, guilt, and amusement and share the dynamic, temporal characteristics of emotion. Second, What are the theoretical accounts of embarrassment? Three accounts focus on the causes of embarrassment, positioning that it follows the loss of self-esteem, concern for others' evaluations, or absence of scripts to guide interactions. A fourth account focuses on the effects of the remedial actions of embarrassment, which correct preceding transgressions. A fifth account focuses on the functional parallels between embarrassment and nonhuman appeasement. The discussion focuses on unanswered questions about embarrassment.
BACKGROUND: Two core characteristics of pathologic fear are its rapid onset and resistance to cognitive regulation. We hypothesized that activation of the amygdala early in the presentation of fear-relevant visual stimuli would distinguish phobics from nonphobics. METHODS: Chronometry of amygdala activation to phobia-relevant pictures was assessed in 13 spider phobics and 14 nonphobics using functional magnetic resonance imaging (fMRI). RESULTS: Blood oxygen level-dependent (BOLD) responses in the amygdala early in picture processing consistently differentiated between phobic and nonphobic subjects, as well as between phobogenic and nonphobogenic stimuli among phobics. Furthermore, amygdalar BOLD responses associated with timing but not magnitude of activation predicted affective responses to phobogenic stimuli. Computational modeling procedures were used to identify patterns of neural activation in the amygdala that could yield the observed BOLD data. These data suggest that phobic responses were characterized by strong but brief amygdala responses, whereas nonphobic responses were weaker and more sustained. CONCLUSIONS: Results are discussed in the context of the amygdala's role in rapid threat detection and the vigilance-avoidance hypothesis of anxiety. These data highlight the importance of examining the neural substrates of the immediate impact of phobogenic stimuli for understanding pathological fear.
We assessed whether resting anterior asymmetry would discriminate individual differences in repressive-defensive coping styles. In 2 sessions, resting electroencephalogram was recorded from female adults during 8 60-s baselines. Subjects were classified as repressors or nonrepressors on the basis of scores on the Marlowe-Crowne Social Desirability Scale (MC), the State-Trait Anxiety Inventory (STAI), and the Beck Depression Inventory (BDI). In midfrontal and lateral frontal sites, repressors demonstrated relative left hemisphere activation when compared with other groups. The MC, but not the STAI or the BDI, contributed unique variance to frontal asymmetry. Relative left frontal activation may be linked to a self-enhancing regulatory style that promotes lowered risk for psychopathology.
Individuals differ dramatically in the quality and intensity of their response to affectively evocative stimuli. On the basis of prior theory and research, we hypothesized that these individual differences are related to variation in activation of the left and right frontal brain regions. We recorded baseline brain electrical activity from subjects on two occasions 3 weeks apart. Immediately following the second recording, subjects were exposed to brief positive and negative emotional film clips. For subjects whose frontal asymmetry was stable across the 3-week period, greater left frontal activation was associated with reports of more intense positive affect in response to the positive films, whereas greater right frontal activation was associated with more intense reports of negative affect in response to the negative film clips. The methodological and theoretical implications of these data are discussed.
The relation between brain activity and the immune system was evaluated by assessing immune responses in 20 healthy women who manifested extreme differences in the asymmetry of frontal cortex activation. One group showed extreme and stable left frontal activation; the other group showed extreme and stable right frontal activation. As predicted, women with extreme right frontal activation had significantly lower levels of natural killer cell activity (at effector:target cell ratios of 33:1 and 11:1) than did left frontally activated individuals. This difference did not extend to two other immune measures, lymphocyte proliferation and T-cell subsets. However, higher immunoglobulin levels of the M class were observed in the right frontal group. In this study, the immune patterns could not be accounted for by plasma cortisol levels, anxiety- and depression-related symptomatology, or recent health histories. These findings support the hypothesis that there is a specific association between frontal brain asymmetry and certain immune responses.
Recent studies have identified a distributed network of brain regions thought to support cognitive reappraisal processes underlying emotion regulation in response to affective images, including parieto-temporal regions and lateral/medial regions of prefrontal cortex (PFC). A number of these commonly activated regions are also known to underlie visuospatial attention and oculomotor control, which raises the possibility that people use attentional redeployment rather than, or in addition to, reappraisal as a strategy to regulate emotion. We predicted that a significant portion of the observed variance in brain activation during emotion regulation tasks would be associated with differences in how participants visually scan the images while regulating their emotions. We recorded brain activation using fMRI and quantified patterns of gaze fixation while participants increased or decreased their affective response to a set of affective images. fMRI results replicated previous findings on emotion regulation with regulation differences reflected in regions of PFC and the amygdala. In addition, our gaze fixation data revealed that when regulating, individuals changed their gaze patterns relative to a control condition. Furthermore, this variation in gaze fixation accounted for substantial amounts of variance in brain activation. These data point to the importance of controlling for gaze fixation in studies of emotion regulation that use visual stimuli.
This research assessed whether individual differences in anterior brain asymmetry are linked to differences in basic dimensions of emotion. In each of 2 experimental sessions, separated by 3 weeks, resting electroencephalogram (EEG) activity was recorded from female adults during 8 60-s baselines. Mean alpha power asymmetry across both sessions was extracted in mid-frontal and anterior temporal sites. Across both regions, groups demonstrating stable and extreme relative left anterior activation reported increased generalized positive affect (PA) and decreased generalized negative affect (NA) compared with groups demonstrating stable and extreme relative right anterior activation. Additional correlational analyses revealed robust relations between anterior asymmetry and PA and NA, particularly among subjects who demonstrated stable patterns of EEG activation over time. Anterior asymmetry was unrelated to individual differences in generalized reactivity.
Freezing is an adaptive defensive behavior that is expressed in response to an imminent threat. In prior studies with rhesus monkeys, stable individual differences in animals' propensities to freeze have been demonstrated. To understand the factors associated with these individual differences, freezing behavior was examined in infant rhesus monkeys and their mothers, in conjunction with levels of the stress-related hormone cortisol. In both mothers and infants, basal cortisol levels were positively correlated with freezing duration. Additionally, the number of offspring a mother had was negatively correlated with her infant's cortisol level. These findings suggest a link between basal cortisol levels and an animal's propensity to freeze, as well as a mechanism by which maternal experience may affect infants' cortisol levels.
Prior studies assessing the relation between negative affective traits and cortisol have yielded inconsistent results. Two studies assessed the relation between individual differences in repressive-defensiveness and basal salivary cortisol levels. Experiment 1 assessed midafternoon salivary cortisol levels in men classified as repressors, high-anxious, or low-anxious. In Experiment 2, more rigorous controls were applied as salivary cortisol levels in women and men were assessed at 3 times of day on 3 separate days. In both studies, as hypothesized, repressors and high-anxious participants demonstrated higher basal cortisol levels than low-anxious participants. These findings suggest that both heightened distress and the inhibition of distress may be independently linked to relative elevations in cortisol. Also discussed is the possible mediational role of individual differences in responsivity to, or mobilization for, uncertainty or change.
The present study investigated the premise that individual differences in autonomic physiology could be used to specify the nature and consequences of information processing taking place in medial prefrontal regions during cognitive reappraisal of unpleasant pictures. Neural (blood oxygenation level-dependent functional magnetic resonance imaging) and autonomic (electrodermal [EDA], pupil diameter, cardiac acceleration) signals were recorded simultaneously as twenty-six older people (ages 64-66 years) used reappraisal to increase, maintain, or decrease their responses to unpleasant pictures. EDA was higher when increasing and lower when decreasing compared to maintaining. This suggested modulation of emotional arousal by reappraisal. By contrast, pupil diameter and cardiac acceleration were higher when increasing and decreasing compared to maintaining. This suggested modulation of cognitive demand. Importantly, reappraisal-related activation (increase, decrease>maintain) in two medial prefrontal regions (dorsal medial frontal gyrus and dorsal cingulate gyrus) was correlated with greater cardiac acceleration (increase, decrease>maintain) and monotonic changes in EDA (increase>maintain>decrease). These data indicate that these two medial prefrontal regions are involved in the allocation of cognitive resources to regulate unpleasant emotion, and that they modulate emotional arousal in accordance with the regulatory goal. The emotional arousal effects were mediated by the right amygdala. Reappraisal-related activation in a third medial prefrontal region (subgenual anterior cingulate cortex) was not associated with similar patterns of change in any of the autonomic measures, thus highlighting regional specificity in the degree to which cognitive demand is reflected in medial prefrontal activation during reappraisal.
Based on previous findings in humans and rhesus monkeys suggesting that diazepam has asymmetrical effects on frontal lobe activity and other literature supporting a role for the benzodiazepine system in the mediation of individual differences in anxiety and fearfulness, the relation between asymmetrical changes in scalp-recorded regional brain activity in response to diazepam and the temperamental dimension of behavioral inhibition indexed by freezing time in 9 rhesus monkeys was examined. Animals showed greater relative left-sided frontal activation in response to diazepam compared with the preceding baseline. The magnitude of this shift was strongly correlated with an aggregate measure of freezing time (r = .82). The implications of these findings for understanding the role of regional differences in the benzodiazepine system in mediating individual differences in fearfulness are discussed.
Baseline resting electroencephalogram activity was recorded with 3 different reference montages from 15 clinically depressed and 13 control subjects. Power in all frequency bands was extracted by fast Fourier transformation. There was a significant Group X Hemisphere interaction in the mid-frontal region, for the alpha band power only. Depressed subjects had less left-sided activation (i.e., more alpha activity) than did normal control subjects. This pattern of diminished left-sided frontal activation is interpreted as indicating a deficit in approach mechanisms in depressed subjects.
OBJECTIVE: Positron emission tomography was used to investigate the neural substrates of normal human emotional and their dependence on the types of emotional stimulus. METHOD: Twelve healthy female subjects underwent 12 measurements of regional brain activity following the intravenous bolus administration of [15O]H2O as they alternated between emotion-generating and control film and recall tasks. Automated image analysis techniques were used to characterize and compare the increases in regional brain activity associated with the emotional response to complex visual (film) and cognitive (recall) stimuli. RESULTS: Film- and recall-generated emotion were each associated with significantly increased activity in the vicinity of the medial prefrontal cortex and thalamus, suggesting that these regions participate in aspects of emotion that do not depend on the nature of the emotional stimulus. Film-generated emotion was associated with significantly greater increases in activity bilaterally in the occipitotemporparietal cortex, lateral cerebellum, hypothalamus, and a region that includes the anterior temporal cortex, amygdala, and hippocampal formation, suggesting that these regions participate in the emotional response to certain exteroceptive sensory stimuli. Recall-generated sadness was associated with significantly greater increases in activity in the vicinity of the anterior insular cortex, suggesting that this region participates in the emotional response to potentially distressing cognitive or interoceptive sensory stimuli. CONCLUSIONS: While this study should be considered preliminary, it identified brain regions that participate in externally and internally generated human emotion.
In this article, the authors elaborate on 3 ideas advanced in P. Rozin and A. B. Cohen's (2003) innovative study of facial expression. Taking a cue from their discovery of new expressive behaviors (e.g., the narrowed eyebrows), the authors review recent studies showing that emotions are conveyed in more channels than usually studied, including posture, gaze patterns, voice, and touch. Building on their claim that confusion has a distinct display, the authors review evidence showing distinct displays for 3 self-conscious emotions (embarrassment, shame, and pride), 5 positive emotions (amusement, desire, happiness, love, interest), and sympathy and compassion. Finally, the authors offer a functional definition of emotion to integrate these findings on "new" displays and emotions.
Positive affect elicited in a mother toward her newborn infant may be one of the most powerful and evolutionarily preserved forms of positive affect in the emotional landscape of human behavior. This study examined the neurobiology of this form of positive emotion and in so doing, sought to overcome the difficulty of eliciting robust positive affect in response to visual stimuli in the physiological laboratory. Six primiparous human mothers with no indications of postpartum depression brought their infants into the laboratory for a photo shoot. Approximately 6 weeks later, they viewed photographs of their infant, another infant, and adult faces during acquisition of functional magnetic resonance images (fMRI). Mothers exhibited bilateral activation of the orbitofrontal cortex (OFC) while viewing pictures of their own versus unfamiliar infants. While in the scanner, mothers rated their mood more positively for pictures of their own infants than for unfamiliar infants, adults, or at baseline. The orbitofrontal activation correlated positively with pleasant mood ratings. In contrast, areas of visual cortex that also discriminated between own and unfamiliar infants were unrelated to mood ratings. These data implicate the orbitofrontal cortex in a mother's affective responses to her infant, a form of positive emotion that has received scant attention in prior human neurobiological studies. Furthermore, individual variations in orbitofrontal activation to infant stimuli may reflect an important dimension of maternal attachment.
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