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Emotion Regulation

BP Doré and KN Ochsner, Columbia University, New York, NY, USA


ã 2015 Elsevier Inc. All rights reserved.

Glossary Emotional response A psychological state that involves


Amygdala A subcortical emotion generation region that subjective experience (e.g., feelings of valence, arousal, and/
registers arousal and stimulus salience. or particular categories of emotion), behavior (e.g., facial,
Domain-general control regions Prefrontal, parietal, and bodily, and verbal), and peripheral physiology (e.g., heart
cingulate regions of the brain that support controlled rate and respiration).
processing in general, including the implementation of Inferior parietal cortex (iPC) A parietal control region that
emotion regulation strategies like reappraisal. supports controlled shifts of attention.
Dorsal anterior cingulate cortex (dACC) A prefrontal Insula A cortical emotion generation region that supports
control region that detects conflict and signals adjustments integration of affective and visceral–somatic information.
in controlled processing. Lateral temporal cortex A brain region that supports
Dorsolateral prefrontal cortex (dlPFC) A prefrontal reappraisal and is known to reflect semantic and perceptual
control region that supports maintenance and manipulation processing.
of information in working memory. Reappraisal A particularly powerful and well-studied
Dorsomedial prefrontal cortex (dmPFC) A brain region emotion regulation strategy that involves changing the way
that supports reappraisal and is known to underlie mental- one thinks about the meaning of a stimulus.
state judgments. Ventral striatum A subcortical emotion generation region
Emotion generation The process by which a situation is that supports encoding and construction of stimulus value.
selected, attended to, and appraised as goal-relevant, giving Ventrolateral prefrontal cortex (vlPFC) A prefrontal
rise to an emotional response. control region that supports the inhibition and controlled
Emotion generation regions Cortical and subcortical selection of semantic information.
regions of the brain that support emotion generation and are Ventromedial prefrontal cortex (vmPFC) A brain region
modulated by regulation strategies like reappraisal. that may underlie individual or group differences in
Emotion regulation The recruitment of strategies that reappraisal ability and is known to underlie fear extinction,
influence one or more stages of the emotion generation value computation, and self-processing.
process and thereby modify emotional responses.

Many of history’s greatest minds hold that life is tragic. Some of Lazarus, 1991). Given that we are at a relatively early stage of
life’s tragedies are major, like losing a loved one. And many of brain-imaging research on emotion and its regulation, we take
them are small, like arriving at work and realizing that you an inclusive stance and group both terms under the umbrella
forgot your keys at home. These events elicit emotional category of emotional responses that can be regulated.
responses ranging from grief to sadness, anger, and frustration, As shown in Figure 1, contemporary views hold that emo-
among many others. How, exactly, does the brain enable us to tion generation involves at least four distinct stages (adapted
regulate these emotions? The answer to this question has from Gross, 1998; Ochsner, Silvers, & Buhle, 2012). An emo-
implications for our mental and physical well-being (DeSteno tion is generated when (1) a situation is encountered (i.e.,
et al., 2013; Gross & Munoz, 1995) and our scientific under- stimuli embedded within a context are perceived), (2) one or
standing of brain function. In recent years, an entire field of more features of this situation are attended to, (3) these fea-
research has emerged devoted to understanding this ability. tures are appraised as positive or negative in a variety of ways
Here, we summarize this research and outline a brain-based depending on their relevance to current and chronic goals, and
model of emotion regulation that focuses on the neural sys- (4) a corresponding pattern of experience, behavior, and/or
tems underlying the cognitive control of emotion. peripheral physiology is produced. Although in prototypical
emotional responses this pattern is described as coordinated
(e.g., feelings of fear, escape behavior, and increased heart rate
The Process Model of Emotion arise together upon seeing a snake), research on this topic
suggests that coherence of these variables may be less than
Emotion Generation
originally hypothesized (Mauss, Levenson, McCarter, Wilhelm,
Before turning to how emotions can be regulated, it is useful to & Gross, 2005).
consider what emotions are and how they arise. Some theorists
distinguish between affective responses, which are valenced (i.e.,
Emotion Regulation
positive or negative) evaluations that reflect an assessment of
the momentary goodness or badness of a stimulus, and emo- Emotion regulation involves the modification of emotional
tions, which entail a richer appraisal of the meaning of the responses via the recruitment of strategies that influence par-
stimulus along multiple dimensions (see, e.g., Barrett, 2012; ticular stages of the emotion generation process. Figure 1

Brain Mapping: An Encyclopedic Reference http://dx.doi.org/10.1016/B978-0-12-397025-1.00153-6 53


54 INTRODUCTION TO SOCIAL COGNITIVE NEUROSCIENCE | Emotion Regulation

Emotion regulation
Situation selection Attention Cognitive Response
and modification deployment change modulation

Situation Attention Appraisal Response


Emotion generation
Figure 1 The process model of emotion. See text for details.

Domain-general control
Emotion generation
and other relevant regions

Figure 2 Brain mechanisms of emotion regulation. See text for details.

illustrates four such families of emotion regulation strategies, Diekhof, Geier, Falkai, & Gruber, 2011; Kalisch, 2009) in
organized in terms of the stage of the emotion generation healthy adults suggest that implementing reappraisals involves
process that they impact. Note that although the goal to regu- recruitment of domain-general control regions, including the pre-
late one’s emotions and the strategies used to do so could frontal, parietal, and cingulate cortices, in order to influence
theoretically be activated and operate implicitly and automat- activity in the subcortical and cortical emotion generation
ically, almost all neuroscience studies of emotion regulation regions, including the amygdala, ventral striatum, and insula
have focused on strategies that are explicitly motivated, cued, (Figure 2).
and implemented (Gyurak, Gross, & Etkin, 2011).
The most forward-thinking families of strategies, situation
Emotion Generation Regions
selection and modification, entail efforts to influence the kinds of
situations that one will experience or to modify relevant fea- First, we consider brain regions that support the valenced eval-
tures of those situations once you are in them. Next, attention uation of stimuli and thereby the generation of emotional
deployment involves directing attention toward or away from responses. Activity in these regions is associated with the expe-
features of a given situation, as when distracting oneself. After rience of emotional states (Kober et al., 2008) and is modulated
that, cognitive change strategies target our appraisals, changing by regulation strategies like reappraisal (Buhle et al., 2013;
the ways we think about an attended stimulus in order to Diekhof et al., 2011; Kalisch, 2009; Ochsner et al., 2012).
change its emotional impact. The prototypical cognitive The amygdala. The amygdala is a subcortical region that
change strategy, reappraisal, involves deliberately changing detects affectively salient stimuli, modulates activity in percep-
one’s interpretation of and/or personal connection to a stimu- tual and memory systems for further processing of these stim-
lus. Finally, response modulation strategies target and modulate uli, and can trigger appropriate behaviors (Cunningham, Van
the behavioral component of the emotional response, for Bavel, & Johnsen, 2008; Whalen, 1998). Although early theo-
example, emotional facial expressions. ries of amygdala function emphasized its role in negative affect
(e.g., LeDoux, 1998), it has become clear that this region also
responds to positive, novel, and ambiguous stimuli (e.g.,
Brain Mechanisms of Emotion Regulation Phelps, 2006). Amygdala activity is modulated by reappraisal
in studies that use negative stimuli as well as studies using
Although behavioral studies have examined how the strategies positive stimuli, although studies of the latter are far less repre-
listed in the preceding text differ in terms of their impact on the sented in the current literature (Ochsner et al., 2012). In the
emotional response (e.g., Gross & John, 2003; Kross & Ayduk, context of emotion regulation, amygdala activity is thought to
2008; Richards & Gross, 2000), to date, the majority of brain- reflect the current salience and arousal level elicited by the
imaging research has focused on the prototypical cognitive viewed or regulated stimulus.
change strategy, reappraisal. Recent meta-analyses of over 40 The ventral striatum. The ventral portion of the striatum is
brain-imaging studies of reappraisal (Buhle et al., 2013; involved in encoding and constructing representations of
INTRODUCTION TO SOCIAL COGNITIVE NEUROSCIENCE | Emotion Regulation 55

stimulus value that guide learning and motivate behavior. As a language production (Snyder et al., 2010) and, more generally,
key region of the mesolimbic dopamine reward pathway, the the selection and controlled retrieval of information from
ventral striatum receives dopaminergic input from the midbrain semantic memory (Badre & Wagner, 2007). This region is
ventral tegmental area (Haber & Knutson, 2009; Schultz, Dayan, also known to support controlled inhibition of prepotent
& Montague, 1997). Functional imaging studies show that the behavioral responses (Aron, Robbins, & Poldrack, 2004; Levy
ventral striatum responds to expectation and receipt of a wide & Wagner, 2011). Reappraisal is theorized to depend deeply on
variety of rewards, including sweet liquids, money, music, memory and language processes to support the generation of
attractive faces, and social fairness (e.g., Cloutier, Heatherton, alternative narratives about the meaning of affective stimuli,
Whalen, & Kelley, 2008; King-Casas et al., 2005; O’Doherty, which also involves the inhibition of representations that gen-
2004; Salimpoor, Benovoy, Larcher, Dagher, & Zatorre, 2011). erate one’s initial affective response. As such, it is likely that
Although emotion regulation studies have reported modulation vlPFC activation in studies of reappraisal reflects computations
of the ventral striatum much less commonly than modulation supporting some or all of these functions.
of the amygdala (Buhle et al., 2013) – in large part because they The dorsal anterior cingulate cortex. Implementing control dur-
have infrequently utilized appetitive/positive stimuli – studies ing reappraisal may require detection of conflict between com-
employing mediation analysis support the idea that the reap- peting responses. Models of dACC function emphasize its role
praisal of aversive stimuli can rely on independent pathways of as a nexus region that detects and signals the need for adjust-
amygdala and ventral striatum modulation (Kober et al., 2010; ments in cognitive control (Botvinick, Braver, Barch, Carter, &
Wager, Davidson, Hughes, Lindquist, & Ochsner, 2008). Cohen, 2001; Miller & Cohen, 2001; Ridderinkhof, Ullsperger,
The insula. The insula is a cortical region hidden beneath Crone, & Nieuwenhuis, 2004). In the context of emotion regu-
overlying folds of the temporal and parietal cortices. Based on lation, dACC may serve to signal the conflict between one’s
patterns of anatomical connectivity (Augustine, 1996; Craig, initial appraisal of a stimulus and a desired reappraisal of it or
2009) and meta-analyses of neuroimaging studies (e.g., Chang, conflicts between current and desired levels of affect.
Yarkoni, Khaw, & Sanfey, 2013), it has been proposed
that the insula supports the integration of affective and
Other Regions Relevant to Emotion Regulation
visceral–somatic information involved in interoceptive states,
like emotion (Garfinkel et al., 2013; Zaki, Davis, & Ochsner, Two regions that do not map cleanly onto cognitive control
2012), including the experience of disgust and pain. In studies systems – the dorsomedial prefrontal cortex (dmPFC) and the
of reappraisal, insula modulation has been observed far less lateral temporal cortex – also are consistently observed to
commonly than modulation of either the amygdala or ventral increase their activity during reappraisal. A third region, the
striatum (Ochsner et al., 2012). ventromedial prefrontal cortex (vmPFC), has not been consis-
tently observed during reappraisal (Buhle et al., 2013), but has
been variably proposed to support affect generation or con-
Domain-General Control Regions
trolled regulation or to act as a mediator of the effects of
Next, we turn to a set of brain regions previously identified as control regions on emotion generation regions (see Diekhof
critical in the ability to exert cognitive control over the contents et al., 2011; Roy, Shohamy, & Wager, 2012).
of memory, focus of attention, and language functions. Meta- The dorsomedial prefrontal cortex. The dmPFC is a cortical
analyses support the idea that reappraisal is implemented by region known to support mental-state judgments about tran-
these domain-general control regions (Buhle et al., 2013; sient or enduring qualities of oneself or other people (Denny,
Diekhof et al., 2011; Kalisch, 2009), including the dorsal ante- Kober, Wager, & Ochsner, 2012; Mitchell, 2009). Activation in
rior cingulate cortex (dACC), dorsolateral prefrontal cortex this region is also reliably observed during reappraisal, alongside
(dlPFC), inferior parietal cortex (iPC), ventrolateral prefrontal domain-general control regions, suggesting that processes
cortex (vlPFC), and dorsomedial prefrontal cortex (dmPFC). recruited to support introspection about and assessments of
The dorsolateral prefrontal cortex and inferior parietal cortex. mental states also support emotion regulation. This observed
When reappraising an affectively salient stimulus, a set of cog- activity may reflect attempts to judge one’s own emotional state
nitive control regions are engaged. Chief among these are the during the reappraisal period or to gauge and reconsider the
dlPFC and iPC, which together form the brain’s frontoparietal mental states of individuals depicted in the evocative stimuli.
network thought to underlie goal-directed (i.e., top-down) con- The lateral temporal cortex. Reappraisal also consistently
trol of attention and working memory (Corbetta & Shulman, engages a swath of the lateral temporal cortex, an area of the
2002; Wager, Jonides, & Reading, 2004). Of the two compo- brain thought to support abstract representation of semantic
nents of this network, studies suggest that the dlPFC is involved and perceptual information (Binder & Desai, 2011; Visser &
primarily in the maintenance and manipulation of information Ralph, 2011). This pattern of findings suggests that the cogni-
in working memory (Ptak, 2012; Wager & Smith, 2003) whereas tive control of emotion involves using control systems to mod-
the iPC is more involved in triggering shifts of attention between ulate semantic and perceptual processing of a stimulus, which
external stimuli and internal representations (Ptak, 2012; Wager in turn leads to modulation of emotion generation regions like
& Smith, 2003). In the context of emotion regulation, the dlPFC the amygdala (Buhle et al., 2013).
and iPC may work together to attend to, maintain, and manip- The ventromedial prefrontal cortex. The vmPFC is known to
ulate semantic information related to stimulus qualities, directly track reward outcome magnitude in a variety of contexts
appraisal representations, and regulatory goals. and is thought to compute an integrative value signal that takes
The ventrolateral prefrontal cortex. Another region subserving into account stimulus history and context (Fehr & Rangel, 2011;
mechanisms of controlled processing, the vlPFC is involved in Schoenbaum, Saddoris, & Stalnaker, 2007). Interestingly, this
56 INTRODUCTION TO SOCIAL COGNITIVE NEUROSCIENCE | Emotion Regulation

region is crucial to reversal learning and extinction of condi- studies of reappraisal, may generalize to diverse social and
tioned fear, as well as placebo analgesia (Delgado, Gillis, & emotional contexts, but the particular regions recruited (and
Phelps, 2008; Diekhof et al., 2011; Schiller & Delgado, 2010). the interplay between these regions) are likely to vary across
Although several theorists have proposed that the vmPFC plays emotion categories, stimulus types, regulatory goals, and spe-
a role in emotion regulation broadly construed, this region is cific regulatory tactics.
not consistently activated in studies of reappraisal (Buhle et al., Although research on this topic has expanded dramatically in
2013). However, this region is known to relate to individual recent years, several areas merit even closer attention. An impor-
and group differences in reappraisal efficacy (Erk et al., 2010; tant goal for translational research will be to understand how
Johnstone, van Reekum, Urry, Kalin, & Davidson, 2007; Wager dysfunction in either bottom-up emotion generation or top-
et al., 2008) and may play a general role in implicit forms of down control might underlie patterns of emotional changes
emotion regulation (Gyurak et al., 2011). observed in psychopathology. Such a direction is already being
pursued in studies of emotion regulation across various disorders,
including borderline personality disorder (e.g., Koenigsberg et al.,
Extending the Model to Other Regulatory Phenomena 2009), major depression (e.g., Heller et al., 2009), bipolar disor-
der (e.g., Townsend et al., 2012), and anxiety disorders (e.g., Etkin
Having established a model of the basic brain mechanisms
& Wager, 2007). Similarly, there will be a growing value in
underlying emotion regulation, research has begun to ask
research that investigates brain changes underlying life-span
how these brain systems may contribute to other forms of
changes in emotion experience from childhood and adolescence
regulatory and social cognitive phenomena.
(e.g., McRae, Ciesielski, & Gross, 2012; Silvers et al., 2012) to
Affect labeling. Even in the absence of a goal to regulate,
young adulthood and old age (e.g., Mather, 2012).
simply verbally labeling feelings can have a regulatory effect in
In addition to these translational directions, several basic
some circumstances. Recent research demonstrates that engag-
questions about the regulation of emotion remain understu-
ing in affect labeling when viewing an affective stimulus leads to
died. First, among the constellation of distinct regions that
increases in vlPFC activity, decreases in amygdala activity
support reappraisal, the mechanistic contributions of particu-
(Torrisi, Lieberman, Bookheimer, & Altshuler, 2013), and
lar regions remain relatively unclear; this could be addressed by
attenuated intensity of self-reported emotional experience
applying conjunction analyses to reappraisal and cognitive
(Lieberman, Inagaki, Tabibnia, & Crockett, 2011), suggesting
control tasks and by developing more controlled paradigms
that verbally labeling affective responses leads to the incidental
that fractionate reappraisal into distinct component processes
regulation of affective responses to them.
(see Cohen, Berkman, & Lieberman, 2012). Second, it will be
Self-serving biases. Self-enhancement, or the motivated ten-
important for future studies to determine the impact of stimu-
dency to judge oneself in an unrealistically positive light, repre-
lus valence and regulatory goals on activity in control regions
sents another pervasive phenomenon that relates to affect
and emotion generation regions, as well as intermediary
regulation (Taylor & Brown, 1994). Interestingly, the most con-
regions that may mediate these effects. Third, it will be impor-
sistent neural marker of unrealistically positive social cognition
tant to use neuroscience to address fundamentally new ques-
seems to be a reduction in vmPFC activity (Hughes & Beer,
tions about how emotion regulation operates. For example,
2012; Somerville, Kelley, & Heatherton, 2010), which has
although theories of emotion regulation emphasize the impor-
been interpreted as reflecting reductions in integration processes
tance of recognizing appropriate contexts to regulate and acti-
that can be used to overcome self-serving biases (Beer, 2012).
vating and sustaining a goal to implement regulation, little is
Self-regulation. Given that many domains of regulation rely
known about the mechanisms underlying these abilities. For
on similar regions of the brain, recent studies that have
another, while it is clear that regulation strategies have large
attempted to use brain imaging to examine the neural basis
immediate effects on emotional responses, few studies have
of the prominent limited resource model of self-regulation hold
attempted to determine the long-term effects of practicing
that repeated attempts at regulation draw on and eventually
regulation on responses to previously regulated or novel stim-
exhaust a common regulatory resource (Baumeister & Heather-
uli. Lastly, although virtually all of the research on this topic
ton, 1996; Heatherton, 2011). Recent imaging work supports
looks at explicit regulation, it will be important to understand
this idea by showing that engaging in an effortful self-
the neural mechanisms underlying more implicit or automatic
regulation task leads to exaggerated amygdala responses to
forms of regulation, which may be less prone to failure under
negative images and reduced functional connectivity between
certain circumstances. In addressing these and other questions,
the amygdala and the vmPFC (Wagner & Heatherton, 2013).
it is our hope that this field will continue to inspire theoretical
and empirical work that builds a nuanced and comprehensive
understanding of the contribution of specific brain mecha-
Conclusion and Future Directions nisms to emotion regulation successes and failures.

In this article, we have outlined research demonstrating that


the brain mechanisms of emotion regulation involve interac-
tions between emotion generation regions that support emo- See also: INTRODUCTION TO SOCIAL COGNITIVE
tional responding and domain-general control regions that act NEUROSCIENCE: Emotional Experience; Self-Regulation and Self-
to modulate activity in these generation regions. Available Regulation Failure; INTRODUCTION TO SYSTEMS: Brain Mapping
research suggests that this model, derived primarily from of Control Processes; Emotion; Reward; Salience Network.
INTRODUCTION TO SOCIAL COGNITIVE NEUROSCIENCE | Emotion Regulation 57

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