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.
The nature of the affective deficit that characterizes social anhedonia is not well understood. Emotionally evocative visual stimuli were presented to undergraduates identified as anhedonic or normal, based on their scores on the revised Social Anhedonia Scale. The affective stimuli were chosen to elicit positive and negative emotion; a subset of slides were specifically chosen to include social-interpersonal content. In the acoustic startle paradigm, participants were administered startle probes (50-ms 95 dB white noise bursts) while viewing images from the International Affective Picture System. Socially anhedonic individuals did not differ from normally hedonic individuals in terms of their physiological response to the stimuli, regardless of the nature of the content of the stimuli. However, on the self-report measures of trait affectivity, the socially anhedonic individuals reported significantly lower levels of positive affect and higher levels of negative affect. These findings suggest that the affective deficits reported by socially anhedonic individuals are not global in nature.
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.
Social relations between humans critically depend on our affective experiences of others. Oxytocin enhances prosocial behavior, but its effect on humans' affective experience of others is not known. We tested whether oxytocin influences affective ratings, and underlying brain activity, of faces that have been aversively conditioned. Using a standard conditioning procedure, we induced differential negative affective ratings in faces exposed to an aversive conditioning compared with nonconditioning manipulation. This differential negative evaluative effect was abolished by treatment with oxytocin, an effect associated with an attenuation of activity in anterior medial temporal and anterior cingulate cortices. In amygdala and fusiform gyrus, this modulation was stronger for faces with direct gaze, relative to averted gaze, consistent with a relative specificity for socially relevant cues. The data suggest that oxytocin modulates the expression of evaluative conditioning for socially relevant faces via influences on amygdala and fusiform gyrus, an effect that may explain its prosocial effects.
Examined were electroencephalogram (EEG) asymmetries during the presence of discrete facial signs of emotion among 10-month-old infants who were tested in a standard stranger- and mother-approach paradigm that included a brief separation from mother. Data underscore the usefulness of EEG measures of hemispheric activation in differentiating among certain emotional states. (RH)
<p>Measures of left-right asymmetry in resting brain activity were derived from spectral estimates of electroencephalogram (EEG) alpha-band power density in 13 homologous scalp electrode pairs from 81 right-handed individuals (43 F) on two occasions separated by 6 weeks. At a third, later session, these individuals completed a cognitive task, comparing word-pairs that systematically differed in affective tone. For an extended series of paired-comparisons, the subject chose the one word-pair that 'went together best'. Objectively, associative strength was comparable for both word-pairs. Individuals with relatively greater left-sided anterior frontal resting activity were more likely to select the more pleasant word-pair. Relations between word-pair selection and asymmetry in resting brain activity at central and posterior sites were not significant.</p>
Significant progress has been made in our understanding of the neural substrates of emotion and its disorders. Neuroimaging methods have been used to characterize the circuitry underlying disorders of emotion. Particular emphasis has been placed on the prefrontal cortex, anterior cingulate, parietal cortex, and the amygdala as critical components of the circuitry that may be dysfunctional in both depression and anxiety.
Although once considered disruptive, self-conscious emotions are now theorized to be fundamentally involved in the regulation of social behavior. The present study examined the social regulation function of self-conscious emotions by comparing healthy participants with a neuropsychological population--patients with orbitofrontal lesions--characterized by selective regulatory deficits. Orbitofrontal patients and healthy controls participated in a series of tasks designed to assess their social regulation and self-conscious emotions. Another task assessed the ability to infer others' emotional states, an appraisal process involved in self-conscious emotion. Consistent with the theory that self-conscious emotions are important for regulating social behavior, the findings show that deficient behavioral regulation is associated with inappropriate self-conscious emotions that reinforce maladaptive behavior. Additionally, deficient behavioral regulation is associated with impairments in interpreting the self-conscious emotions of others.
OBJECTIVE Deficits in positive affect and their neural bases have been associated with major depression. However, whether reductions in positive affect result solely from an overall reduction in nucleus accumbens activity and fronto-striatal connectivity or the additional inability to sustain engagement of this network over time is unknown. The authors sought to determine whether treatment-induced changes in the ability to sustain nucleus accumbens activity and fronto-striatal connectivity during the regulation of positive affect are associated with gains in positive affect. METHOD Using fMRI, the authors assessed the ability to sustain activity in reward-related networks when attempting to increase positive emotion during performance of an emotion regulation paradigm in 21 depressed patients before and after 2 months of antidepressant treatment. Over the same interval, 14 healthy comparison subjects underwent scanning as well. RESULTS After 2 months of treatment, self-reported positive affect increased. The patients who demonstrated the largest increases in sustained nucleus accumbens activity over the 2 months were those who demonstrated the largest increases in positive affect. In addition, the patients who demonstrated the largest increases in sustained fronto-striatal connectivity were also those who demonstrated the largest increases in positive affect when controlling for negative affect. None of these associations were observed in healthy comparison subjects. CONCLUSIONS Treatment-induced change in the sustained engagement of fronto-striatal circuitry tracks the experience of positive emotion in daily life. Studies examining reduced positive affect in a variety of psychiatric disorders might benefit from examining the temporal dynamics of brain activity when attempting to understand changes in daily positive affect.
Depression has been associated with dysfunctional executive functions and abnormal activity within the anterior cingulate cortex (ACC), a region critically involved in action regulation. Prior research invites the possibility that executive deficits in depression may arise from abnormal responses to negative feedback or errors, but the underlying neural substrates remain unknown. We hypothesized that abnormal reactions to error would be associated with dysfunctional rostral ACC activity, a region previously implicated in error detection and evaluation of the emotional significance of events. To test this hypothesis, subjects with low and high Beck Depression Inventory (BDI) scores performed an Eriksen Flanker task. To assess whether tonic activity within the rostral ACC predicted post-error adjustments, 128-channel resting EEG data were collected before the task and analyzed with low-resolution electromagnetic tomography (LORETA) using a region-of-interest approach. High BDI subjects were uniquely characterized by significantly lower accuracy after incorrect than correct trials. Mirroring the behavioral findings, high BDI subjects had significantly reduced pretask gamma (36.5-44 Hz) current density within the affective (rostral; BA24, BA25, BA32) but not cognitive (dorsal; BA24', BA32') ACC subdivision. For low, but not high, BDI subjects pretask gamma within the affective ACC subdivision predicted post-error adjustments even after controlling for activity within the cognitive ACC subdivision. Abnormal responses to errors may thus arise due to lower activity within regions subserving affective and/or motivational responses to salient cues. Because rostral ACC regions have been implicated in treatment response in depression, our findings provide initial insight into putative mechanisms fostering treatment response.
This article assessed whether resting electroencephalographic (EEG) asymmetry in anterior regions of the brain can predict affective responses to emotion elicitors. Baseline EEG was recorded from 32 female adults, after which Ss viewed film clips preselected to elicit positive or negative affect. Resting alpha power asymmetry in the frontal region significantly predicted self-reported global negative affect in response to clips and predicted the difference between global positive and negative affect. Analyses of discrete emotions revealed a strong relation between frontal asymmetry and fear responses to films. Effects were independent of Ss mood ratings at the time at which baseline EEG was measured. Resting anterior asymmetry may be a state-independent index of the individual's predisposition to respond affectively.
Resting respiratory sinus arrhythmia (RSAREST) indexes important aspects of individual differences in emotionality. In the present investigation, the authors address whether RSAREST is associated with tonic positive or negative emotionality, and whether RSAREST relates to phasic emotional responding to discrete positive emotion-eliciting stimuli. Across an 8-month, multiassessment study of first-year university students (n = 80), individual differences in RSAREST were associated with positive but not negative tonic emotionality, assessed at the level of personality traits, long-term moods, the disposition toward optimism, and baseline reports of current emotional states. RSAREST was not related to increased positive emotion, or stimulus-specific emotion, in response to compassion-, awe-, or pride-inducing stimuli. These findings suggest that resting RSA indexes aspects of a person's tonic positive emotionality.
Although several studies have examined anterior asymmetric brain electrical activity and cortisol in infants, children, and adults, the direct association between asymmetry and cortisol has not systematically been reported. In nonhuman primates, greater relative right anterior activation has been associated with higher cortisol levels. The current study examines the relation between frontal electroencephalographic (EEG) asymmetry and cortisol (basal and reactive) and withdrawal-related behaviors (fear and sadness) in 6-month-old infants. As predicted, the authors found that higher basal and reactive cortisol levels were associated with extreme right EEG asymmetry. EEG during the withdrawal-negative affect task was associated with fear and sadness behaviors. Results are interpreted in the context of the previous primate work, and some putative mechanisms are discussed.
Drawing on recent claims in the study of relationships, attachment, and emotion, the authors hypothesized that romantic love serves a commitment-related function and sexual desire a reproduction-related function. Consistent with these claims, in Study 1, brief experiences of romantic love and sexual desire observed in a 3-min interaction between romantic partners were related to distinct feeling states, distinct nonverbal displays, and commitment- and reproductive-related relationship outcomes, respectively. In Study 2, the nonverbal display of romantic love was related to the release of oxytocin. Discussion focuses on the place of romantic love and sexual desire in the literature on emotion.
<p>Appropriate social problem solving constitutes a critical skill for individuals and may rely on processes important for self-generated thought (SGT). The aim of the current study was to investigate the link between SGT and social problem solving. Using the Means-End Problem Solving task (MEPS), we assessed participants' abilities to resolve daily social problems in terms of overall efficiency and number of relevant means they provided to reach the given solution. Participants also performed a non-demanding choice reaction time task (CRT) and a moderately-demanding working memory task (WM) as a context in which to measure their SGT (assessed via thought sampling). We found that although overall SGT was associated with lower MEPS efficiency, it was also associated with higher relevant means, perhaps because both depend on the capacity to generate cognition that is independent from the hear and now. The specific content of SGT did not differentially predict individual differences in social problem solving, suggesting that the relationship may depend on SGT regardless of its content. In addition, we also found that performance at the WM but not the CRT was linked to overall better MEPS performance, suggesting that individuals good at social processing are also distinguished by their capacity to constrain attention to an external task. Our results provide novel evidence that the capacity for SGT is implicated in the process by which solutions to social problems are generated, although optimal problem solving may be achieved by individuals who display a suitable balance between SGT and cognition derived from perceptual input.</p>
This brief commentary highlights seven sins in the study of emotion that are explicitly treated in contemporary affective neuroscience. These sins are (1) Affect and cognition are subserved by separate and independent neural circuits; (2) Affect is subcortical; (3) Emotions are in the head; (4) Emotions can be studied from a purely psychological perspective; (5) Emotions are similar in structure across age and species; (6) Specific emotions are instantiated in discrete locations in the brain; and (7) Emotions are conscious feeling states. Each of these is briefly discussed and evidence from affective neuroscience that bears on these sins is noted. The articles in this Special Issue underscore the vitality of research in affective neuroscience and illustrate how some of these sins can be addressed and rectified using concepts and methods from affective neuroscience.
This paper reports three studies showing sex differences in EEG asymmetry during self-generated cognitive and affective tasks. In the first experiment, bilateral EEG, quantified for alpha on-line, was recorded from right-handed subjects while they either whistled, sang or recited lyrics of familiar songs. The results revealed significant asymmetry between the whistle and talk conditions only for subjects with no familial left-handedness and, within this group, only for females and not for males. In the second experiment, bilateral EEG was recorded while right-handed subjects (with no familial left-handedness) self-induced covert affective and non-affective states. Results revealed significantly greater relative right-hemisphere activation during emotion versus non-emotion trials only in females; males showed no significant task-dependent shifts in asymmetry between conditions. The third experiment was designed to test the hypothesis that females show greater percent time asymmetry than males during biofeedback training for symmetrical and asymmetrical EEG patterns. Results confirmed this prediction as well as indicating that females show better control of such asymmetrical cortical patterning. These findings provide new neuropsychological support for the hypothesis of greater bilateral flexibility in females during self-generation tasks.
The development of functional neuroimaging of emotion holds the promise to enhance our understanding of the biological bases of affect and improve our knowledge of psychiatric diseases. However, up to this point, researchers have been unable to objectively, continuously and unobtrusively measure the intensity and dynamics of affect concurrently with functional magnetic resonance imaging (fMRI). This has hindered the development and generalizability of our field. Facial electromyography (EMG) is an objective, reliable, valid, sensitive, and unobtrusive measure of emotion. Here, we report the successful development of a method for simultaneously acquiring fMRI and facial EMG. The ability to simultaneously acquire brain activity and facial physiology will allow affective neuroscientists to address theoretical, psychiatric, and individual difference questions in a more rigorous and generalizable way.
OBJECTIVES: Affective neuroscience research that investigates core symptoms of pediatric bipolar disorder (PBD) may be effective in differentiating PBD phenotypes. The current study used affect-modulated startle to examine potential differences in reactivity to emotional stimuli (reward and punishment) in narrow and broad phenotype PBD and controls. METHODS: Thirty children meeting DSM-IV bipolar disorder criteria (i.e. narrow phenotype PBD with defined manic episodes with elevated/expansive mood), 19 children meeting criteria for severe mood dysregulation (i.e. broad phenotype with chronic irritability, hyper-reactivity, and hyperarousal), and 19 controls completed a lottery startle paradigm involving reward (money) and punishment (loud noise). Startle probes were presented during anticipation of the emotional stimulus, immediately following the presentation of the stimulus, or during return to baseline following the stimulus. RESULTS: By self-report, patients and controls found the putative punishment to be preferable to the neutral condition. In the reward condition, patient samples reported greater arousal than did controls, but no between-group differences were found on the magnitude of startle response during the reward, punishment, or neutral conditions. CONCLUSIONS: The failure to find differences in affect-modulated startle between control children and those with narrow or broad PBD phenotypes speaks to the methodological challenges associated with studying reward mechanisms in PBD. Alternative paradigms that focus on different aspects of reward mechanisms are discussed.
Facial expressions of pain are an important part of the pain response, signaling distress to others and eliciting social support. To evaluate how voluntary modulation of this response contributes to the pain experience, 29 subjects were exposed to thermal stimulation while making standardized pain, control, or relaxed faces. Dependent measures were self-reported negative effect (valence and arousal) as well as the intensity of nociceptive stimulation required to reach a given subjective level of pain. No direct social feedback was given by the experimenter. Although the amount of nociceptive stimulation did not differ across face conditions, subjects reported more negative effects in response to painful stimulation while holding the pain face. Subsequent analyses suggested the effects were not due to preexisting differences in the difficulty or unpleasantness of making the pain face. These results suggest that voluntary pain expressions have no positively reinforcing (pain attenuating) qualities, at least in the absence of external contingencies such as social reinforcement, and that such expressions may indeed be associated with higher levels of negative affect in response to similar nociceptive input. PERSPECTIVE: This study demonstrates that making a standardized pain face increases negative affect in response to nociceptive stimulation, even in the absence of social feedback. This suggests that exaggerated facial displays of pain, although often socially reinforced, may also have unintended aversive consequences.
This commentary provides reflections on the current state of affairs in research on EEG frontal asymmetries associated with affect. Although considerable progress has occurred since the first report on this topic 25 years ago, research on frontal EEG asymmetries associated with affect has largely evolved in the absence of any serious connection with neuroscience research on the structure and function of the primate prefrontal cortex (PFC). Such integration is important as this work progresses since the neuroscience literature can help to understand what the prefrontal cortex is "doing" in affective processing. Data from the neuroscience literature on the heterogeneity of different sectors of the PFC are introduced and more specific hypotheses are offered about what different sectors of the PFC might be doing in affect. A number of methodological issues associated with EEG measures of functional prefrontal asymmetries are also considered.
For survivors of childhood sexual abuse (CSA), verbal disclosure is often complex and painful. The authors examined the voluntary disclosure-nondisclosure of CSA in relation to nonverbal expressions of emotion in the face. Consistent with hypotheses derived from recent theorizing about the moral nature of emotion, CSA survivors who did not voluntarily disclose CSA showed greater facial expressions of shame, whereas CSA survivors who voluntarily disclosed CSA expressed greater disgust. Expressions of disgust also signaled sexual abuse accompanied by violence. Consistent with recent theorizing about smiling behavior, CSA nondisclosers made more polite smiles, whereas nonabused participants expressed greater genuine positive emotion. Discussion addressed the implications of these findings for the study of disclosure of traumatic events, facial expression, and the links between morality and emotion.