The temporal dynamics of volitional emotion regulation

Happiness, anger, surprise, irritation… if we note down the emotions that we go through on a given day, the list will most probably be quite long. A surge of studies on the bidirectional interaction between emotion and cognition suggests that we need emotional appraisals in order to lead a successful life and maintain our personal, social and economic integrity (Bechara, 2005; Damasio, 1994; Fox, 2008; Gross & Thompson, 2007; Walter, 2005). And yet, we seldom ‘just’ experience emotions, but often try to influence them to best fit our current goals. Based on the assumption that emotional reactions entail changes on various levels, and that these changes happen in- or outside of our awareness, affective science has adopted emotion regulation as one of its major research topics (Beauregard, Levesque, & Paquette, 2004; Gross, 1999; Ochsner, 2007). In fact, neural (e.g. amygdala activation) and behavioral (e.g. feeling of negativity) correlates of emotional reactions are effectively reduced by top-down processes of explicit and implicit control (Drabant, McRae, Manuck, Hariri, & Gross, 2009; Levesque, et al., 2003; Ochsner, Ray, et al., 2004). Furthermore, evidence from studies investigating voluntary thought control suggests that control strategies may have lasting and paradoxical consequences (Abramowitz, Tolin, & Street, 2001; Wegner, 2009). In a very recent investigation, lasting effects of regulation were also shown after the cognitive control of emotions: the activation timecourse of the amygdala was significantly increased immediately following regulation, and this difference was also related to the activation of the amygdala to the same stimuli a few minutes later (Walter, et al., 2009). Aside from these contextual or qualitative influences, emotional processing also differs between individuals: genetic variation within the serotonergic system for instance is known to affect emotional reactivity both on the behavioral and on the neural level (Hariri, et al., 2005; Hariri, et al., 2002; Lesch, et al., 1996). In the present work, the temporal dynamics of volitional emotion regulation were investigated in three studies. It was hypothesized that both the subjective experience of negativity and the amygdala activation can be attenuated by the detachment from negative emotions, which in turn leads to an immediate neural aftereffect after the offset of regulation. Furthermore, volitional emotion regulation was expected to be capable of reducing or even obliterating genetically mediated amygdala hyperreactivity to negative emotional cues. Similar to previous investigations (Walter, et al., 2009), pictures of aversive or neutral emotional content were presented while participants were instructed to react naturally to half of the pictures, and to regulate their emotional response upon the other half of the stimuli. The first two studies of the present work were designed to further characterize the immediate aftereffect of volitional regulation in the amygdala: Study 1 included behavioral ratings of negativity at picture offset and at fixation offset in order to provide behavioral measures of experiential changes, while in Study 2, participants continued to experience or regulate their emotions during a “maintain” phase after picture offset. The primary goal of Study 3 was to evaluate whether volitional emotion regulation can reduce genetically mediated amygdala hyperreactivity to aversive emotional material in individuals with the short variant of the serotonin transporter genotype (Hariri, et al., 2005; Hariri, et al., 2002), and whether the immediate aftereffect is also influenced by the serotonin transporter genotype. In all three studies, the amygdala was significantly activated by aversive versus neutral stimuli, while cognitive emotion regulation attenuated the activation in the amygdala and increased the activation in a frontal-parietal network of regulatory brain regions. This neural effect was complemented by the behavioral ratings which show that the subjective experience of negativity was also reduced by detachment (Study 1). Also in all three studies, an immediate aftereffect was observed in the amygdala following the end of regulation. Moreover, the preoccupation with the previously seen pictures after the scanning session varied across the experimental conditions (Studies 2 and 3). Volitional regulation proved effective in reducing amygdala activation to negative stimuli even in 5-HTTLPR short allele carriers that show an increased reactivity to this type of cue. At the same time, functional coupling of the ventrolateral and medial orbital prefrontal cortex, the subgenual and the rostral anterior cingulate with the amygdala was higher in the s-group. However, in Study 3 the immediate aftereffect was found only in l/l-homozygote individuals following the regulation of fear. Taken together, the results of the three studies clearly show that volitional regulation is effective in reducing behavioral and neural correlates of the experience of negative emotions (Levesque, et al., 2003; Ochsner, Bunge, Gross, & Gabrieli, 2002; Ochsner, Ray, et al., 2004), even in the case of a genetically mediated hyperreactivity to such materials. Thus, it seems reasonable to assume that conscious will can effectively counteract genetic determinants of emotional behavior. Moreover, the present results suggest that the temporal dynamics of volitional emotion regulation are characterized by a paradoxical rebound in amygdala activation after regulation, and that the immediate aftereffect is a marker of the efficiency of the initial and the sustained effects of emotion regulation (Walter, et al., 2009). In summary, the successful replication of the immediate aftereffect of emotion regulation in all three studies of this dissertation opens up exciting new research perspectives: a comparison of the short- and long-term effects of different regulatory strategies, and the investigation of these effects also in positive emotions would complement the present results, since the neural mechanisms involved in these processes show some characteristic differences (Ochsner, 2007; Staudinger, Erk, Abler, & Walter, 2009). A comprehensive characterization of this neural marker and its implications for emotional experience might also be useful with respect to clinical applications. The detailed examination of the various time scales of emotional regulation might for instance inform the diagnostic and therapeutic interventions in affective disorders that are associated with emotional dysfunctions (Brewin, Andrews, & Rose, 2000; Johnstone, van Reekum, Urry, Kalin, & Davidson, 2007). Ultimately, we might thus come to understand the neural underpinnings of what the feelings we have today have to do with the feelings we had yesterday – and with the feelings with might have tomorrow

The Temporal Dynamics of Voluntary Emotion Regulation

Background: Neuroimaging has demonstrated that voluntary emotion regulation is effective in reducing amygdala activation to aversive stimuli during regulation. However, to date little is known about the sustainability of these neural effects once active emotion regulation has been terminated. Methodology/Principal Findings: We addressed this issue by means of functional magnetic resonance imaging (fMRI) in healthy female subjects. We performed an active emotion regulation task using aversive visual scenes (task 1) and a subsequent passive viewing task using the same stimuli (task 2). Here we demonstrate not only a significantly reduced amygdala activation during active regulation but also a sustained regulation effect on the amygdala in the subsequent passive viewing task. This effect was related to an immediate increase of amygdala signal in task 1 once active emotion regulation has been terminated: The larger this peak postregulation signal in the amygdala in task 1, the smaller the sustained regulation effect in task 2. Conclusions/Significance: In summary, we found clear evidence that effects of voluntary emotion regulation extend beyond the period of active regulation. These findings are of importance for the understanding of emotion regulation i

The temporal dynamics of volitional emotion regulation

Happiness, anger, surprise, irritation… if we note down the emotions that we go through on a given day, the list will most probably be quite long. A surge of studies on the bidirectional interaction between emotion and cognition suggests that we need emotional appraisals in order to lead a successful life and maintain our personal, social and economic integrity (Bechara, 2005; Damasio, 1994; Fox, 2008; Gross & Thompson, 2007; Walter, 2005). And yet, we seldom ‘just’ experience emotions, but often try to influence them to best fit our current goals. Based on the assumption that emotional reactions entail changes on various levels, and that these changes happen in- or outside of our awareness, affective science has adopted emotion regulation as one of its major research topics (Beauregard, Levesque, & Paquette, 2004; Gross, 1999; Ochsner, 2007). In fact, neural (e.g. amygdala activation) and behavioral (e.g. feeling of negativity) correlates of emotional reactions are effectively reduced by top-down processes of explicit and implicit control (Drabant, McRae, Manuck, Hariri, & Gross, 2009; Levesque, et al., 2003; Ochsner, Ray, et al., 2004). Furthermore, evidence from studies investigating voluntary thought control suggests that control strategies may have lasting and paradoxical consequences (Abramowitz, Tolin, & Street, 2001; Wegner, 2009). In a very recent investigation, lasting effects of regulation were also shown after the cognitive control of emotions: the activation timecourse of the amygdala was significantly increased immediately following regulation, and this difference was also related to the activation of the amygdala to the same stimuli a few minutes later (Walter, et al., 2009). Aside from these contextual or qualitative influences, emotional processing also differs between individuals: genetic variation within the serotonergic system for instance is known to affect emotional reactivity both on the behavioral and on the neural level (Hariri, et al., 2005; Hariri, et al., 2002; Lesch, et al., 1996). In the present work, the temporal dynamics of volitional emotion regulation were investigated in three studies. It was hypothesized that both the subjective experience of negativity and the amygdala activation can be attenuated by the detachment from negative emotions, which in turn leads to an immediate neural aftereffect after the offset of regulation. Furthermore, volitional emotion regulation was expected to be capable of reducing or even obliterating genetically mediated amygdala hyperreactivity to negative emotional cues. Similar to previous investigations (Walter, et al., 2009), pictures of aversive or neutral emotional content were presented while participants were instructed to react naturally to half of the pictures, and to regulate their emotional response upon the other half of the stimuli. The first two studies of the present work were designed to further characterize the immediate aftereffect of volitional regulation in the amygdala: Study 1 included behavioral ratings of negativity at picture offset and at fixation offset in order to provide behavioral measures of experiential changes, while in Study 2, participants continued to experience or regulate their emotions during a “maintain” phase after picture offset. The primary goal of Study 3 was to evaluate whether volitional emotion regulation can reduce genetically mediated amygdala hyperreactivity to aversive emotional material in individuals with the short variant of the serotonin transporter genotype (Hariri, et al., 2005; Hariri, et al., 2002), and whether the immediate aftereffect is also influenced by the serotonin transporter genotype. In all three studies, the amygdala was significantly activated by aversive versus neutral stimuli, while cognitive emotion regulation attenuated the activation in the amygdala and increased the activation in a frontal-parietal network of regulatory brain regions. This neural effect was complemented by the behavioral ratings which show that the subjective experience of negativity was also reduced by detachment (Study 1). Also in all three studies, an immediate aftereffect was observed in the amygdala following the end of regulation. Moreover, the preoccupation with the previously seen pictures after the scanning session varied across the experimental conditions (Studies 2 and 3). Volitional regulation proved effective in reducing amygdala activation to negative stimuli even in 5-HTTLPR short allele carriers that show an increased reactivity to this type of cue. At the same time, functional coupling of the ventrolateral and medial orbital prefrontal cortex, the subgenual and the rostral anterior cingulate with the amygdala was higher in the s-group. However, in Study 3 the immediate aftereffect was found only in l/l-homozygote individuals following the regulation of fear. Taken together, the results of the three studies clearly show that volitional regulation is effective in reducing behavioral and neural correlates of the experience of negative emotions (Levesque, et al., 2003; Ochsner, Bunge, Gross, & Gabrieli, 2002; Ochsner, Ray, et al., 2004), even in the case of a genetically mediated hyperreactivity to such materials. Thus, it seems reasonable to assume that conscious will can effectively counteract genetic determinants of emotional behavior. Moreover, the present results suggest that the temporal dynamics of volitional emotion regulation are characterized by a paradoxical rebound in amygdala activation after regulation, and that the immediate aftereffect is a marker of the efficiency of the initial and the sustained effects of emotion regulation (Walter, et al., 2009). In summary, the successful replication of the immediate aftereffect of emotion regulation in all three studies of this dissertation opens up exciting new research perspectives: a comparison of the short- and long-term effects of different regulatory strategies, and the investigation of these effects also in positive emotions would complement the present results, since the neural mechanisms involved in these processes show some characteristic differences (Ochsner, 2007; Staudinger, Erk, Abler, & Walter, 2009). A comprehensive characterization of this neural marker and its implications for emotional experience might also be useful with respect to clinical applications. The detailed examination of the various time scales of emotional regulation might for instance inform the diagnostic and therapeutic interventions in affective disorders that are associated with emotional dysfunctions (Brewin, Andrews, & Rose, 2000; Johnstone, van Reekum, Urry, Kalin, & Davidson, 2007). Ultimately, we might thus come to understand the neural underpinnings of what the feelings we have today have to do with the feelings we had yesterday – and with the feelings with might have tomorrow

The temporal dynamics of volitional emotion regulation

Happiness, anger, surprise, irritation… if we note down the emotions that we go through on a given day, the list will most probably be quite long. A surge of studies on the bidirectional interaction between emotion and cognition suggests that we need emotional appraisals in order to lead a successful life and maintain our personal, social and economic integrity (Bechara, 2005; Damasio, 1994; Fox, 2008; Gross & Thompson, 2007; Walter, 2005). And yet, we seldom ‘just’ experience emotions, but often try to influence them to best fit our current goals. Based on the assumption that emotional reactions entail changes on various levels, and that these changes happen in- or outside of our awareness, affective science has adopted emotion regulation as one of its major research topics (Beauregard, Levesque, & Paquette, 2004; Gross, 1999; Ochsner, 2007). In fact, neural (e.g. amygdala activation) and behavioral (e.g. feeling of negativity) correlates of emotional reactions are effectively reduced by top-down processes of explicit and implicit control (Drabant, McRae, Manuck, Hariri, & Gross, 2009; Levesque, et al., 2003; Ochsner, Ray, et al., 2004). Furthermore, evidence from studies investigating voluntary thought control suggests that control strategies may have lasting and paradoxical consequences (Abramowitz, Tolin, & Street, 2001; Wegner, 2009). In a very recent investigation, lasting effects of regulation were also shown after the cognitive control of emotions: the activation timecourse of the amygdala was significantly increased immediately following regulation, and this difference was also related to the activation of the amygdala to the same stimuli a few minutes later (Walter, et al., 2009). Aside from these contextual or qualitative influences, emotional processing also differs between individuals: genetic variation within the serotonergic system for instance is known to affect emotional reactivity both on the behavioral and on the neural level (Hariri, et al., 2005; Hariri, et al., 2002; Lesch, et al., 1996). In the present work, the temporal dynamics of volitional emotion regulation were investigated in three studies. It was hypothesized that both the subjective experience of negativity and the amygdala activation can be attenuated by the detachment from negative emotions, which in turn leads to an immediate neural aftereffect after the offset of regulation. Furthermore, volitional emotion regulation was expected to be capable of reducing or even obliterating genetically mediated amygdala hyperreactivity to negative emotional cues. Similar to previous investigations (Walter, et al., 2009), pictures of aversive or neutral emotional content were presented while participants were instructed to react naturally to half of the pictures, and to regulate their emotional response upon the other half of the stimuli. The first two studies of the present work were designed to further characterize the immediate aftereffect of volitional regulation in the amygdala: Study 1 included behavioral ratings of negativity at picture offset and at fixation offset in order to provide behavioral measures of experiential changes, while in Study 2, participants continued to experience or regulate their emotions during a “maintain” phase after picture offset. The primary goal of Study 3 was to evaluate whether volitional emotion regulation can reduce genetically mediated amygdala hyperreactivity to aversive emotional material in individuals with the short variant of the serotonin transporter genotype (Hariri, et al., 2005; Hariri, et al., 2002), and whether the immediate aftereffect is also influenced by the serotonin transporter genotype. In all three studies, the amygdala was significantly activated by aversive versus neutral stimuli, while cognitive emotion regulation attenuated the activation in the amygdala and increased the activation in a frontal-parietal network of regulatory brain regions. This neural effect was complemented by the behavioral ratings which show that the subjective experience of negativity was also reduced by detachment (Study 1). Also in all three studies, an immediate aftereffect was observed in the amygdala following the end of regulation. Moreover, the preoccupation with the previously seen pictures after the scanning session varied across the experimental conditions (Studies 2 and 3). Volitional regulation proved effective in reducing amygdala activation to negative stimuli even in 5-HTTLPR short allele carriers that show an increased reactivity to this type of cue. At the same time, functional coupling of the ventrolateral and medial orbital prefrontal cortex, the subgenual and the rostral anterior cingulate with the amygdala was higher in the s-group. However, in Study 3 the immediate aftereffect was found only in l/l-homozygote individuals following the regulation of fear. Taken together, the results of the three studies clearly show that volitional regulation is effective in reducing behavioral and neural correlates of the experience of negative emotions (Levesque, et al., 2003; Ochsner, Bunge, Gross, & Gabrieli, 2002; Ochsner, Ray, et al., 2004), even in the case of a genetically mediated hyperreactivity to such materials. Thus, it seems reasonable to assume that conscious will can effectively counteract genetic determinants of emotional behavior. Moreover, the present results suggest that the temporal dynamics of volitional emotion regulation are characterized by a paradoxical rebound in amygdala activation after regulation, and that the immediate aftereffect is a marker of the efficiency of the initial and the sustained effects of emotion regulation (Walter, et al., 2009). In summary, the successful replication of the immediate aftereffect of emotion regulation in all three studies of this dissertation opens up exciting new research perspectives: a comparison of the short- and long-term effects of different regulatory strategies, and the investigation of these effects also in positive emotions would complement the present results, since the neural mechanisms involved in these processes show some characteristic differences (Ochsner, 2007; Staudinger, Erk, Abler, & Walter, 2009). A comprehensive characterization of this neural marker and its implications for emotional experience might also be useful with respect to clinical applications. The detailed examination of the various time scales of emotional regulation might for instance inform the diagnostic and therapeutic interventions in affective disorders that are associated with emotional dysfunctions (Brewin, Andrews, & Rose, 2000; Johnstone, van Reekum, Urry, Kalin, & Davidson, 2007). Ultimately, we might thus come to understand the neural underpinnings of what the feelings we have today have to do with the feelings we had yesterday – and with the feelings with might have tomorrow

Oxytocin modulates neural reactivity to children's faces as a function of social salience

Oxytocin (OT) enhances social behaviors such as attachment and parental caretaking. Neural correlates of maternal attachment are found in reward-related brain regions, for example, in the globus pallidus (GP). The present work investigates the effects of OT on the neural correlates of parental attachment. Fathers viewed pictures of their own child (oC), a familiar child (fC), and an unfamiliar child (uC) after intranasal application of OT vs placebo. OT reduced activation and functional connectivity of the left GP with reward- and attachment-related regions responsive to pictures of the oC and the uC. The present results emphasize the key role of OT in human parental attachment and suggest that OT reduces neural reactivity to social cues as a function of social salience. Our results together with previous findings speak to a selective reduction of neural reactivity to social stimuli, irrespective of their valence. We argue that one major pathway by which OT exerts its positive effects on affiliative and social behaviors is the attenuation of automatic neural responses, which in turn leads to increased approach behaviors and decreased social avoidance

Attachment representation modulates oxytocin effects on the processing of own-child faces in fathers

Oxytocin (OT) plays a crucial role in parental-infant bonding and attachment. Recent functional imaging studies reveal specific attachment and reward related brain regions in individuals or within the parent-child dyad. However, the time course and functional stage of modulatory effects of OT on attachment-related processing, especially in fathers, are poorly understood. To elucidate the functional and neural mechanisms underlying the role of OT in paternal-child attachment, we performed an event-related potential study in 24 healthy fathers who received intranasal OT in a double-blind, placebo-controlled, within-subject experimental design. Participants passively viewed pictures of their own child (oC), a familiar (fC) and an unfamiliar child (ufC) while event-related potentials were recorded. Familiarity of the child’s face modulated a broad negativity at occipital and temporo-parietal electrodes within a time window of 300–400 ms, presumably reflecting a modulation of the N250 and N300 ERP components. The oC condition elicited a more negative potential compared to the other familiarity conditions suggesting different activation of perceptual memory representations and assignment of emotional valence. Most importantly, this familiarity effect was only observed under placebo (PL) and was abolished under OT, in particular at left temporo-parietal electrodes. This OT induced attenuation of ERP responses was related to habitual attachment representations in fathers. In summary, our results demonstrate an OT-specific effect at later stages of attachment-related face processing presumably reflecting both activation of perceptual memory representations and assignment of emotional value

Task 1 (active regulation).

<p>Upper row left: Amygdala activation was significantly attenuated during regulation (p<0.05 FWE corrected for ROI). This regulation related decrease of amygdala activation was positively correlated with a regulation related increase in DLPFC activation (Upper row, right). Bottom row, left: Time course of left Amygdala, showing a significant postregulation rebound and a significant interaction of regulation and period (bar plot bottom row, middle). Note: all effects here shown for the left amygdala, are also significant for the right amygdala. Bottom row, right: Positive correlation between peak activation during relax period (rebound) in left amygdala and individual scores in the WBSI.</p

Delay discounting without decision-making: Medial prefrontal cortex and amygdala activations reflect immediacy processing and correlate with impulsivity and anxious-depressive traits

Humans value rewards less when these are delivered in the future as opposed to immediately, a phenomenon referred to as delay discounting. While delay discounting has been studied during the anticipation of rewards and in the context of intertemporal decision-making, little is known about its neural correlates in the outcome phase (during reward delivery) and their relation to personality. Personality traits that have been associated with increased delay discounting include impulsivity and, potentially, anxious-depressive traits. Here we performed functional magnetic resonance imaging (fMRI) in 72 healthy participants while they carried out a monetary incentive delay task with a delay manipulation. In sixty percent of the experimental trials, participants won rewards that differed in magnitude (0.05€, 0.50€ or 1€) and delay until delivery (immediately, 10 days, or 100 days). A factor analysis on questionnaires yielded two factors reflecting Impulsivity and Anxiety/Depression, which we used to examine potential relationships between personality and delay discounting. When winning a reward, medial prefrontal cortex (mPFC) activation was higher for immediate compared to delayed rewards. Moreover, amygdala activation correlated with reward magnitude for immediate but not for delayed rewards. Amygdala activation to winning immediate rewards was higher in more impulsive participants, while mPFC activation to winning immediate rewards was higher in more anxious-depressed participants. Our results uncover neural correlates of delay discounting during reward delivery, and suggest that impulsivity and subclinical anxious-depressive traits are related to stronger neural responses for winning immediate relative to delayed rewards

Results of Task 1 (Active Regulation).

<p>All results: p<0.05 FWE corrected for multiple comparisons; ^*p<0.05 FWE corrected for region of interest; BA Brodmann area; x,y,z, respective coordinates of MNI template.</p