Guendelman, MEdeiros &rampes PDF

REVIEW
published: 06 March 2017

doi: 10.3389/fpsyg.2017.00220
Mindfulness and Emotion Regulation:

Insights from Neurobiological,
Psychological, and Clinical Studies
Simón Guendelman 1*, Sebastián Medeiros 2, 3 and Hagen Rampes 4
1
Social Cognition Group, Berlin School of Mind and Brain, Humboldt Universität, Berlin, Germany, 2 Research Unit on
Psychotherapeutic Interventions and Change Processes, Millennium Institute for Research in Depression and Personality,
Santiago, Chile, 3 Health Psychology, Department of Psychology, Pontificia Universidad Católica de Chile, Santiago, Chile,
4
Community Mental Health Team East, Central North West London Foundation NHS Foundation Trust, London, UK
There is increasing interest in the beneficial clinical effects of mindfulness-based

interventions (MBIs). Research has demonstrated their efficacy in a wide range of
psychological conditions characterized by emotion dysregulation. Neuroimaging studies
have evidenced functional and structural changes in a myriad of brain regions mainly
involved in attention systems, emotion regulation, and self-referential processing. In
this article we review studies on psychological and neurobiological correlates across
different empirically derived models of research, including dispositional mindfulness,
mindfulness induction, MBIs, and expert meditators in relation to emotion regulation.
From the perspective of recent findings in the neuroscience of emotion regulation,
we discuss the interplay of top-down and bottom-up emotion regulation mechanisms
Edited by: associated with different mindfulness models. From a phenomenological and cognitive
Alessandro Grecucci, perspective, authors have argued that mindfulness elicits a “mindful emotion regulation”
University of Trento, Italy
strategy; however, from a clinical perspective, this construct has not been properly
Reviewed by:
Eric S. Allard,
differentiated from other strategies and interventions within MBIs. In this context
Cleveland State University, USA we propose the distinction between top-down and bottom-up mindfulness based
Stéphane Dandeneau,
emotion regulation strategies. Furthermore, we propose an embodied emotion regulation
Université du Québec à Montréal,
Canada framework as a multilevel approach for understanding psychobiological changes due
*Correspondence: to mindfulness meditation regarding its effect on emotion regulation. Finally, based on
Simon Guendelman clinical neuroscientific evidence on mindfulness, we open perspectives and dialogues
[email protected]
regarding commonalities and differences between MBIs and other psychotherapeutic
Specialty section: strategies for emotion regulation.
This article was submitted to
Keywords: mindfulness, emotion regulation, neuroimaging, top down and bottom up processing, emotion
Emotion Science,
dysregulation disorders, embodied cognition
a section of the journal
Frontiers in Psychology
Received: 01 May 2016 Increasing interest has emerged about the therapeutic effects of mindfulness meditation and its
Accepted: 06 February 2017
clinical applications. Several studies have shown positive results in fostering emotional mental
Published: 06 March 2017
health among clinical and healthy populations (Bohlmeijer et al., 2010; Fjorback et al., 2011; Gotink
Citation: et al., 2015). Neurobiological studies indicate that this type of mental training may have an effect
Guendelman S, Medeiros S and
on the plasticity of brain structure and functioning (Tomasino et al., 2013; Fox et al., 2014). Some
Rampes H (2017) Mindfulness and
Emotion Regulation: Insights from
of the main neurocognitive mechanisms implicated in mindfulness meditation include attention
Neurobiological, Psychological, and control, emotion regulation, and self-awareness (Tang et al., 2015). In this article, we will focus
Clinical Studies. Front. Psychol. 8:220. on the relationship between mindfulness and emotion regulation, taking into account diverse
doi: 10.3389/fpsyg.2017.00220 psychological, clinical and neuroimaging evidence.
Frontiers in Psychology | www.frontiersin.org 1 March 2017 | Volume 8 | Article 220

Guendelman et al. Mindfulness and Emotion Regulation
Unlike other reviews on the topic, this article does not focus Taking into account how individual differences in emotion
on the problematic aspects involved in the operationalization regulation strategies influence mental health, and the extensive
and definition of mindfulness itself. Instead, the intention is role of emotion dysregulation in many psychopathological
to offer a comprehensive perspective linking different empirical conditions, it is reasonable to believe that clinical interventions
models including mindfulness as a trait, mindfulness inductions, focused on emotion regulation/dysregulation might have
MBIs and mindfulness experts, and emotion regulation-related substantial benefits for these psychological disorders. This
mechanisms including psychological and top-down/bottom-up argument is in line with several studies in which MBIs seem to
brain systems. Moreover, we propose a preliminary framework be particularly effective in clinical and non-clinical conditions
for better understanding of emotion regulation changes due to characterized by distress and negative emotions.
mindfulness practice, tackling problematic aspects of the notion Mindfulness meditation has its origin in the Buddhist
of “mindful emotion regulation” widely used in mindfulness psychology tradition, more specifically in the texts known as
clinical research, and complex involvement of top-down and Satipatthana Sutra (Analayo, 2003) and the Abhidharma (from
bottom-up mechanisms in MBIs. Sanskrit, means higher teachings), a cycle of teachings concern
about how the mind, including emotions and consciousness work
(Trungpa, 2001; Analayo, 2003; Rapgay and Bystrisky, 2009). The
MINDFULNESS, EMOTION REGULATION, word “mindfulness” corresponds to the translation of the original
AND CLINICAL APPLICATIONS terms smrti (from Sanskrit) or sati (Pali), which captures the
capacity to retain an object in the mind, but in a broad sense also
Contemporary psychology considers emotion regulation a implies being aware of and attentive to the present moment (Lutz
central component of mental health, and its imbalances might et al., 2015). In clinical and research contexts, mindfulness as a
underlie several mental disorders (Berenbaum et al., 2003; specific type of meditation practice has been described as a “non-
Mennin and Farach, 2007). Emotion regulation includes all of elaborative, non-judgmental awareness” of present-moment
the conscious and non-conscious strategies we use to increase, to experience (Kabat-Zinn, 2005), a non-reactive awareness that
maintain or decrease one or more components of an emotional emerges as a result of intentionally paying attention to
response (Gross, 1998). Originally, trying to bring together present experience, and a capacity that can be trained through
ideas from psychoanalysis and the field of stress and coping formal meditation practice. Several MBIs have been developed,
behaviors, Gross developed a process or time model of emotion including mindfulness meditation and other components, such
regulation, in which emotions can be modulated in five different as body awareness, yoga, and psychoeducation. These are
stages: selecting a situation, modifying a situation, deployment group interventions, specially designed for targeting specific
of attention, changing cognition (cognitive reappraisal), and psychopathological substrates (like emotion dysregulation), in
modulating the experience, behavior or physiological response particular those related to psychiatric conditions (Shonin et al.,
(Gross, 2001). Gross and John in a correlational study 2013).
demonstrated that individual differences in the usage of these The mindfulness-based stress reduction (MBSR) program
strategies (more cognitive reappraisal) were related to better was developed by Jon Kabat-Zinn during the late seventies
emotional health, well-being and interpersonal functioning (Kabat-Zinn, 2005). Several revisions and meta-analyses have
(Gross and John, 2003). highlighted its robust benefits for healthy subjects, increasing
In line with this approach, Aldao et al. performed a well-being, and decreasing stress and negative emotions (Eberth
meta-analytic review focused on how emotion regulation and Sedlmeier, 2012). For clinical population, highlights
strategies, measured by self-report scales, vary across different the decrease in pain intensity, stress, and psychological
psychopathological conditions. The main findings showed complaints among patients suffering from diverse chronic
that avoidance, rumination, and suppression (as strategies) pain/inflammatory diseases (Cramer et al., 2012; Lauche et al.,
were each positively associated with anxiety, depression and 2013) and cancer (Ledesma and Kumano, 2009). Recently, a
eating disorders. Problem-solving was negatively associated standardized review of meta-analysis of randomized controlled
with anxiety, depression and eating disorders. Reappraisal trials (RCTs) for MBSR and mindfulness-based cognitive therapy
and acceptance-based strategies were negatively associated, but (MBCT) demonstrated a significant improvement in different
not significantly, with anxiety and depression (Aldao et al., domains (calculated as Cohen’s d effect sizes): depressive
2010). Emotion dysregulation has been recognized as a core symptoms (d = 0.37), anxiety (d = 0.49), stress (d = 0.51), quality
psychopathological factor in many other psychological disorders of life (d = 0.39), physical functioning (d = 0.27; Gotink et al.,
such as borderline personality disorder (BPD; Linehan, 1993; 2015).
Schore, 2003), emotional trauma (Corrigan et al., 2011), attention MBCT is a program derived from MBSR, developed for
deficit hyperactivity disorder (ADHD; Shaw et al., 2014), bipolar preventing recurrence/relapse in recurrent major depressive
disorder (Van Rheenen et al., 2015), and anorexia and bulimia disorder (MDD; Segal et al., 2002). Several RCT and systematic
nervosa (Lavender et al., 2015). Emotion dysregulation has been reviews have demonstrated its effectiveness in relapse prevention
demonstrated to mediate the link between child abuse/neglect and residual symptoms (Chiesa and Serretti, 2011; Piet and
and later depressive disorder (Crow et al., 2014), and also the Hougaard, 2011; Clarke et al., 2015), and lately, also, in
link between cumulative adversity in lifetime and depressive depressive symptoms in MDD (Jain et al., 2015). Another
symptoms (Abravanel and Sinha, 2015). MBI is mindfulness-based relapse prevention (MBRP), which

TABLE 1 | Summary of mindfulness-based interventions (MBIs) and main lead to decreased mental proliferation (rumination narrative
evidence-based targeted conditions. based), and through this to salutary effects (Grabovac et al., 2011).
MBI Main conditions with evidence support for MBI
Other authors have proposed neurocognitive models,
integrating psychological and neuroscientific data. Vago and
MBSR Stress, burnout (health professions) Silbersweig proposed that mindfulness leads to changes in
Chronic pain (low-back pain, fibromyalgia) self-processing, through the development of self-awareness
Cancer (meta-awareness), self-regulation (modulation of behavior),
MBCT MDD (relapse prevention and acute treatment). BD and self-transcendence (prosocial characteristics). These
MBRP Substance use disorders (relapse prevention) changes reflect modulation in neurocognitive networks
ACT Chronic pain, anxiety and depressive disorders related to intention and motivation, attention and emotion
DBT Borderline personality disorder, substance use disorders regulation, extinction and reconsolidation, prosociality,
non-attachment, and decentering (Vago and Silbersweig,
MBSR, mindfulness based stress reduction; MBCT, mindfulness based cognitive therapy;
2012). Hölzel et al. proposed that mindfulness enacts its
MBRP, mindfulness based relapse prevention; ACT, acceptance and commitment
therapy; DBT, dialectical behavioral therapy. effects through plastic changes of mental and brain functions
related to attention regulation, body awareness, emotion
regulation and self-perspectives (Hölzel et al., 2011a).
is designed for preventing relapse in substance use disorders Recently, Lutz et al. developed a multidimensional model
(Bowen et al., 2010). Available studies have demonstrated its for understanding mindfulness in expert meditators and
efficacy in reducing relapse into drug and drinking use, as well as MBIs, proposing a neurophenomenological “matrix model”
substance usage after a period of abstinence (Bowen et al., 2014; in which categorical orthogonal dimensions, including object
for summary of results, see Table 1). orientation, dereification and meta-awareness, are central
Acceptance and commitment therapy (ACT) is a particular cognitive mechanisms underlying contemplative practices (Lutz
psychotherapeutic orientation developed from behavioral et al., 2015; for summary of models, see Table 2).
analysis, with mindfulness and acceptance as core principles As can be seen, the nature and usage of the construct of
(Hayes et al., 1999), whose effectiveness is similar to that of mindfulness are complex and elusive. In order to understand
cognitive behavioral therapy (CBT) for relevant mental disorders the myriad of studies reviewed in this article, it’s necessary
(A-Tjak et al., 2015). Dialectical behavioral therapy (DBT) was to clarify the different usage of the mindfulness construct.
developed within a CBT framework, and combines mindfulness Dispositional mindfulness is understood as a mental trait or
and ACT elements. It is organized as a yearlong program, stable characteristic of personality, which can vary between
targeting self-harm, and chronic suicidal behavior in BPD and within individuals across time. Mindfulness as practice
(Linehan, 1993). Systematic reviews of ACT find decreases in refers to the concrete practice of mindfulness meditation,
impulsivity and suicidal attempts, and improvements in general the deployment (and training) of a non-elaborative (non-
mental health (Stoffers et al., 2012). Interestingly, for the MBIs conceptual), present-centered, exploratory and non-judgmental
clinical programs, the central aim is to target dysfunctional (non-valorative) awareness. Mindfulness as a state corresponds to
strategies of emotion regulation, which are claimed to drive the the actual proper first-person experience of the non-elaborative,
maintenance and recurrence of these disorders. In this sense, the present-centered, non-judgmental awareness (Chambers et al.,
claim is that mindfulness might re-establish emotion regulation 2009; Davidson, 2010).
capacities, which leads to symptomatic and clinical recovery. Although most of these models include cognitive, self-
awareness, emotional, and attitudinal components, none of them
provide an in-depth understanding of the relationship between
PSYCHOLOGICAL MECHANISMS OF mindfulness and emotion regulation changes. As can be derived
EMOTION REGULATION INVOLVED IN from previous section, a lot of clinical evidence indicates that
MINDFULNESS MBIs seem to be particularly effective in psychological conditions
characterized by different forms of emotion dysregulation
Despite the effectiveness of MBIs in different psychological (see Table 1). In accordance with this, authors studying the
disorders, the underlying psychological and neurobiological psychological mechanisms underlying mindfulness as a trait or
mechanisms are still unclear. Several authors have proposed as a practice have focused specially on the relationship between
psychological models to account for the therapeutic effects of mindfulness and its capacity to enhance emotion regulation as a
MBIs. Shapiro et al. claim that mindfulness might act through key route to yielding mental health benefits.
changing attention, intention, and attitude (Shapiro et al., 2006).
Others suggest that positive effects of MBIs could be explained Cross-Sectional Studies
by mechanisms such as observing, describing, acting with Studies measuring dispositional mindfulness consist of cross-
awareness, non-judging of inner experiences, and non-reactivity sectional surveys using self-report scales in a healthy population.
to inner experiences (Baer et al., 2006). Based on an integration The frequency of these studies has grown exponentially and
of Buddhist psychology and empirical evidence, Grabovac et al. their scope has moved beyond psychiatry and psychology
proposed a model in which changes in acceptance, attention issues to include several other positive health-related outcomes.
regulation, ethical practice, and attachment/aversion to feelings For example, recent studies suggest that higher dispositional

TABLE 2 | Psychological and neurocognitive models of mechanisms of MBIs.
Author Type of model Components
Shapiro et al., 2006 Psychological Attention, attitude, intention

Baer et al., 2006 Psychological Observing, describing, acting with awareness, non-judging of inner experiences and
non-reactivity to inner experiences
Grabovac et al., 2011 Psychological Acceptance, attention regulation, ethical practice and decreased attachment/aversion to
feelings. Final pathway: decreased mental proliferation (rumination narrative based)
Vago and Silbersweig, 2012 Psychological-Neurocognitive Intention and motivation, attention and emotion regulation, extinction, and reconsolidation,
prosociality, non-attachment, and decentering. Final pathway: increasing self-awareness,
self-regulation, self-trascendence
Hölzel et al., 2011a Psychological-Neurocognitive Attention regulation, body awareness, emotion regulation, and change in perspective of the
self. Final pathway: increasing self-regulation
Lutz et al., 2015 Phenomenological-Neurocognitive Primary (orthogonal) dimensions: object orientation, dereification, and meta-awareness.
Secondary qualities: aperture, clarity, stability, and effort
mindfulness is correlated to improved self-care behaviors attention and acceptance of experience (for mindfulness), clarity
(Slonim et al., 2015), and among people with adverse childhood about one’s internal experience, and the ability to manage
experiences, mindfulness as a trait is related to fewer medical negative emotions (for emotion regulation). A path analysis
conditions, and better health behaviors (Whitaker et al., 2014). supported the stance that mindfulness (including the factors
Giluk performed a meta-analysis of 29 studies investigating “present-centered attention” and “acceptance of experience”),
the relationship between mindfulness and personality (Big Five) through clarity about one’s own experience, improves the ability
and aspects of affect/mood, finding a negative correlation to deal with negative emotions (the model had a good data
between mindfulness, neuroticism and negative affect, and a fit, having a RMSEA of 0.059; p < 0.0001). The authors also
positive correlation between mindfulness and conscientiousness found that clarity about experience was negatively correlated to
and positive affect (Giluk, 2009). Feltman et al., in a study with rumination and psychological distress, and positively related to
289 participants, found that mindfulness and neuroticism were flourishing (Coffey et al., 2010). Acknowledging methodological
independent and inverse predictors of depressive symptoms and limitations, studies using dispositional mindfulness as a trait or
trait anger; importantly the relationship between neuroticism personality characteristic (statistically as independent variable
and symptoms was stronger with low mindfulness, suggesting or predictor) provide interesting preliminary evidence that
that mindfulness might play a role in buffering the negative mindfulness, even though partially overlapping with emotion
emotionality of neuroticism (Feltman et al., 2009). In line with regulation constructs, might exert its beneficial salutary effects
this, Wupperman et al. found that deficits in mindfulness predict through higher emotion regulation capacities.
borderline symptoms in a healthy population, independently of
neuroticism (Wupperman et al., 2008). Longitudinal Studies
Other studies have evaluated what factors mediate the In the area of clinical and psychotherapy research, the question
effect of mindfulness on emotion symptomatology. Bao et al. of change mechanisms, or “active ingredients,” that drive
found a mediation effect of mindfulness, through increases therapeutic effects has been a central concern over the last 20
in emotional intelligence (including factors such as emotion years (Kazdin, 2007; Nock, 2007). As we stated in previous
regulation) over perceived stress (Bao et al., 2015). Selby et al. sections, hundreds of longitudinal studies have demonstrated the
looked at how borderline symptoms predict low mindfulness efficacy of MBIs in a healthy or clinical population, but also
levels. Performing a bootstrapping mediation analysis revealed a studies have evaluated change factors that might mediate the
significant effect of rumination as a mediator between borderline salutary effects of these interventions.
features and mindfulness deficits, indicating the maladaptive role Recently, Gu et al. performed a systematic review and
of rumination as a regulatory strategy (Selby et al., 2016). These meta-analysis only of MBSR and MBCT studies that included
results are congruent with intervention studies that highlight the mediation analysis. Starting from 169 trials and ending with
positive effect of DBT and ACT in the BPD population (Gratz and 20 included in further analyses, the authors found consistent
Gunderson, 2006; Stoffers et al., 2012). and strong evidence of emotional and cognitive reactivity,
Looking to further clarify and understand psychological repetitive negative thinking (such as rumination and worry),
mechanisms of mindfulness, Coffey et al. conducted a and mindfulness itself as change factors/mechanisms. Only for
correlational study with 399 healthy people using the five- mechanisms with sufficient studies (mindfulness and repetitive
factor mindfulness questionnaire, the difficulties in emotion negative thinking) was quantitative synthesis using two-
regulation scale and the trait meta-mood scale. Using factor stage meta-analytic structural equation modeling used, further
analysis and structural equation modeling, the authors found that confirming mindfulness and rumination/worry as mediators
mindfulness and emotion regulation corresponded to shared and of the effects of MBIs (Gu et al., 2015). In the same vein,
distinct constructs, distinguishing four factors: present-centered intending to understand change mechanisms using MBCT

TABLE 3 | Evidence-based putative psychological mechanisms of MBIs (pre-SMA and SMA) and parietal cortex are involved in explicit
(MBSR/MBCT). emotion regulation. These regions conform to the so-called
Author Emotional Cognitive Attitudinal
central executive network (CEN), usually involved in top-down
emotion regulation, but also in attention and voluntary cognitive
Gu et al., 2015 <Emotional reactivity <Cognitive reactivity >Mindfulness control. Finally, the ventral-anterior cingulate cortex (vACC) and
<Rumination the ventro-medial prefrontal cortex (vMPFC) are involved in
<Worry implicit emotion regulation, the outside of awareness processing
of emotion, but also in encoding subjective value of the stimuli
Van der Velden et al., >Self-compassion >Meta-awareness >Mindfulness or condition experienced by the subject (Frank et al., 2014; Kohn
2015
et al., 2014; Etkin et al., 2015). From now on, we will refer to the
<Worry
explicit emotion regulation system as the top-down system, and
<Rumination to the emotion generation and the implicit emotion regulation
systems as both part of a bottom-up system, since both feed up
the top-down system with information regarding arousal, visceral
for recurrent depressive disorder, Maj van der Velden et al. homeostasis, aversiveness and rewardingness of a given stimuli or
performed a systematic review of mediation studies. Out of 23 situation, among others.
studies, 12 showed that mindfulness skills, worry, rumination, It has been stated that different emotion regulation strategies
self-compassion and meta-awareness mediated or predicted might differentially activate these brain systems implicated in
treatment outcomes of MBCT (Van der Velden et al., 2015). emotion regulation processes. For example, Dörfel et al. found
From these meta-analytic reviews, including high-quality that detachment, distraction (two forms of reappraisal), and
RCT mediation studies, it is possible to state that mindfulness, expressive suppression increase brain activation in the same
emotional and cognitive reactivity, rumination/worry, self- regions of the right fronto-parietal network, reducing activation
compassion, and meta-awareness might be mechanisms of the left amygdala. This suggests a common underlying
underlying the therapeutic effects of MBIs (for summary neural process for these strategies, but somewhat contrary to
of mechanisms, see Table 3). On the one hand, increases theoretical predictions, since expressive suppression as a less
in mindfulness, self-compassion, and meta-awareness might adaptive strategy might have a different neural correlate from
account for adaptive emotion regulation strategies; on the reappraisal strategies. Interestingly, only reinterpretation induced
other hand, decreases in emotional, cognitive reactivity, a different activation pattern, recruiting the left vLPFC and
and rumination/worry might represent the dismantling of orbitofrontal gyrus, but not decreasing amygdala activation
dysfunctional emotional-cognitive and self-processing strategies (Dörfel et al., 2014). In another study comparing reappraisal
of emotion regulation. This evidence is concordant with the and affect labeling, authors found a common activation pattern
work of Aldao et al. in which avoidance, rumination, and including activation in the right and left dLPFC, right and left
suppression as emotion regulation strategies were correlated to vLPFC, and pre-SMA, and decreased amygdala and vMPFC
anxiety, depression, and eating disorders (Aldao et al., 2010). activation (Burklund et al., 2014). Recently, a meta-analysis of 48
Therefore, MBIs might target specific emotion regulation deficits studies of cognitive reappraisal emotion regulation neuroimaging
of emotion-related disorders. studies concluded that this strategy particularly activates the
bilateral dLPFC, vLPFC, dMPFC, posterior parietal cortex,
and left-middle temporal gyrus, and deactivates the amygdala
NEURAL MECHANISMS OF EMOTION bilaterally. Clearly involving the explicit emotion regulation
REGULATION INVOLVED IN network. Unexpectedly, no other regions related to emotion
MINDFULNESS reactivity decreased their activation level during reappraisal
down regulation (Buhle et al., 2014).
As we have stated before, emotion regulation can be defined Interestingly, some studies have demonstrated that the top-
as all the conscious and non-conscious strategies we use to down or explicit emotion regulation system (dLPFC, vLPFC,
increase, maintain or decrease one or more components of parietal cortex) can also be involved in generating emotional
an emotional response (Gross, 2001), including implicit, non- states and not only in controlling them, in conjunction or in
conscious, and automatic processes, as well as explicit, voluntary parallel with the implicit emotion generation system (Ochsner
and conscious mental processes (Gyurak et al., 2011). From et al., 2009; McRae et al., 2012). In particular, in two studies,
a neural perspective, these processes are realized by different applying cognitive reappraisal to emotions generated via implicit
and complex distributed brain systems. Subcortical regions stimulation resulted in a paradoxical increased activation of the
like the amygdala, periaqueductal gray, ventral striatum (VS), amygdala (Herwig et al., 2010; McRae et al., 2012). In Herwig
anterior insula (AI), and dorsal-anterior cingulate cortex (dACC) et al.’s study, the usage of emotional body-awareness strategy
are involved in emotional reactivity, as emotion generation decreased amygdala activation compared to reappraisal strategy
regions leading changes in arousal and valence regarding the (Herwig et al., 2010). These studies highlight the question of
triggering stimuli. Cortical regions such as the dorso-lateral whether top-down emotion regulation strategies are always the
prefrontral cortex (dLPFC), the ventro-lateral prefrontral cortex most appropriate, and whether there are other effective forms of
(vLPFC), the pre-supplementary and supplementary motor area emotion regulation that are not based on top-down mechanism.

Of particular interest for the mindfulness-based emotion because of the nature of brain structural imaging, it is not
regulation field is the notion of bottom-up emotion regulation. At possible to derive any information about brain regions’ functions.
the brain mechanisms level, the main assumption of this model Another limitation of these studies is the variability of hours of
is that the bottom-up systems implying emotional generation meditation practice within this population, ranging from 1,000 to
regions (like the amygdala, dACC and AI) and implicit emotion 10,000 or more hours. Nevertheless, they might offer preliminary
regulation regions (like the vMPFC) can also be modulated evidence of the effects of long-term mindfulness practice on brain
without the involvement of cognitive control (like the v- plasticity.
d LPFC), or semantic processing regions (temporal cortex). During the last few years, longitudinal studies have assessed
Several authors have argued that mindfulness might exert a the impact of MBIs on brain morphology, particularly the
unique emotion regulation strategy, termed “mindful emotion MBSR 8-week program. Hölzel et al., using MRI voxel-based
regulation,” different from cognitive reappraisal (based on top- morphometry (VBM), found changes in gray matter density in
down system), mainly through the privileged engagement of the left hippocampus, posterior cingulate cortex, right temporo-
these bottom-up emotion regulation systems (Chambers et al., parietal junction (TPJ), some small regions in the brainstem,
2009; Farb and Segal, 2012; Chiesa et al., 2013; Grecucci and cerebellum (Hölzel et al., 2011b). In a similar uncontrolled
et al., 2015a). Nevertheless, whether mindfulness-based emotion longitudinal study with MBSR, the authors found that decreases
regulation is a unique phenomena, and whether it only relies in perceived stress were correlated to a decreased gray matter
on the involvement of bottom-up systems excluding cognitive density in the right amygdala (Hölzel et al., 2009). They also
control regions (top-down systems), and what the exact brain found a correlation between major psychological well-being
signature of mindfulness is as an emotion regulation strategy, and plastic changes in the brainstem (Singleton et al., 2014).
among other questions, are still a matter of debate and will be Santarnecchi et al. performed a controlled longitudinal study
addressed in the following sections of the article. with MBSR, finding a significant increase in cortical thickness in
two clusters: the right SSC and right paracentral lobule, and AI
and right inferior frontal gyrus (operculum). The authors found
Structural Brain Changes in Mindfulness a significant interaction between structural changes in the right
Experts and Mindfulness-Based insula and a decrease in alexithymia levels, suggesting “body or
Interventions interoceptive awareness” as a possible mechanism responsible for
Several studies have investigated the effect of MBIs and long-term salutary effects of mindfulness practice (Santarnecchi et al., 2014).
mindfulness meditation practice using structural brain imaging, These studies suggest that an 8-week MBI (MBSR)
like morphometry-based magnetic resonance imaging (MRI) might induce neuroplastic changes in key areas for
techniques. Cross-sectional design studies comparing healthy emotional reactivity (amygdala, insula), body awareness or
controls with expert meditators (EMs) from different meditation interoception/exteroception (insula, somatosensory cortex),
traditions have demonstrated structural MRI changes in: the self-consciousness (posterior cingulate cortex, pons), mood, and
hippocampus (Hölzel et al., 2008; Luders et al., 2009; Kang et al., arousal regulation (brainstem regions—locus coeruleus, and
2013); right anterior insula (AI; Lazar et al., 2005; Hölzel et al., raphe nuclei), perspective taking (TPJ) and memory systems
2008); orbitofrontal cortex (OFC; Hölzel et al., 2008; Luders et al., (hippocampus, cerebellum). Interestingly, none of these studies
2009; Kang et al., 2013); anterior cingulate cortex (ACC; Grant suggest changes in PFC areas or regions involved in the top-
et al., 2013); left temporal pole (TP; Hölzel et al., 2008; Luders down emotion regulation system, thereby indicating that salutary
et al., 2009; Kang et al., 2013); left frontal gyrus (Vestergaard- effects of MBI might be mediated mainly by changes in particular
Poulsen et al., 2009; Kang et al., 2013); right frontal sulcus relevant subcortical and cortical regions related to bottom-up or
(Lazar et al., 2005); corpus callosum (Luders et al., 2012; Kang non-emotion regulation related functional systems.
et al., 2013); and regions in the brainstem (Vestergaard-Poulsen
et al., 2009). Moreover, a study using machine learning structural
pattern recognition analysis estimated that brains of meditators Functional Brain Changes in Emotion
were 7.5 years younger than matched control subjects (Luders Tasks in Mindfulness Studies
et al., 2016). Dispositional Mindfulness
As can be seen, covering a wide range of brain regions, Cross-sectional studies in healthy populations have investigated
according to recent reviews and meta-analysis of neural bases how individual differences in mindfulness as trait might be
of emotion regulation (Frank et al., 2014; Kohn et al., 2014; related to specific brain functions during emotion elicitation
Etkin et al., 2015), would partially overlap with emotion reactivity task experiments. Creswell et al., in an affect labeling task
(AI, ACC), and with implicit emotion regulation regions (OFC during fMRI, found that levels of dispositional mindfulness
and vMPFC), and very loosely with explicit emotion regulation were related to higher activations in the right vMPFC and right
(medial PFC, but not lateral PFC regions) systems. From this, vLPFC and major deactivation of the right amygdala (Creswell
if mindfulness meditation would involve cognitive reappraisal, et al., 2007). In a similar study, participants were asked to
or top-down emotion regulation strategies, one would expect observe emotional faces during fMRI, and higher levels of DM
changes in lateral PFC morphometry. It is important to note that were correlated to less amygdala reactivity. Using resting-state
due to the design of the studies, it is not possible to infer causality functional connectivity (rs-fMRI) analysis, the authors found
between brain changes and long-term meditation practice; also, a relationship between higher dispositional mindfulness and

decreased connectivity within the midline regions, including time, no involvement of the PFC was detected (Westbrook et al.,
the PCC and MPFC (Way et al., 2010). Importantly, the 2013). Interestingly, sg-ACC, AI, and VS correspond to emotion
midline regions like the MPRC, PCC, precuneus, ACC, and generation regions, but are also implicated in other relevant
parietal cortex are part of the so-called default mode network affective functions such as craving and reward processing (VS),
(DMN; Raichle and Snyder, 2007), which has been related to processing of salient stimuli and interoception (AI), and the
mind-wandering (task-unrelated thought) and self-referential subjective encoding of value and processing of emotional conflict
processing (Qin and and Northoff, 2011). Brown et al. assessed (sgACC; Wilcox et al., 2016).
46 participants with an electro-encephalogram (EEG) while Lutz et al., in a cross-sectional study with healthy participants,
viewing emotionally laden pictures, particularly looking at compared one group applying mindfulness with a no-strategy
the late positive potential (LPP) as a marker of affective group while looking at a set of emotional pictures during
processing. Authors found that higher dispositional mindfulness fMRI. When expecting negative pictures, the mindfulness group
correlated to lower LPP during high-arousal negative images displayed increased activation of the left AI, right and left
(Brown et al., 2013). Finally, Kong et al., using rs-fMRI and dMPFC, and left dLPFC. During perception of negative pictures,
local synchronization measurements (estimated by regional the mindfulness group showed reduced activation in the right
homogeneity) with 290 subjects, found that major dispositional amygdala and parahippocampal gyrus, with no involvement
mindfulness correlated to local synchronization in the right of the PFC (Lutz J. et al., 2013). The same researchers also
insula, left OFC, left parahippocampal gyrus (regions involving compared groups using mindfulness vs. cognitive reappraisal
emotion reactivity, implicit emotion regulation), and decreased using the same emotional task as in fMRI. During the expectation
local synchronization with the inferior frontal gyrus (IFG; of negative pictures, both groups showed a similar pattern
related to explicit emotion-regulation). Furthermore, levels of of activation of the MPFC and the amygdala, and during
local synchronization in the OFC predicted positive emotions, the perception of negative images, decreased activation of
and in the IFG predicted a sense of meaning and purpose in the head of the right caudate in the mindfulness group was
life, both effects mediated by DM (Kong et al., 2016). This the only difference (Opialla et al., 2014). Interestingly, the
study suggests that local synchronization in key regions of first experiment comparing mindfulness vs. baseline conditions
emotion regulation might engage differently in subjects high suggests a bottom-up (targeting emotion reactivity regions, with
in dispositional mindfulness, accounting for positive emotions’ no changes in PFC) mechanism of mindfulness as emotion
salutary effects. Also it shows no correlation between lateral regulation strategy; instead, when adding an active regulatory
PFC local synchrony and dispositional mindfulness in emotion strategy as comparison, it is almost impossible to differentiate at
regulation-related variables, suggesting that individuals high the neural level between the two emotion regulation strategies.
in dispositional mindfulness might engage in emotion-related However, the observed deactivation of the right caudate head
processes involving different regulatory systems than top-down might index decreased engagement of automated cognitive and
ones (for summary of results, see Table 4). motor responses (Parent and Hazrati, 1995), which might be
Interestingly, these findings are concordant with linked to decreased automatic cognitive reactivity, known as a
psychological studies linking dispositional mindfulness to mindfulness mechanism (Gu et al., 2015).
better emotional life outcomes (positive affect and emotional Interestingly, this draws attention to the fact that even a
intelligence and minor neuroticism, negative affect, rumination, short mindfulness induction, in people naive about meditation,
and borderline symptoms) thereby providing preliminary can induce a distinguishable bottom-up brain activation pattern
support for the construct validity of DM. These studies face many when comparing mindfulness as a strategy to baseline or no-
limitations, such as the difficulty in deriving causal inferences, strategy condition. Nevertheless, when compared to cognitive
and disentangling relevant confounders such as psychological reappraisal, differences seem to vanish. This suggests that
traits and biological differences. Another problematic claim of mindfulness meditation in naive practitioners is performed with
these studies is the assumption that dispositional mindfulness the engagement of widespread brain regions including top-down
really reflects daily-life mindful attitudes. At this time, to the best and bottom-up regulatory systems. From the clinical perspective,
of our knowledge, no study has empirically clarified this point. these studies provide a valuable outlook for understanding
neurobiological substrates of brief meditation practices, which
Mindfulness Inductions are central components of many MBIs, like MBCT, ACT, or DBT,
Studies using brief meditation practice, or mindfulness that intend to elicit “mindfulness states” to face difficult emotions
inductions, have started to explore the clinical utility and emotion dysregulation states.
(effectiveness) and neural underpinnings of these types of As previously stated, these studies share limitations with
interventions. Westbrook et al. performed a cross-sectional study cross-sectional design studies. These investigations raise
with smokers looking to stop smoking. Participants were asked particularly relevant problems in the discussion of mindfulness
to watch specific craving-inducing images during fMRI, using and emotion regulation mechanisms, starting with the question
“mindful attention” vs. “passive viewing” as strategies. When of the acquisition of the so-called mindfulness emotion
applying “mindful attention,” subjects reported less craving regulation strategies—in other words, when and how a person
impulse; additionally, they presented decreased activation in the acquires the capacity to elicit a “mindfulness state,” different
subgenual ACC (sg-ACC), and reduced functional connectivity from other mental states. And also, when and how a person
between this same region and bilateral AI and VS. At the same acquires the capacity to use mindfulness as an emotion

TABLE 4 | Summary of neuroimaging studies using emotion-task experiments in different mindfulness conditions.
Mindfulness condition Study design Population Sample size: Experimental task and Main finding: summarized in terms of References
(different models) vs mindfulness (M) neuro-imaging method: resting brain, and/or physiological response
control condition vs control state functional magnetic changes
(Waiting list, or active groups (C) resonance imaging (rs-fMRI), task
Guendelman et al.
control) based functional magnetic

resonance imaging (fMRI),
electroencephalography (EEG)
DISPOSITIONAL MINDFULNESS
Dispositional Cross sectional/Uncontrolled Healthy M: 27 Affect labeling task during fMRI Level of DM mediates the relationship Creswell et al., 2007
mindfulness study between right vMPFC, right vLPFC
activation and right amygdala deactivation
Frontiers in Psychology | www.frontiersin.org

Dispositional Cross sectional/Uncontrolled Healthy M: 27 Viewing negative emotional faces Higher DM correlated with less amygdala Way et al., 2010
mindfulness study during fMRI + rs-fMRI reactivity. Also with less resting connectivity
in midline brain regions (self-referential
processing)
Dispositional Cross sectional/Uncontrolled Healthy M: 46 Viewing negative/positive pictures Higher DM correlated to lower LPP during Brown et al., 2013
mindfulness study during EEG (LPP: late positive potential) high-arousal negative emotions
Dispositional Cross sectional/Uncontrolled Healthy M: 290 rs-fMRI—local synchronization Higher DM correlated to local synch in left Kong et al., 2016
mindfulness study OFC, left parahippocampal gyrus, right
insula. Local synch in OFC-predicted
positive affect, and in IFG-predicted
purpose/meaningful life
MINDFULNESS INDUCTION
8
Mindfulness induction Cross sectional/Uncontrolled Smokers looking for M: 47 Cue-induced craving during fMRI. Reduced neural activity in sg-ACC Westbrook et al., 2013
study treatment to stop [craving-related—emotion reactivity region]
smoking and a reduced functional connectivity
between this same region with the bilateral
insula and ventral striatum with no direct
involvement of PFC regions (*)
Mindfulness induction Cross Healthy M: 24/C: 22 Cued expectation and perception of During expectation major activations in Lutz A. et al., 2013
sectional/Non-randomized negative pictures during fMRI prefrontal regions: left AI, right and left
controlled study dMPFC and left dLPFC. During perception
reduced activation in right amygdala and
parahippocampal gyrus [emotion processing
reactivity] (*)
Mindfulness induction vs Cross Healthy M: 24/C: 23 Cued expectation and perception of Both groups: similar activity of the m-PFC Opialla et al., 2014
Reappraisal strategy sectional/Non-randomized negative pictures during fMRI and the amygdala. Major activations in MI
controlled study group, during expectation: vLPFC, vLPFC,
Supramarginal gyrus and left insula. During
perception: major activity in the caudate in
the cognitive group
MINDFULNESS-BASED INTERVENTIONS
Mindfulness-based Cross Healthy M: 20/C: 16 Self-reference task during fMRI Significant difference in the neural correlates Farb et al., 2007
stress reduction sectional/Non-randomized of the self-reference task, during experiential
controlled study (novice vs focus an increased activation in right brain
those who attended the regions: lateral PFC, insula, second
course) somatosensory area, and IPL
Mindfulness and Emotion Regulation
March 2017 | Volume 8 | Article 220

(Continued)
TABLE 4 | Continued
Guendelman et al.

Mindfulness-based Longitudinal/Non-randomized Healthy M: 20/C: 16 Sadness induction paradigm during MBI group changed activation pattern in key Farb et al., 2010
stress reduction vs controlled study fMRI emotion regulation regions: major activation
Waiting list in the right anterior insula, r-lPFC and
sg—ACC.

Mindfulness-based Longitudinal/Non-Controlled Social Phobia M: 16 Breath focus task during fMRI Reduced amygdala activity, major activation Goldin and Gross,
stress reduction trial in precuneus, SPL, IPL compared to 2010
distraction focus task
Mindfulness Training (4 Longitudinal/Non-Controlled Healthy (pain) M: 15 Breath focus meditation during noxious MBI reduction in pain intensity: major Zeidan et al., 2011
days) trial stimulation task in fMRI activation in ACC, anterior insula. MBI
reduction in pain unpleasantness: major
activation in OFC and thalamus (*)
Mindfulness-based Longitudinal/Randomized Social Phobia M: 31/C: 25 Self-reference task during fMRI MBIs during negative self-view: major Goldin et al., 2012
stress reduction vs controlled trial activation in PCC, and dMPFC
Aerobic exercise activity-associated less social anxiety
disability and mindfulness level
Mindfulness-based Longitudinal/Randomized Social Phobia M: 31/C: 25 Emotion regulation of negative MBI regulating negative self-beliefs: fewer Goldin et al., 2013
9
stress reduction vs controlled trial self-beliefs task during fMRI negative emotions, major activation in R-IPL,
Aerobic exercise R-SPL
Mindfulness-based Longitudinal/Randomized Generalized Anxiety M: 15/C: 11 Affect labeling of emotional expressions Both groups less amygdala activation. MBI Hölzel et al., 2013
stress reduction vs controlled trial Disorder during fMRI major activation in vLPFC. Increase
Stress management functional connectivity between amygdala
education and PFC regions
Mindfulness Training (6 Longitudinal/Randomized Healthy M: 30/C: 31 Affective Stroop conflict resolution task Both groups improved significantly in a Allen et al., 2012
weeks) vs Shared controlled trial during fMRI response inhibition task. MBI reduced
reading and listening emotional interference, in negative emotion
group processing: increased bilateral dLPFC, right
anterior insula and m-PFC (*)
Mindfulness Training (8 Longitudinal/Randomized Healthy M: 12/C: Observation of emotional pictures In MBI: decrease in right amygdala Desbordes et al., 2012
weeks) vs Compassion controlled trial 12/Compassion during fMRI activation (all valences). In Compassion
training vs Health Training: 12 Training: trend increase in right amygdala
discussion group response in negative pictures (*)
Mindfulness Training (4 Longitudinal/Randomized Healthy (pain) M: 80 Pain regulation strategy during noxious MBI reduction in pain intensity: major Zeidan et al., 2015
days) vs Sham controlled trial (four-arm) stimulation task in fMRI activation in sg—ACC, anterior insula, OFC.
mindfulness vs Placebo Placebo analgesia: major activation in
vs Control DLPFC and secondary somatosensory
cortex (*)
(Continued)

TABLE 4 | Continued
Guendelman et al.

EXPERT MEDITATORS
Tibetan Buddhist monks Cross sectional/Case-control Healthy M: 14/C: 16 Auditory stimuli during focus attention EM: amygdala deactivation Brefczynski-Lewis
study task in fMRI et al., 2007

Tibetan Buddhist monks Cross sectional/Case-control Healthy M: 15/C: 15 Auditory stimuli during active EM: increased activation in the anterior Lutz et al., 2008a
study compassion meditation in fMRI insula and ACC, proportional to compassion
experience intensity
Zen Western vs novices Cross sectional/Case-control Healthy M: 12/C: 8 Observation of emotional pictures EM during meditation: major deactivation of Taylor et al., 2011
meditators study during active meditation in fMRI m-PFC and PCC. Relative deactivation of
amygdala and insula vs novice meditators.
Novice during meditation: downregulation of
amygdala
Zen Western Cross sectional/Case-control Healthy M: 13/C: 13 Noxious stimulus during fMRI EM during pain: reduced activation in PFC, Grant et al., 2011
study amygdala, hippocampus. Major activations
in ACC, insula, thalamus.
Vipassana Cross sectional/Case-control Healthy M: 17/C: 17 Noxious stimulus during fMRI EM during pain in meditation: reduced Gard et al., 2012
study activation in lateral PFC, major activation in
10
ACC, R-posterior insula
Tibetan tradition Cross sectional/Case-control Healthy M: 14/C: 14 Noxious stimulus during fMRI EM: equal pain, less unpleasantness. During Lutz A. et al., 2013
study pain: major AI, ACC. Minor baseline
activation AI, ACC, amygdala
Buddhist Western Cross sectional/Case-control Healthy M: 18/C: 26 Dictator Game (DG) and Ultimatum EM: in DG reduced arousal, distress and Grecucci et al., 2015b
study Game (UG) during Skin Conductance SCL. In UG accept more unfair offers
Level (SCL)
Buddhist Western Cross sectional/Case-control Healthy M: 26/C: 40 Ultimatum game during fMRI EM: in UG accept more unfair offers. Major Kirk et al., 2011
study activation of the posterior insula
(interoception) versus anterior insula
(emotion reactivity) in controls; major
activation in somatosensory and posterior
superior temporal cortex
Zen Western Cross sectional/Case-control Healthy M: 34/C: 44 Monetary incentive delay during fMRI EM during reward anticipation: reduced Kirk et al., 2011
study activation in caudate nucleus, major
activation in bilateral posterior insula. During
reward receipt: reduced activation in vMPFC
Buddhist Western Cross sectional/Case-control Healthy M: 28/C: 30 Passive conditioning task during fMRI EM during reward prediction: reduced Kirk et al., 2015
study positive and negative prediction error BOLD
in putamen. Major activation in posterior
insula
*Finding indicates bottom-up mechanisms.


regulation strategy. Finally, the question of how this learning Other studies have evaluated the impact of MBIs on pain
process can be distinctly measured from behavioral and brain processing. Zeidan et al. performed a longitudinal uncontrolled
signatures. These are central questions that future studies need to study with 4-day MBI training, using Artetial Spin Labeling
unravel. (ASL), a technique for estimating cerebral blood flow with MRI
across time points. After the intervention, during a breathing
Mindfulness-Based Interventions: Longitudinal meditation task, the authors found decreased perfusion of the
Studies on Emotion, Pain, and Anxiety MPFC and PCC (DMN), and a major activation of the AI, ACC,
Over the last few years, longitudinal studies using fMRI have used pre-SMA, OFC, VS, SSC, and posterior insula (PI). During a
a myriad of experimental tasks investigating emotion regulation pain induction paradigm, minor activation of the contra-lateral
changes secondary to MBIs. Farb et al. studied the impact of SSC and increased activation in the ACC, AI, PI, and fronto-
MBSR using fMRI under a sadness induction paradigm. After parietal operculum were reported. It is worthy of note that
the intervention, the mindfulness group changed the activation participants reported a significant decrease in pain intensity and
pattern in key diverse emotion regulation regions: comparatively unpleasantness (Zeidan et al., 2011). Later, the same authors
increased activation in the right AI, right LPFC and sg-ACC. performed a four-arm RCT comparing MBI vs. placebo vs.
The control group showed major activation in the left PFC, sham mindfulness using a pain induction paradigm with ASL
left superior temporal sulcus (STS), precuneus, and PCC, areas MRI. Interestingly, all groups showed a significant reduction in
usually involved in self-awareness and semantic processing (Farb pain intensity and unpleasantness, but the MBI demonstrated
et al., 2010). From the same lab, using a self-referential task a unique brain mechanism including greater activation of
(self-narrative vs. self-experiential) during fMRI, an increased the OFC, sg-ACC, and AI. In line with previous evidence,
activation was found in similar right brain regions, LPFC, AI, these studies highlight emotion reactivity (AI, ACC, VS) and
second SSC and inferior parietal lobule (IPL), for the self- implicit emotion regulation (OFC, vMPFC) systems as the main
experiential focus. Conversely, a self-narrative focus engaged emotion regulation targets of MBIs, again notably without any
major activation in the left vMPFC, dMPFC, and PCC, all midline major involvement of PFC-related systems (top-down emotion
regions that mainly correspond with the DMN (Farb et al., 2007). regulation).
These studies indicate a different engagement of brain regions Other researchers have explored the effects of MBIs in
during emotion regulation; although both groups displayed top- clinical populations. In one of the first such studies, Goldin
down mechanisms linked to explicit emotion regulation systems and Gross conducted an MBSR longitudinal study with people
(right or left LPFC), only the MBI groups employed regions suffering from social anxiety disorder (SAD). Comparing two
related to emotion reactivity (AI, ACC), interoception (AI) and emotion regulation strategies using an anxiogenic task with
somatosensory awareness (SSC, IPL). negative self-beliefs, the authors found that being breathing-
Attempting to unravel the involvement of different emotion focused (vs. distraction-focused) produced minor negative
regulation systems implicated in mindfulness meditation, Allen emotional experiences, decreases in amygdala activation, and
et al. performed an RCT comparing a 6-week mindfulness increased activation of the PCC, SPL, and IPL (areas involved
training and an active control (sharing and listening training). in top-down emotion regulation, but also in self-awareness
Despite both groups improving significantly in a response and attention processing; Goldin and Gross, 2010). The same
inhibition task, only the MBI group showed reduced emotional authors performed an RCT comparing MBSR with aerobic
interference under an affective Stroop conflict resolution exercise (AE), also in SAD patients, in this case comparing
paradigm (a task known to activate implicit emotion regulation mindful attention (metacognitive perspective of mental content)
processes). The authors found no differences between groups and reacting (thinking according to negative self-beliefs) as
in behavioral and neural activations during negative affect strategies for dealing with negative-self-belief-induced emotions.
processing. Nevertheless, the greater amount of mindfulness During the task, the MBSR group reported fewer negative
practice predicted increased activation of bilateral dACC, right emotions, and showed differential engagement of attention
AI, and MPFC during implicit negative emotional processing, regulation areas, with increased activation of the right IPL and
suggesting both implicit and explicit emotion regulation SPL, and decreased activation of the culmen and left lingual
plasticity as mechanisms underlying mindfulness training (Allen gyrus (Goldin et al., 2013), areas involved in the orienting-
et al., 2012). Another RCT study compared the effects of attention network, implicated in early spatial detection of stimuli
an 8-week Mindful Attention Training (MAT) vs. Cognitively (Posner et al., 2006). The authors interpreted this finding as
Based Compassion Training (CBCT) vs. active control while suggesting that MBIs enhance approaching behavior/attention
participants passively viewed affective pictures during fMRI. In a toward anxiogenic stimuli, a core deficit in SAD (Goldin et al.,
region of interest analysis, the authors found decreased activation 2013). In the context of the same trial condition, both groups
in the right amygdala in the MAT group in response to images significantly decreased social anxiety symptoms, disability and
of all valences. Interestingly, a trend increase in activation of negative self-attribution, while also increasing positive self-views.
the right amygdala when viewing negative images in the CBCT Examining the neural correlate of self-views, the MBSR group
group was found, and the extent of this increase was significantly displayed larger responses in the PCC, and dMPFC, which
correlated to reductions in depressive symptoms (Desbordes correlated with minor social anxiety, disability, and increased
et al., 2012). Although not conclusive, both RCT studies provide mindfulness (Goldin et al., 2012). Finally, Hölzel et al. ran an RCT
evidence that MBIs might exert their effects on the level of with generalized anxiety disorder (GAD) patients, comparing
emotion reactivity and implicit emotion regulation. MBSR and psychoeducation treatment groups performing an

emotion labeling task during fMRI. The findings highlighted Other studies have used reward or economic behavioral
small increases in amygdala activation in both groups, and major paradigms for studying emotion processing in EMs. Grecucci
increases of activity in the vLPFC, as well as increased functional et al. compared EMs with a control group contrasting a
connectivity between these regions (Hölzel et al., 2013). These “cognitive” vs. an “experiential” emotion regulation strategy
studies point toward the idea that MBIs target basic cognitive during two monetary distribution tasks. While receiving offers in
processes broadly involved in attention regulation, including the dictator game, EMs showed decreased emotion arousal and
information updating, response inhibition, and goal maintenance physiological reactivity, with no effect of the strategy observed.
(Malinowski, 2013). Interestingly, these are core functions for the While receiving unfair offers in the ultimatum game (UG), EMs
CEN, and for the top-down emotion regulation system (Okon- accepted more unfair offers and performed less punishment,
Singer et al., 2015). In sum, these studies provide evidence particularly during the “experiential” emotion regulation strategy
that MBIs might exert their effects through top-down/cognitive- (Grecucci et al., 2015a). Another study used fMRI during the
control emotion regulation mechanisms. Besides sample size, execution of the UG. Compared to controls, EMs accepted
noteworthy limitations of these studies include the lack of control more unfair offers, and during that particular condition engaged
of basal cognitive deficits in patients, and of personality and a particular functional brain response with greater activation
comorbidity factors, which might influence basal neuroimaging of the PI than the AI, and major activation in the SSC and
results. posterior superior temporal cortex (Kirk et al., 2011). Note that
the PI is preferentially involved in interoception and the AI in
Expert Meditators (EMs): Cross-Sectional Studies on emotion reactivity/generation and emotional awareness (Craig,
Emotion, Pain, and Reward 2009; Gu et al., 2013). These studies show that during socially
Lutz et al. used an annoying auditory task during fMRI, induced negative emotions, EMs showed stronger modulation
comparing Tibetan monks and controls during active of their interactive behavior (less punishment) and greater
compassion meditation. They found increased activity in emotion regulation, which was mediated via increased activation
the AI and ACC, which were proportional to first-person of interoception and exteroception brain regions, modulating
experience of compassion intensity (Lutz et al., 2008a). Using emotion generation regions.
the same experimental task, but during focused-attention Kirk et al. used the monetary incentive delay task in EMs
meditation, researchers also found a direct relationship between during fMRI, looking to disentangle the neural differences
meditation expertise (total hours of practice) and amygdala between anticipation and receipt of monetary reward. Compared
deactivation (Brefczynski-Lewis et al., 2007). Taylor et al. to controls, during the anticipation phase EMs displayed
compared Western EMs with novel meditators using emotional decreased activation of the bilateral caudate, and increased
pictures during fMRI, and observed a decrease in activation activation of the bilateral PI. During the encoding of gains of
levels of the PCC and MPFC (DMN) during active meditation in reward, a minor activation of the vMPFC was seen (Kirk et al.,
EMs. During passive observation, beginner meditators showed 2015), indicating a dampening of the reward system. The same
major amygdala activation increases for negative affective authors used a passive conditioning task (pairing a yellow light
pictures (Taylor et al., 2011). These studies highlight a specific to juice intake) to evaluate how changes in the predictability
modulation of the emotion generation system in EMs during of reward, encoded by the prediction error (PR) neural signal,
emotion tasks. differ between EMs and matched controls. In this task, the delay
Other studies have explored the effects of EMs in pain of the reward decreases PE (negative PE), while the intake of
processing. Gard et al. compared Western EMs with controls, unexpected reward generates an increase in PE signal (positive
contrasting active meditation, and resting state using a pain PE). EMs were found to be less prone to positive and negative
induction paradigm with fMRI. The authors found no differences PE signals in the putamen (part of the striatum and the reward
between groups in pain intensity, but in active meditation system), which again was associated to major activation in the
during pain induction, EMs referred less unpleasantness and PI (Kirk and Montague, 2015). Interestingly, both studies show
a major activation in the right AI and a deactivation in the a specific modulation in value reward processing in the striatum
right and left inferior PFC (Gard et al., 2012). Grant et al. also and vMPFC, from interoceptive body awareness regions (PI) that
compared EMs with controls during a pain induction task in correspond to bottom-up emotion regulation systems, in line
fMRI. EMs showed decreased activation of the PFC, amygdala with the bottom-up mechanism hypothesis of emotion regulation
and hippocampus, and increased activity in the AI, ACC, and changes derived from mindfulness practice.
thalamus. Interestingly, the decreased functional connectivity
between PFC and AI and ACC predicted lower pain in EMs
(Grant et al., 2011). In a similar study, EMs showed lower
INTEGRATING PSYCHOLOGICAL,
baseline activation in the AI, ACC, and amygdala, and during CLINICAL AND NEUROSCIENCE
pain induction higher activation of AI and ACC regions than EVIDENCE ON EMOTION REGULATION IN
controls (Lutz A. et al., 2013). These studies indicate that EMs MINDFULNESS RESEARCH
specifically increases activation of subcortical emotion generation
regions, related to affective processing of pain, and deactivates The field of contemplative science, the scientific study of the
top-down mechanisms, evidencing a unique emotion regulation effects of mindfulness, and contemplative practices in mental
bottom-up mechanism. health and biological functions, is fairly new but growing quickly.

In this article we have focused exclusively on the relationship mindfulness might be mediated by emotion regulation capacities
between mindfulness practices, using diverse empirical models (Coffey et al., 2010). Similarly, dispositional mindfulness has
(dispositional mindfulness, mindfulness inductions, MBIs, and been linked to a higher right PFC, minor amygdala activation and
EMs), and emotion regulation functions from psychological changes in rs-fMRI in regions from all the emotion regulation
and neurobiological perspectives. A range of MBIs have systems (see Table 4). These studies provide evidence of top-
demonstrated utility in several clinical conditions (see Table 1), down regulation mechanisms. As stated early, several limitations
targeting a myriad of emotion dysregulation symptoms (Gotink preclude an unequivocal interpretation of these findings in the
et al., 2015). context of mindfulness and emotion regulation research.
With the aim of understanding mechanisms underlying Two studies using mindfulness inductions (mindfulness
mindfulness health benefits, authors have proposed several as emotion regulation strategy) have provided preliminary
psychological and neurocognitive models (see Table 2) that evidence of direct bottom-up regulation engagement, changing
cover attention, emotion, and self-awareness systems as target the emotion generation system, with no involvement of the
mechanisms (Tang et al., 2015). Here we focused particularly PFC. However, these studies lack an alternative cognitive
on emotion regulation mechanisms targeted by mindfulness emotion regulation strategy for contrasting the specificity of the
meditation, reviewing different studies using psychological and strategy (see Table 4). In addition to the noted methodological
neuroimaging measurements, ranging from correlational to limitations, we argue that using a unique mindfulness induction
randomized longitudinal designs. session might be insufficient for eliciting a “mindful emotion
In the field of mindfulness and emotion regulation, one main regulation” strategy and the recruitment of the bottom-up brain
claim is that mindfulness might elicit a particular type of emotion systems. Secondly, central to this discussion is the question of
regulation strategy often called “mindful emotion regulation” how mindfulness as an emotion regulation strategy is defined and
that relies on bottom-up mechanisms, in contrast to cognitive operationalized. Is it a formal practice, identical or derived from
reappraisal, which relies on a top-down mechanism. Although mindfulness meditation? Or is it a particular state, related to the
there is no single definition, mindful emotion regulation notion of mindfulness as a transient state? We will return to this
is conceived as a unique emotion regulation strategy, that discussion in the next section.
results from encountering diverse emotional states from a Longitudinal studies have yielded mixed results regarding
mindful mental state, which includes awareness and acceptance the involvement of different emotion regulation systems (top-
(Chambers et al., 2009; Farb and Segal, 2012; Chiesa et al., 2013; down vs. bottom-up). Studies with healthy populations using
Grecucci et al., 2015a). In particular, it is stated that bottom-up self-experiential focus recruit emotion-generation (AI, sg-ACC)
emotion regulation strategies (like those implied in mindfulness) and body-awareness (AI, SSC) systems. Well-designed RCTs
don’t require PFC and top-down mechanisms (Chambers et al., with active control groups have mostly (but not exclusively)
2009; Farb and Segal, 2012; Chiesa et al., 2013; Grecucci et al., demonstrated changes in emotion generation (amygdala, AI,
2015a). In terms of neurobiological emotion regulation systems, ACC) and implicit emotion regulation systems (v-MPFC, OFC),
these strategies might rely on modification of implicit emotion while being effective in regulating negative emotions. Clinical
regulation and emotion generation systems, but not on changes studies with anxiety disorder populations have shown major
in the explicit emotion regulation system. In this section, in involvement of explicit emotion regulation systems (see Table 4).
accordance with the reviewed studies, we will assess whether this It is worth noting that these differences might be due to
claim and its assumptions are met. methodological limitations (e.g., simple size), but also to the
Studies measuring structural brain changes in EMs highlight specific cognitive demands of the experimental tasks (such as
changes in the MPFC and diverse subcortical regions, including self-reference, regulation of self-beliefs or affect labeling tasks)
regions devoted to meta-awareness, memory consolidation, that by nature require top-down regulation mechanisms. Overall,
extero-interoception, and emotion regulation (Fox et al., 2014), changes in bottom-up neural mechanisms are in line with the
with no exact matching to bottom-up systems, but with findings of psychological studies of MBIs, in which decreases
no involvement of typical LPFC. Longitudinal studies with in emotional cognitive reactivity, and rumination strategies,
MBIs have also implicated regions typically involved in the and increases in mindfulness skills, self-compassion, and meta-
same functions described above (like the AI and amygdala), awareness emotion regulation strategies, appear to underlie the
but no changes in the MPFC and LPFC have been found, beneficial effects of MBIs (see Table 3).
regions known for top-down emotion regulation. Strikingly, Finally, studies with EMs using emotion and pain paradigms
only AI and brainstem regions overlap between EM and MBIs have consistently demonstrated changes in bottom-up emotion
studies, suggesting neuroplasticity in key areas for emotion generation systems (amygdala, AI, sg-ACC), with reported
generation, interoception, mood, and viscerosomatic processing. deactivations in, or no involvement of, the PFC. In some
As mentioned, no inference about causality (in EM studies), nor studies involving social emotion or reward processing tasks,
about brain functions, can be derived from these studies. EMs displayed increased engagement of interoception brain
Studies measuring dispositional mindfulness have found system (mainly PI), modulating emotion generation, and
negative correlations with negative affect and positive implicit emotion regulation systems of reward-related areas
correlations with positive affect traits; factorial analysis has (caudate, putamen, v-MPFC; see Table 4), providing evidence
pointed out the distinct and interrelated nature of mindfulness of the engagement of a bottom-up emotion regulation system
and emotion regulation as constructs. Mental health outcomes of in EMs.

From the reviewed studies, we argue that there is support should the focus of attention be (external or internal stimuli)?
for the claim that mindfulness practice changes the bottom-up And, in terms of behavior, what exactly should be done to
emotion regulation systems (emotion generation and implicit perform the regulation (approach, stop, or hold back)? From
emotion regulation systems), although this effect diverges a psychological perspective, there is not a clear commitment
across different empirical models dispositional mindfulness, regarding the unique (or common) involvement of attentional or
mindfulness inductions, MBIs and EM studies. In line with emotional or body awareness processes. Thus, in line with clinical
Chiesa et al. (2013), studies with EMs show a clearer engagement evidence (Table 3), it is not clear whether “mindful emotion
pattern of bottom-up systems, suggesting that these types of regulation” is properly a unique emotion regulation strategy, with
strategies are developed through long-term meditation training. a unique neurocognitive underlying mechanism.
However, intervention studies with a RCT design are better In light of this debate, we argue that “mindful emotion
suited for providing evidence about a causal relationship between regulation” entails a variety of emotion regulation processes,
mindfulness training and bottom-up emotion regulation system including top-down processes which are cognitively based,
changes. involving attention and voluntary cognitive control, conscious
monitoring, and explicit regulatory functions; and bottom-up
processes, which are affect driven, based on emotion functions
The Problem of Mindful Emotion that modulates arousal, valence and the encoding of subjective
Regulation value regarding the triggering stimuli. We argue that “mindful
From psychological studies, including theoretical and evidence- emotion regulation” entails as well a variety of emotion
based psychological models (Table 3), as well as neuroimaging regulation strategies, in accordance with the different strategies
studies (Table 4), it becomes evident that mindfulness (in taught within MBIs and EMs trainings. In this context, we
MBIs and EMs) also engages and requires top-down emotion propose a distinction between primarily top-down mindfulness-
regulation. As Lutz et al. stated, mindfulness meditation based emotion regulation strategies and bottom-up mindfulness-
can be conceived as “a family of complex emotional and based emotion regulation strategies. Since emotions are multi-
attentional regulatory strategies developed for various ends” componential processes (Thompson, 1990), and like Gross’s
(Lutz et al., 2008b). From a traditional Buddhist psychology classification of emotion regulation strategies, our distinction is
perspective, the development and refinement of attention based on the primary component of the emotional response that
(attention regulation; Grabovac et al., 2011), and the capacity is targeted and drive the regulation of the emotional state (Koole,
for monitoring and labeling affective states (Analayo, 2003), 2009).
are central for achieving the intended effects of mindfulness Top-down mindfulness-based emotion regulation strategies
meditation. From this viewpoint, and taking into account models correspond to affect labeling, mindful detachment, dereification,
of different emotion regulation brain systems and different meta-awareness, and cognitive reappraisal, among others, for
emotion regulation strategies, the notion of “mindful emotion which cognitions and thought process are the primary targets of
regulation” (Chambers et al., 2009; Farb and Segal, 2012; Chiesa the strategy. Within this group we can find impulses control and
et al., 2013; Grecucci et al., 2015a) seems to imply certain emotion dysregulation managing strategies, like those delivered
problematic aspects. in MBIs (like in DBT and ACT) in which subjects use intentional
The notion of “mindful emotion regulation” entails two efforts to increase their attention and awareness capacities for
problematic aspects. The first refers to the nature and definition better regulation and control of emotions (Linehan, 1993; Hayes
of the construct “mindful emotion regulation” itself, and the et al., 1999). In this group, dereification and meta-awareness
second refers to its brain correlates or engagement/functioning would correspond to more sophisticated strategies, since they
of emotion regulation systems, which we will address separately. involve the development of insight into the nature of the
Although we have extensively shown that emotion regulation thought process itself (e.g., see thoughts not as facts; Dahl
is (somehow) enhanced by mindfulness practice, we argue et al., 2015). Using the process model of emotion regulation by
that the notion of “mindful emotion regulation” has not been Gross, we can understand that increases of mindfulness can
accurately and properly defined. Is “mindful emotion regulation” indeed modulate any of the five stages: selecting or modifying a
a psychological trait, stable in time, that diverges across subjects? situation, deployment of attention, changing cognition (cognitive
Or is it a particular mental practice derived from mindfulness? reappraisal), modulating the experience, and behavior, or
Or is it a mental state, like a transient moment of mindfulness? physiological response (Gross, 2001). This distinction is in line
Generally, the common view across authors is that “mindful with findings in MBIs (Table 4), and by Chiesa et al. (2013), and
emotion regulation” is a somehow unique emotion regulation is consistent with the claim that novel practitioners in MBIs use
strategy, the result of encountering diverse emotional states from primarily top-down emotion regulation strategies.
a mindful mental state, including awareness and acceptance In bottom-up mindfulness-based emotion regulation
(Chambers et al., 2009; Farb and Segal, 2012; Chiesa et al., 2013; strategies, sensory-perception and interoceptive-proprioception
Grecucci et al., 2015a). From a first-person perspective, this are the primary aspects of the emotional response targeted
definition does not make explicit specifications regarding what by the strategies. The bottom-up strategies are characterized
the practitioner should do while engaging within the emotional by the intentional stance to directly feel (instead of think) or
state, only succinctly suggesting the gradual development of to experience, thus targeting primarily the feeling processes
experiential qualities (attentiveness, acceptance, etc.). What (sensory-perception and interoceptive-proprioception).

Bottom-up mindfulness-based emotion regulation strategies nature of emotions (Damasio, 1999; Prinz, 2004). As Colombetti
include concrete experiential explorations that focus for example et al. noted, cognitivist theories of emotions have neglected the
on unimodal body sensations, like feeling the temperature of the role of the body in the generation of emotional states (Colombetti
skin, or exteroceptive sensations, feeling the peri-personal space and Thompson, 2008), and as we argue, as well in the regulation
around, to interoceptive sensations, like feeling the internal of emotional states.
sensations of the body. Other strategies focus on the broad multi- In this context, one of the problematic aspects of Gross’s
modal sensory perception of the body, in which interoceptive, “process model” of emotion regulation is the assumption of a
exteroceptive sensations, and basic sensory (auditory and visual) linear fixed sequence through which emotions are generated,
perceptions are used as a whole as the main focus of intentional starting from attention to relevant external stimuli, cognitive
experiential explorations (Kabat-Zinn, 2005). appraisals, to emotional responses and behaviors as secondarily
From the above, the bottom-up mindfulness-based emotion generated (Koole, 2009). Nevertheless, relevant stimuli can
regulation strategies range from the titrated exposure to negative trigger emotions without cognitive reappraisal (e.g., Neumann
sensations (e.g., physical pain), to different body and perception et al., 2003) and emotions can be generated from the bottom-
modalities conscious explorations, to the exposure to the up systems (Ochsner et al., 2009; McRae et al., 2012). Using
complete range of negative and positive emotions without magneto-encephalography Rudrauf et al. showed that emotional
holding or avoiding/rejecting, which are thought within MBIs stimuli elicited early brain activation in the visual cortex,
and EMs trainings. In sum, there is an explicit intention of spreading through the ventral visual stream, temporopolar
experiential exploration of bodily sensations (e.g., the felt sense) regions, to OFC/vMPFC, ACC, and SSC. This early activation
underlying all type of emotion and mental content (Hölzel et al., was correlated to arousal ratings and heart beats changes
2011a). For example in the MBCT program, participants are (Rudrauf et al., 2009). Also, it is known that bodily movements
instructed to use the “opening the door of the body” strategy, can actively influence emotions (Strack et al., 1988; Niedenthal
which invites to be aware of the body sensations that accompany et al., 2005), the manipulation of body posture can alter the
any intense emotions, stepping back from cognitive analysis and regulation of mood (Veenstra et al., 2016), and intentional
rumination and thus cultivating “intimacy” with the raw and movement can regulate emotional states (Shafir et al., 2013).
usually rejected experience of emotions (Segal et al., 2002). As we From this, even more relevant is the fact that previous emotional
have argued, these strategies are primarily the result of changes in states can strongly influence cognitions and attention processes
bottom-up emotion regulation systems (e.g., exposure to painful (Okon-Singer et al., 2015), which then will drive the emotion
feelings), and can be present in mindfulness inductions, MBIs regulation process. We argue that this model is fairly reductionist
and EMs. (neurocentric), since it denies the constitutive interwoven nature
We further noted that studies applying cognitive reappraisal of body and brain and that their widely known continuous bi-
to emotions generated via bottom-up stimulation can result in a directional interactions are essential for adaptive behavior (Chiel
paradoxical increase in amygdala reactivity (Herwig et al., 2010; and Beer, 1997).
McRae et al., 2012), which in turn can be related to ruminative We argue that the cognitivist “neurocentric” model also
or repetitive negative thinking as maladaptive cognitive emotion disregard the complex reciprocal influences between cortical
regulation strategies (see Table 3), characteristic of anxiety and (high-order) and subcortical (low-order) regions (Okon-Singer
depression disorders (Aldao et al., 2010). Dysfunctional top- et al., 2015). This “corticocentric” model of the brain, in which
down emotion regulation in psychiatric conditions such as MDD “high”-order regions dominate “low”-order regions (Parvizi,
(Johnstone et al., 2007) might be related to dysfunctional forms 2009), fits very well with the “process model” of emotion
of self-evaluative processes such as rumination and worry (Farb regulation, in which only the cortical top-down emotion
and Segal, 2012). In this sense, emotions can be generated from regulation system has a privileged role for regulating emotional
top-down and bottom-up systems (Ochsner et al., 2009; McRae states. As we have shown in this article, bottom-up (mindfulness-
et al., 2012), and the way/pathway emotions are generated seems based) emotion regulation strategies modulate sensory-perception
to play a crucial role in the successfulness of emotion regulation and interoceptive-proprioception components of the emotional
strategies. Bottom-up-generated emotional states, as pain and state, due to changes in bottom-up emotion regulation systems.
reward in EM studies reveal, might be best targeted by bottom-up These subcortical systems are central in the homeostatic
mindfulness emotion regulation strategies (see Table 4). regulation of neuro-vegetative and visceral functions which
provide the bodily aspect of emotion experience (Bechara et al.,
2000; Critchley et al., 2002).
Embodied Emotions and Emotion The enactive approach to mind-brain considers cognition,
Regulation emotion, and body functions as parts of an integrated system at
Classical theories of emotions from Aristoteles, Spinoza, and neurobiological, psychological, and phenomenological domains
Hume have highlighted the importance of the body and (Thompson and Stapleton, 2009). One of its central principles
physiological aspects of emotions, conceiving them essentially is the notion of embodiment, or embodied cognition, which
as psychosomatics states (Colombetti and Thompson, 2008). in simple terms claims that the whole body (not only the
Post Jamesian contemporary authors like Damasio and Prinz brain) is involved in building up cognition (Varela et al., 1991;
assert that emotions are basically the perception of the actual Kiverstein, 2012), and in this particular case the experience
physiological condition, affirming in a broad sense the embodied of emotions (Colombetti and Thompson, 2008; Slaby et al.,

2013; Colombetti, 2014). From this perspective, the emotional or states and regulatory mechanisms as inseparable, relying in
affective dimension is connatural and constitutive of organism’s shared neural networks. It offers a preliminary new framework
adaptation and agency in the world. Organisms have to be for integrating neurobiological, psychophysiological, and
“sensible” to their environment in order to make sense and psychological systems perspectives on emotion regulation
adaptively respond to new demands, in this account emotions are and clinical interventions. It aims to be a multilevel and
inseparable from cognitions (Colombetti, 2014). Central for the non-reductive paradigm to advance the understanding of
affective constitution of organisms, three interrelated activities emotion dysregulation psychopathologies and their changes
characterize the embedded body-brain system: the capacity of in the context of various biological and psychological
self-regulation of internal states, sensorimotor coupling with the treatments.
environment and intersubjective interaction with other agents
(Thompson and Varela, 2001).
In this context, we argue that emotions are the ensuing and CLINICAL IMPLICATIONS: EMOTION
guiding state of the organism engagement with the environment REGULATION, MINDFULNESS, AND
(world), in which the regulation of its own internal homeostatic PSYCHOTHERAPY
states (humoral, visceral, somatic-motor) is inseparable from
the emotional state itself (that is targeted with the regulation). As we have seen, MBIs have shown efficacy in a myriad of
As an example, we cannot think that body temperature (the psychological disorders, characterized by emotion dysregulation
target of the regulation) is something separate and distinct psychopathology (see Table 1). From the perspective of
from the homeostatic mechanisms that continuously regulate longitudinal, clinical, and affective neuroscience studies, we
body functions to keep the temperature constant (regulation hypothesize that changes in bottom-up emotion regulation
mechanism). In fact, the actual body temperature emerges as systems might be a key differential feature of MIBs vs.
the result of the reciprocal interactions of diverse regulatory the usual Western psychotherapeutic approaches—more
mechanisms. Derived from this, we propose a preliminary specifically, not in the sense that only MBIs elicit changes
account of emotion regulation as an embodied process, basically in these systems (which is not the case), but in the sense
rejecting the dualism between emotional states (and its somatic that MBIs explicitly involve the engagement of bottom-up
expressions, motor and autonomic systems), and the processes mindfulness emotion regulation strategies, using the sensory and
and mechanisms of emotion regulation. Emotions and its interoceptive components of emotions as targets and vehicles for
experience are the result of the continuous reciprocal interactions emotion regulation (according to embodied emotion regulation
of top-down, bottom-up, sensory-perception and interoception account).
processes, in which top-down and bottom-up systems can serve From a clinical psychotherapeutic perspective, this means
as generative and regulatory mechanisms. As we have reviewed that the therapist (or MBI instructor) will be able to guide the
in this paper, both emotion systems participate in the generation patient/client into the application of different top-down and
and expression of emotional states (Ochsner et al., 2009; McRae bottom-up mindfulness based strategies. In the case of bottom-
et al., 2012), at the same time, both are engaged in the regulation up strategies, the clinician encourages the participants to focus
of internal homeostatic states (humoral, visceral) and expressive on the “bodily” components of different emotional state, always
somatic-motor responses (Frank et al., 2014; Kohn et al., 2014; conveying the attitudinal stance of acceptance and openness.
Etkin et al., 2015). In this way, discouraging the intend to control and subjugate
The embodied approach to emotion regulation regarding the negative emotional states, but more importantly, discouraging
problem of “mindful emotion regulation” allows us to conceive the use of maladaptive top-down emotion regulation strategies
top-down and bottom-up mindfulness based strategies in a like avoidance, rumination, and suppression among others.
dimensional and continuous way. These strategies primarily In this sense, there is a constant incentive to shift from a
target different aspects of the emotional state, cognitions self-narrative perspective (ruminative), based on past or future
and thought process, sensory-perception, and interoceptive- stories, to a self-experiential present-centered perspective, so the
proprioception, and their corresponding neural substrates, in experience of emotion is decoupled from maladaptive evaluative
this way, at the same time regulating and ensuing the current cognitions. As stated by Chambers, one main difference between
emotional state. From this, it is possible to understand that psychotherapeutic interventions like psychoanalysis and CBT,
even mindfulness induction and MBIs can deploy bottom-up and MBIs, is that the former aim to change the content of
regulation strategies, and also EMs can use top-down emotion emotional states (self-narratives and cognitions), while MBIs
regulation strategies as part of their repertoire. At the same time, focus on changing the relationship (and not the content) with the
different mindfulness related practices (as samatha, vipashyana emotional (painful) states (Chambers et al., 2009); changing the
and compassion, etc.), as taught within MBIs and EMs trainings perspective from which it is experienced, encouraging acceptance
might differentially engage the components of the emotional state and curiosity about the experience itself (self-experiential focus).
(Dahl et al., 2015). From an emotional learning perspective, this process can be seen
In sum, our approach to emotions and emotion regulation as an exercise of exposure (to certain emotions or experiences),
intends to overcome the “neurocentrism” and “corticocentrism” extinction of maladaptive cognitions or reactive responses, and
of current cognitivist model of emotion regulation. Our reconsolidation as a new relationship pattern regarding own
embodied account of emotion regulation considers emotional experiences or daily life problems (Hölzel et al., 2011a).

Mindfulness and Mentalization in the mentalization interventions, in which the focus is to explore,
Context of Psychotherapy cognitively understand and change mental contents, which
Mindfulness and mentalization can be conceived as different may be referred to future or past life events, but also to
heuristics and approaches to understand mental health, clinical emotions and dysregulated emotional states (Allen, 2006). In
interventions, and psychopathological developments. The notion line with this, mentalization as an emotion regulation strategy
of mentalization has a heterogeneous origin, starting from has been considered a top-down strategy, relying in the
the construct of theory of mind developed in the field of explicit emotion regulation and in the theory of mind brain
etiology/cognitive science (Premack and Wooddruff, 1978), the systems (Fonagy and Luyten, 2009; Vrticka and Vuilleumier,
concept of symbolization from psychoanalysis (Choi-Kain and 2012). As we have stated, MBIs engages bottom-up emotion
Gunderson, 2008) and the notion of meta-cognition from novel regulation strategies, which constitutes the distinctive ingredient
developments in the empirical study of attachment (Main, 1991). from other forms of psychotherapies. From our perspective,
In clinical terms mentalization is defined as the capacity to mindfulness and mentalization have common and different
understand one’s own actions and those of others in terms psychobiological functions, which are complementary in the
of intentional mental states like desires, needs, and feelings context of treatments for diverse psychopathologies related to
(Choi-Kain and Gunderson, 2008). According to psychodynamic emotion dysregulation and mentalization deficits. Nevertheless,
theories, mentalization is a developmental capacity that depends further research needs to be done with a view to achieving a better
on the quality of the early mother–infant relationship, the understanding of the biological and psychological differences
development of secure attachment in the infant and a mother’s between these constructs, as well as integrating them properly in
capacities for mentalization (Fonagy et al., 2002). Originally psychotherapeutic treatments.
developed to understand BPD psychopathology, actually its
deficit has been implicated in a wide range of conditions
including autism and schizophrenia, among others (Roffman CONCLUSIONS AND FUTURE
et al., 2012). Enhancing mentalization is viewed as a common DIRECTIONS
factor responsible for psychotherapeutic change processes, not
only in psychodynamic approaches, but also in other clinical Over the last few years, research on contemplative and affective
perspectives (Björgvinsson and Hart, 2006 for CBT; Lewis, 2006 sciences has grown considerably. In this article we have
for DBT). Moreover, in patients with BPD, increased capacity for shown how mindfulness is related to emotion regulation using
mentalization is considered the central mechanism of change in different theoretically and empirically derived models. The main
all effective treatments (Fonagy and Bateman, 2006). hypothesis explored is that emotion regulation changes are a
Exploring the common ground between mindfulness core mechanism underlying the salutary effects of mindfulness
and mentalization, Goodman (2014) uses four aspects of and MBIs. Nevertheless, many of the psychological and
mentalization: (1) observing mental phenomena, (2) describing neurocognitive theoretical models of mindfulness’s mechanisms
or labeling mental phenomena, (3) describing the meaning and are not properly and empirically validated. At the same time,
motivation of one’s own and others’ behavior as the product empirical studies face many methodological limitations as well.
of mental states, and (4) understanding the intrinsic linkage One important problem is the notion of mindfulness itself.
and mutual influence of mental states in oneself and others. As was mentioned, it has been used for referring to a wide
Taking into account Baer et al.’s models (see Table 2), Goodman range of psychological phenomena, like a trait (or dispositional
suggests that mentalization and mindfulness overlap in two key mindfulness), a proper meditation practice or a mental state
areas: observing mental phenomena, and labeling/describing (Davidson, 2010). Even the concept of mindfulness lacks a unique
mental phenomena. From the perspective of emotion regulation operationalization, since many authors have proposed different
systems, both mental processes correspond to top-down definitions, understanding it as an attention capacity, an attitude,
emotion regulation strategies, such as metacognitive awareness a characteristic type of awareness, or even a combination of
and affect labeling. However, the capacity for attributing these (Quaglia et al., 2015). As Grossman states, the complexity
intentionality to mental states and for understanding the of the concept seems more related to a lack of consensus
interpersonal influences of mental states, are distinctive factors between experts, among other critical issues that constructors of
of mentalization (Goodman, 2014). Given the interpersonal inventories might disregard (Grossman, 2008).
nature of psychotherapy, mentalization capacities constitute On one side, studies measuring dispositional mindfulness
central skills for the therapist (to work with patients) and for using self-report scales have demonstrated good reliability and
the patients (to be developed within the treatment; Fonagy and convergent validity (Quaglia et al., 2015) and a preliminary
Bateman, 2006). coherent putative neural correlate (see Table 4). Coffey et al.
Another important difference between mindfulness and have demonstrated that mindfulness and emotion regulation
mentalization, is the type of relationship intended with mental correspond to related but different constructs (Coffey et al.,
contents and temporality of life events. As we stated, MBIs don’t 2010). Nevertheless, the construct of dispositional mindfulness
intend to change mental contents, neither explore life events from entails several problematic aspects, starting from the assumption
the past or future possibilities, its main focus is the present- that self-report mindfulness scales (basically the self-perception
centered non-evaluative awareness of the self-experience. Unlike of a person) actually tap into the proper practices of mindfulness

(Grossman, 2011). For instance, the specificity of the instruments cognitive and emotion processes within homeostatic regulatory
to MBIs is unknown, e.g., other interventions not based on mechanisms, as well as a non-hierarchical view for conceiving
mindfulness might change the mindfulness level (Lutz et al., cortical and subcortical systems, as well as brain and body
2015). Finally, using these instruments in the context of MBIs interactions. Further developments might complement this
might induce biased responses because of the verbal exposure framework integrating first-person phenomenological accounts
to the word and concept of mindfulness itself, and not because of emotions and emotion regulation, looking for further integrate
of any actual acquired capacity (Van Dam et al., 2012). Another experiential and subjective reports with psychophysiological
problematic issue with dispositional mindfulness is the wide and neurobiological measurements (see Colombetti, 2014, for
range of confounders or variables that actually impact the affective neuro-physiophenomenology).
dispositional “mindfulness level,” including other overlapping In line with these recommendations and limitations, from the
and related psychological traits that also vary within normative perspective of methodological and measurement techniques, we
and clinical populations, like: attention and emotional functions, suggest that research on mindfulness and emotion regulation
attitudinal and biased dispositions, prior socialization with the should take advantage of mobile device technologies, for example
construct and experience with related practices (like yoga or using experience sampling methods, or biological measurements
psychotherapy; Quaglia et al., 2015). Future studies will have including mobile EEGs or galvanic response devices, thereby
to control for these factors to better disentangle the nature of increasing the ecological validity of measurements, variables and
dispositional mindfulness as a construct itself. constructs of interest. Serum biological markers of inflammatory
For longitudinal clinical studies, RCTs with active control response and neuroplasticity (BDNF, for example) are also of
groups and multi-arm designs seems to be methodologically relevance as putative biological mechanisms of MBIs. As regards
the “gold standard” for unraveling the efficacy and effectiveness neuroimaging technologies, future studies might integrate
of a given therapeutic intervention, either for inferiority or different methods, taking advantage of the specificity of each, for
superiority studies. As in Zeidan et al. (2015), comparing example combining the spatial resolution of MRIs with positron
mindfulness, sham mindfulness, placebo, and control could emission tomography (PET), which might help to disentangle
demonstrate the efficacy of all interventions for pain relief, but differences in neurotransmitters or neuroradiological markers
noting a differential brain mechanism in emotion regulation of neuroinflammation. Within MRI techniques, the use of
of pain (Zeidan et al., 2015). For further understanding the computational modeling might help to build and test more
differential engagement of the emotion regulation systems in precise and sophisticated theoretical models for understanding
MBIs, future neuroimaging longitudinal studies will have to cognitive emotional systems underlying mindfulness and
explicitly compare different mindfulness instructions within the emotion regulation. Finally, multivariate pattern analysis is
experimental manipulations (i.e., top-down—attention based situated at a privileged level for decoding mental states (certain
vs. bottom-up bodily-based). Then they can explore the emotion regulation strategies or mindfulness states) from brain
acquisition and development of the strategies and their neural signatures using trained classifiers.
correlates. For avoiding problematic aspects of self-report scales, Clinical applications of MBIs will require a very good
clinical studies should try to include behavioral outcome understanding of what’s better for whom, and distinguishing
measures of mindfulness. For better understanding putative what types of psychological treatments, regular psychotherapy
mechanisms, longitudinal studies should use several prospective (of different types) or MBIs (of different types) are better
measurements of variables of interest to better disentangle how for different types of depression or anxiety disorder. This
changes in independent variables and mediators affect dependent leads to another question regarding how to combine different
variables (Kazdin, 2009). forms of psychotherapy with MBIs in the context of a wider
Using neurobiologically based emotion regulation systems and more comprehensive model of healthcare, even including
as a framework, we have described how top-down strategies psychopharmacological treatments. A better understanding of
(explicit emotion regulation system) and bottom-up strategies emotion regulation mechanisms underlying mindfulness and
(emotion generation and implicit emotion-regulation systems) psychotherapy, from biological and clinical perspectives, will
can be present within novice and expert meditators. In order foster new insights into emotional life and its disturbances,
to deal with the controversy of emotion regulation mechanisms with the purpose of refining and developing better therapeutic
underlying mindfulness in MBIs and EMs, we have proposed interventions for the widespread mental health disorders
the distinction between mindfulness-based top-down emotion characterized by emotion dysregulation.
regulation strategies based on attention and acceptance, vs.
mindfulness-based bottom-up strategies, which target bodily
representations of emotional states. We proposed an embodied AUTHOR CONTRIBUTIONS
perspective on emotion regulation as a preliminary framework
as a means for understanding different emotion regulation SG: conceive the original idea of the article, decided the design
systems, rejecting the dualism between somatic emotional states of each part. Completed the revision of all scientific literature.
and the processes and mechanisms of emotion regulation. Performed all draft versions of the document, as well as the final
From this, the experience of emotional states is build up (approval) version to be published. SG is totally accountable for
from the continuous reciprocal interactions of regulatory all aspects of the work, and ensures that all different questions
mechanisms. This perspective offers an integrative view of regarding any part of the work can be appropriately investigated

and resolved. SM: conceive the original idea of the article, decided questions regarding any part of the work can be appropriately
the design of each part. Completed the revision of all scientific investigated.
literature. Performed all draft versions of the document, as well
as offering important intellectual content for the final version. FUNDING
Also, gave the final approval of the version to be published. HR:
conceive the original idea of the article, decided the design of This review paper was supported by the Fund of the following
each part. Completed the revision of most scientific literature. institutions: CONICYT (National Commission for Scientific and
Participated in the elaboration of almost all draft versions of Technological Research, Chile). Beca-Chile Scholarship. Berlin
the document, as well as offered very important intellectual School of Mind and Brain, Humboldt Universität. Fund for
content for the final version. Also, gave the final approval of Innovation and Competitiveness (FIC) of the Chilean Ministry of
the version to be published. HR and SM are totally accountable Economy, Development and Tourism, through the Millennium
for all aspects of the work, and ensures that all different Scientific Initiative, Grant N◦ IS130005.
REFERENCES Brefczynski-Lewis, J. A, Lutz, A, Schaefer, H. S., Levinson, D. B., and

Davidson, R. J. (2007). Neural correlates of attentional expertise in long-
Abravanel, B. T., and Sinha, R. (2015). Emotion dysregulation term meditation practitioners. Proc. Natl. Acad. Sci. U.S.A. 104, 11483–11488.
mediates the relationship between lifetime cumulative adversity doi: 10.1073/pnas.0606552104
and depressive symptomatology. J. Psychiatr. Res. 61, 89–96. Brown, K. W., Goodman, R. J., and Inzlicht, M. (2013). Dispositional mindfulness
doi: 10.1016/j.jpsychires.2014.11.012 and the attenuation of neural responses to emotional stimuli. Soc. Cogn. Affect.
Aldao, A., Nolen-Hoeksema, S., and Schweizer, S. (2010). Emotion-regulation Neurosci. 8, 93–99. doi: 10.1093/scan/nss004
strategies across psychopathology: a meta-analytic review. Clin. Psychol. Rev. Buhle, J. T., Silvers, J. A., Wage, T. D., Lopez, R., Onyemekwu, C., Kober, H.,
30, 217–237. doi: 10.1016/j.cpr.2009.11.004 et al. (2014). Cognitive reappraisal of emotion: a meta-analysis of human
Allen, J. G. (2006). “Mentalizing in practice,” in Handbook of Mentalization-Based neuroimaging studies. Cereb. Cortex 24, 2981–2990. doi: 10.1093/cercor/bht154
Treatment, eds J. G. Allen and P. Fonagy (Chichester: John Wiley & Sons, Ltd.), Burklund, L. J., David Creswell, J., Irwin, M. R., and Lieberman, M. D. (2014). The
3–30. common and distinct neural bases of affect labeling and reappraisal in healthy
Allen, M., Dietz, M., Blair, K. S., van Beek, M., Rees, G., Vestergaard-Poulsen, adults. Front. Psychol. 5:221. doi: 10.3389/fpsyg.2014.00221
P., and Roepstorff, A. (2012). Cognitive-affective neural plasticity following Chambers, R., Gullone, E., and Allen, N. B. (2009). Mindful emotion
active-controlled mindfulness intervention. J. Neurosci. 32, 15601–15610. regulation: an integrative review. Clin. Psychol. Rev. 29, 560–572.
doi: 10.1523/JNEUROSCI.2957-12.2012 doi: 10.1016/j.cpr.2009.06.005
Analayo (2003). Satipatthana: The Direct Path to Awakening. Birmingham: Chiel, H. J., and Beer, R. D. (1997). The brain has a body: adaptive behavior emerges
Windhorse Publications. from interactions of nervous systems, body and environment. Trends Neurosci.
A-Tjak, J. G. L., Davis, M. L., Morina, N., Powers, M. B., Smits, J. A. J., 20, 553–557. doi: 10.1016/S0166-2236(97)01149-1
and Emmelkamp, P. M. G. (2015). A meta-analysis of the efficacy of Chiesa, A., and Serretti, A. (2011). Mindfulness based cognitive therapy for
acceptance and commitment therapy for clinically relevant mental and psychiatric disorders: a systematic review and meta-analysis. Psychiatry Res.
physical health problems. Psychother. Psychosom. 84, 30–36. doi: 10.1159/0003 187, 441–453. doi: 10.1016/j.psychres.2010.08.011
65764 Chiesa, A., Serretti, A., and Christian Jakobsenc, J. (2013). Mindfulness: top–
Baer, R. A., Smith, G. T., Hopkins, J., Krietemeyer, J., and Toney, L. (2006). Using down or bottom–up emotion regulation strategy? Clin. Psychol. Rev. 33, 82–96.
self-report assessment methods to explore facets of mindfulness. Assessment 13, doi: 10.1016/j.cpr.2012.10.006
27–45. doi: 10.1177/1073191105283504 Choi-Kain, L. W., and Gunderson, J. G. (2008). Mentalization: ontogeny,
Bao, X., Xue, S., and Kong, F. (2015). Dispositional mindfulness and perceived assessment, and application in the treatment of borderline personality disorder.
stress: the role of emotional intelligence. Pers. Individ. Dif. 78, 48–52. Am. J. Psychiatry 165, 1127–1135. doi: 10.1176/appi.ajp.2008.07081360
doi: 10.1016/j.paid.2015.01.007 Clarke, K., Mayo-Wilson, E., Kenny, J., and Pilling, S. (2015). Can non-
Bechara, A., Damasio, H., and Damasio, A. R. (2000). Emotion, decision pharmacological interventions prevent relapse in adults who have recovered
making and the orbitofrontal cortex. Cereb. Cortex 10, 295–307. from depression? A systematic review and meta-analysis of randomised
doi: 10.1093/cercor/10.3.295 controlled trials. Clin. Psychol. Rev. 39, 58–70. doi: 10.1016/j.cpr.2015.04.002
Berenbaum, H., Raghavan, C., Le, H. N., Vernon, L. L., and Gomez, J. J. (2003). Coffey, K. A., Hartman, M., and Fredrickson, B. L. (2010). Deconstructing
A taxonomy of emotional disturbances. Clin. Psychol. Sci. Pract. 10, 206–226. mindfulness and constructing mental health: understanding
doi: 10.1093/clipsy.bpg011 mindfulness and its mechanisms of action. Mindfulness 1, 235–253.
Björgvinsson, T. and Hart, J. (2006). “Cognitive behavioral therapy promotes doi: 10.1007/s12671-010-0033-2
mentalizing,” in Handbook of Mentalization-Based Treatment, eds J. G. Allen Colombetti, G. (2014). The Feeling Body: Affective Science Meets the Enactive Mind.
and P. Fonagy (Chichester: John Wiley & Sons, Ltd.), 157–170. Cambridge, MA: MIT Press.
Bohlmeijer, E., Prenger, R., Taal, E., and Cuijpers, P. (2010). The effects of Colombetti, G., and Thompson, E. (2008). “The feeling body: towards an enactive
mindfulness-based stress reduction therapy on mental health of adults with approach to emotion,” in Developmental Perspectives on Embodiment and
a chronic medical disease: a meta-analysis. J. Psychosom. Res. 68, 539–544. Consciousness, eds W. F. Overton, U. Müller, and J. L. Newman (New York,
doi: 10.1016/j.jpsychores.2009.10.005 NY: Erlbaum), 45–68.
Bowen, S., Chawla, N., Marlatt, G. A. (2010). Mindfulness-Based Relapse Corrigan, F. M., Fisher, J. J., and Nutt, D. J. (2011). Autonomic dysregulation and
Prevention for Addictive Behaviors: a Clinician’s Guide. New York, NY: Guilford the Window of Tolerance model of the effects of complex emotional trauma. J.
Press. Psychopharmacol. 25, 17–25. doi: 10.1177/0269881109354930
Bowen, S., K, W., Clifasefi, S. L., Grow, J., Chawla, N., Hsu, S. H., Craig, A. D. (2009). How do you feel–now? The anterior insula and human
et al. (2014). Relative efficacy of mindfulness-based relapse prevention, awareness. Nat. Rev. Neurosci. 10, 59–70. doi: 10.1038/nrn2555
standard relapse prevention, and treatment as usual for substance use Cramer, H., Haller, H., Lauche, R., and Dobos, G. (2012). Mindfulness-based stress
disorders: a randomized clinical trial. JAMA Psychiatry 71, 547–556. reduction for low back pain. A systematic review. BMC Complement. Altern.
doi: 10.1001/jamapsychiatry.2013.4546 Med. 12:162. doi: 10.1186/1472-6882-12-162

Creswell, J. D., Way, B. M., Eisenberger, N. I., and Lieberman, M. D. (2007). Neural Gard, T., Hölzel, B. K., Sack, A. T., Hempel, H., Lazar, S. W., Vaitl, D., et al.
correlates of dispositional mindfulness during affect labeling. Psychosom. Med. (2012). Pain attenuation through mindfulness is associated with decreased
69, 560–565. doi: 10.1097/PSY.0b013e3180f6171f cognitive control and increased sensory processing in the brain. Cereb. Cortex
Critchley, H. D., Melmed, R. N., Featherstone, E., Mathias, C. J., and Dolan, 22, 2692–2702. doi: 10.1093/cercor/bhr352
R. J. (2002). Volitional control of autonomic arousal: a functional magnetic Giluk, T. L. (2009). Mindfulness, big five personality, and affect : a meta-analysis.
resonance study. Neuroimage 16, 909–919. doi: 10.1006/nimg.2002.1147 Pers. Individ. Dif. 47, 805–811. doi: 10.1016/j.paid.2009.06.026
Crow, T., Cross, D., Powers, A., and Bradley, B. (2014). Emotion dysregulation Goldin, P. R., and Gross, J. J. (2010). Effects of mindfulness-based stress reduction
as a mediator between childhood emotional abuse and current depression in (MBSR) on emotion regulation in social anxiety disorder. Emotion 10, 83–91.
a low-income African-American sample. Child Abuse Neglect 38, 1590–1598. doi: 10.1037/a0018441
doi: 10.1016/j.chiabu.2014.05.015 Goldin, P., Ziv, M., Jazaieri, H., and Gross, J. J. (2012). Randomized controlled trial
Dahl, C. J., Lutz, A., and Davidson, R. J. (2015). Reconstructing and deconstructing of mindfulness-based stress reduction versus aerobic exercise: effects on the
the self: cognitive mechanisms in meditation practice. Trends Cogn. Sci. 19, self-referential brain network in social anxiety disorder. Front. Hum. Neurosci.
515–523. doi: 10.1016/j.tics.2015.07.001 6:295. doi: 10.3389/fnhum.2012.00295
Damasio, A. R. (1999). The Feeling of What Happens: Body and Emotion in the Goldin, P., Ziv, M., Jazaieri, H., Hahn, K., and Gross, J. J. (2013). MBSR vs aerobic
Making of Consciousness. New York, NY: Harcourt Brace. exercise in social anxiety: fMRI of emotion regulation of negative self-beliefs.
Davidson, R. J. (2010). Empirical explorations of mindfulness: conceptual and Soc. Cogn. Affect. Neurosci. 8, 65–72. doi: 10.1093/scan/nss054
methodological conundrums. Emotion 10, 8–11. doi: 10.1037/a0018480 Goodman, G. (2014). “Mentalization: an interpersonal approach to mindfulness,”
Desbordes, G., Negi, L. T., Pace, T. W. W., Wallace, B. A., Raison, C. L., in Mindfulness, Acceptance, and the Psychodynamic Evolution: Bringing Values
and Schwartz, E. L. (2012). Effects of mindful-attention and compassion into Treatment Planning and Enhancing Psychodynamic Work with Buddhist
meditation training on amygdala response to emotional stimuli in an ordinary, Psychology, ed J. M. Stewart (Oakland, CA: Context Press), 111–132.
non-meditative state. Front. Hum. Neurosci. 6:292. doi: 10.3389/fnhum.2012. Gotink, R. A., Chu, P., Busschbach, J. J. V., Benson, H., Fricchione, G. L., and
00292 Hunink, M. G. M. (2015). Standardised mindfulness-based interventions in
Dörfel, D., Lamke, J. P., Hummel, F., Wagner, U., Erk, S., and Walter, healthcare: an overview of systematic reviews and meta-analyses of RCTs. PLoS
H. (2014). Common and differential neural networks of emotion ONE 10:e0124344. doi: 10.1371/journal.pone.0124344
regulation by detachment, reinterpretation, distraction, and expressive Grabovac, A. D., Lau, M. A., and Willett, B. R. (2011). Mechanisms of
suppression: a comparative fMRI investigation. Neuroimage 101, 298–309. mindfulness: a buddhist psychological model. Mindfulness 2, 154–166.
doi: 10.1016/j.neuroimage.2014.06.051 doi: 10.1007/s12671-011-0054-5
Eberth, J., and Sedlmeier, P. (2012). The effects of mindfulness meditation: a Grant, J. A., Courtemanche, J., and Rainville, P. (2011). A non-elaborative
meta-analysis. Mindfulness 3, 174–189. doi: 10.1007/s12671-012-0101-x mental stance and decoupling of executive and pain-related cortices
Etkin, A., Büchel, C., and Gross, J. J. (2015). The neural bases of predicts low pain sensitivity in Zen meditators. Pain 152, 150–156.
emotion regulation. Nat. Rev. Neurosci. 16, 693–700. doi: 10.1038/ doi: 10.1016/j.pain.2010.10.006
nrn4044 Grant, J. A., Duerden, E. G., Courtemanche, J., Cherkasova, M., Duncan, G. H.,
Farb, N. S., Anderson, A. K., Mayberg, H., Bean, J., and McKeon, D. (2010). and Rainville, P. (2013). Cortical thickness, mental absorption and meditative
Minding one’s emotions : mindfulness training alters the neural expression of practice: possible implications for disorders of attention. Biol. Psychol. 92,
sadness. Emotions 10, 25–33. doi: 10.1037/a0017151 275–281. doi: 10.1016/j.biopsycho.2012.09.007
Farb, N. A., and Segal, Z. V. (2012). The mindful brain and emotion Gratz, K. L., and Gunderson, J. G. (2006). Preliminary data on an acceptance-
regulation in mood disorders. Can. J. Psychiatry 57, 70–77. based emotion regulation group intervention for deliberate self-harm among
doi: 10.1177/070674371205700203 women with borderline personality disorder. Behav. Ther. 37, 25–35.
Farb, N. A., Segal, Z. V., Mayberg, H., Bean, J., McKeon, D., Fatima, Z., et al. doi: 10.1016/j.beth.2005.03.002
(2007). Attending to the present: mindfulness meditation reveals distinct Grecucci, A., Pappaianni, E., Siugzdaite, R., Theuninck, A., and Job, R. (2015a).
neural modes of self-reference. Soc. Cogn. Affect. Neurosci. 2, 313–322. Mindful emotion regulation: exploring the neurocognitive mechanisms behind
doi: 10.1093/scan/nsm030 mindfulness. BioMed. Res. Int. 2015:670724. doi: 10.1155/2015/670724
Feltman, R., Robinson, M. D., and Ode, S. (2009). Mindfulness as a moderator Grecucci, A., De Pisapia, N. Kusalagnana Thero, D., Paladino, P. M., Venuti, P.,
of neuroticism-outcome relations: a self-regulation perspective. J. Res. Pers. 43, and Job, R. (2015b). Baseline and strategic effects behind mindful emotion
953–961. doi: 10.1016/j.jrp.2009.08.009 regulation: behavioral and physiological investigation. PLoS ONE 10:e0116541.
Fjorback, L. O., Arendt, M., Ornbol, E., Fink, P., and Walach, H. (2011). doi: 10.1371/journal.pone.0116541
Mindfulness-based stress reduction and mindfulness-based cognitive therapy - Gross, J. J. (1998). The emerging field of emotion regulation: an integrative review.
a systematic review of randomized controlled trials. Acta Psychiatr. Scand. 124, Rev. Gen. Psychol. 2, 271–299. doi: 10.1037/1089-2680.2.3.271
102–119. doi: 10.1111/j.1600-0447.2011.01704.x Gross, J. J. (2001). Emotion regulation in adulthood: timing is everything. Curr.
Fonagy, P., and Bateman, A. W. (2006). Mechanisms of change in mentalization- Dir. Psychol. Sci. 10, 214–219. doi: 10.1111/1467-8721.00152
based treatment of BPD. J. Clin. Psychol. 62, 411–430. doi: 10.1002/jclp. Gross, J. J., and John, O. P. (2003). Individual differences in two emotion regulation
20241 processes: implications for affect, relationships, and well-being. J. Pers. Soc.
Fonagy, P., and Luyten, P. (2009). A developmental, mentalization-based Psychol. 85, 348–362. doi: 10.1037/0022-3514.85.2.348
approach to the understanding and treatment of borderline personality Grossman, P. (2008). On measuring mindfulness in psychosomatic
disorder. Dev. Psychopathol. 21, 1355–1381. doi: 10.1017/S095457940 and psychological research. J. Psychosom. Res. 64, 405–408.
9990198 doi: 10.1016/j.jpsychores.2008.02.001
Fonagy, P., Target, M., Gergely, G., Jurist, E (2002). Affect Regulation, Mentalization Grossman, P. (2011). Defining mindfulness by how poorly I think I pay attention
and the Development of Self. New York, NY: Other Press. during everyday awareness and other intractable problems for psychology’s
Fox, K. C. R., Nijeboer, S., Dixon, M. L., Floman, J. L., Ellamil, M., Rumak, (re)invention of mindfulness: comment on Brown et al. (2011). Psychol. Assess.
S. P., Sedlmeierc, P., et al. (2014). Is meditation associated with altered 23, 1034–1040. doi: 10.1037/a0022713
brain structure? A systematic review and meta-analysis of morphometric Gu, J., Strauss, C., Bond, R., and Cavanagh, K. (2015). How do mindfulness-
neuroimaging in meditation practitioners. Neurosci. Biobehav. Rev. 43, 48–73. based cognitive therapy and mindfulness-based stress reduction improve
doi: 10.1016/j.neubiorev.2014.03.016 mental health and wellbeing ? A systematic review and meta-analysis of
Frank, D. W., Dewitt, M., Hudgens-Haney, M., Schaeffer, D. J., Ball, B. H., mediation studies. Clin. Psychol. Rev. 37, 1–12. doi: 10.1016/j.cpr.2015.
Schwarz, N. F., et al. (2014). Emotion regulation: quantitative meta-analysis of 01.006
functional activation and deactivation. Neurosci. Biobehav. Rev. 45, 202–211. Gu, X., Hof, P. R., Friston, K. J., and Fan, J. (2013). Anterior insular cortex and
doi: 10.1016/j.neubiorev.2014.06.010 emotional awareness. J. Comp. Neurol. 521, 3371–3388. doi: 10.1002/cne.23368

Gyurak, A., Gross, J., and Etkin, A. (2011). Explicit and implicit emotion Lauche, R., Cramer, H., Dobos, G., Langhorst, J., and Schmidt, S. (2013).
regulation: a dual process framework. Cogn. Emot. 25, 400–412. A systematic review and meta-analysis of mindfulness-based stress
doi: 10.1080/02699931.2010.544160 reduction for the fibromyalgia syndrome. J. Psychosom. Res. 75, 500–510.
Hayes, S. C. Strosahl, K. D. Wilson, K. G. (1999). Acceptance and Commitment doi: 10.1016/j.jpsychores.2013.10.010
Therapy: an Experimental Approach to Behavior Change. New York, NY: Lavender, J. M., Wonderlich, S. A., Engel, S. G., Gordon, K. H., Kaye, W. H.,
Guilford Press. and Mitchell, J. E. (2015). Dimensions of emotion dysregulation in anorexia
Herwig, U., Kaffenberger, T., Jäncke, L., and Brühl, A. B. (2010). Self- nervosa and bulimia nervosa: a conceptual review of the empirical literature.
related awareness and emotion regulation. Neuroimage 50, 734–741. Clin. Psychol. Rev. 40, 111–122. doi: 10.1016/j.cpr.2015.05.010
doi: 10.1016/j.neuroimage.2009.12.089 Lazar, S. W., Kerr, C. E., Wasserman, R. H., Gray, J. R., Greve, D. N., Dusek, A., et al.
Hölzel, B. K., Ott, U., Gard, T., Hempel, H., Weygandt, M., Morgen, K., et al. (2005). Meditation experience is associated with increased cortical thickness,
(2008). Investigation of mindfulness meditation practitioners with voxel-based NeuroReport 16, 1893–1897. doi: 10.1097/01.wnr.0000186598.66243.19
morphometry. Soc. Cogn. Affect. Neurosci. 3, 55–61. doi: 10.1093/scan/nsm038 Ledesma, D., and Kumano, H. (2009). Mindfulness-based stress reduction and
Hölzel, B. K., Carmody, J., Evans, K. C., Hoge, E. A., Dusek, J. A., Morgan, L., et al. cancer: a meta-analysis. Psychooncology 18, 571–579. doi: 10.1002/pon.1400
(2009). Stress reduction correlates with structural changes in the amygdala. Soc. Lewis, L. (2006). “Enhancing mentalizing capacity through dialectical behavior
Cogn. Affect. Neurosci. 5, 11–17. doi: 10.1093/scan/nsp034 therapy skills training and positive psychology,” in Handbook of Mentalization-
Hölzel, B. K., Carmody, J., Vangel, M., Congleton, C., Yerramsetti, S. Based Treatment, eds J. G. Allen and P. Fonagy (Chichester: John Wiley & Sons,
M., Gard, T., et al. (2011b). Mindfulness practice leads to increases in Ltd.), 171–182.
regional brain gray matter density. Psychiatry Res. Neuroimaging 191, 36–43. Linehan, M. M. (1993). Cognitive–Behavioral Treatment of Borderline Personality
doi: 10.1016/j.pscychresns.2010.08.006 Disorder. New York, NY: Guilford Press.
Hölzel, B. K., Hoge, E. A., Greve, D. N., Gard, T., Creswell, J. D., Brown, K. W., Luders, E., Cherbuin, N., and Gaser, C. (2016). Estimating brain
et al. (2013). Neural mechanisms of symptom improvements in generalized age using high-resolution pattern recognition: younger brains in
anxiety disorder following mindfulness training. Neuroimage Clin. 2, 448–458. long-term meditation practitioners. Neuroimage. 134, 508–513.
doi: 10.1016/j.nicl.2013.03.011 doi: 10.1016/j.neuroimage.2016.04.007
Hölzel, B. K., Lazar, S. W., Gard, T., Schuman-Olivier, Z., Vago, D. R., and Ott, Luders, E., Phillips, O. R., Clark, K., Kurth, F., Toga, A. W., and Narr, K. L. (2012).
U. (2011a). How does mindfulness meditation work? Proposing mechanisms Bridging the hemispheres in meditation: thicker callosal regions and enhanced
of action from a conceptual and neural perspective. Perspect. Psychol. Sci. 6, fractional anisotropy (FA) in long-term practitioners. Neuroimage 61, 181–187.
537–559. doi: 10.1177/1745691611419671 doi: 10.1016/j.neuroimage.2012.02.026
Jain, F. A., Walsh, R. N., Eisendrath, S. J., Christensen, S., and Rael Cahn, B. (2015). Luders, E., Toga, A. W., Lepore, N., and Gaser, C. (2009). NeuroImage
Critical analysis of the efficacy of meditation therapies for acute and subacute The underlying anatomical correlates of long-term meditation : larger
phase treatment of depressive disorders: a systematic review. Psychosomatics 56, hippocampal and frontal volumes of gray matter. Neuroimage 45, 672–678.
140–152. doi: 10.1016/j.psym.2014.10.007 doi: 10.1016/j.neuroimage.2008.12.061
Johnstone, T., van Reekum, C. M., Urry, H. L., Kalin, N. H., and Davidson, Lutz, A., Brefczynski-Lewis, J., Johnstone, T., and Davidson, R. J. (2008a).
R. J. (2007). Failure to regulate: counterproductive recruitment of top-down Regulation of the neural circuitry of emotion by compassion meditation: effects
prefrontal-subcortical circuitry in major depression. J. Neurosci. 27, 8877–8884. of meditative expertise. PLoS ONE 3:e1897. doi: 10.1371/journal.pone.0001897
doi: 10.1523/JNEUROSCI.2063-07.2007 Lutz, A., Dunne, J. P., and Davidson, J. R. (2008b). “Meditation and the
Kabat-Zinn, J. (2005). Full Catastrophe Living: Using the Wisdom of your Body and neuroscience of consciousness: an introduction,” in Cambridge Handbook of
Mind to Face Stress, Pain, and Illness Fifteenth Anniversary Edition. New York, Consciousness, eds M. M. P. D. Zelazo and E. Thompson (New York, NY:
NY: Bantam Dell. Cambridge University Press), 499–551.
Kang, D.-H., Jo, H. J., Jung, W. H., Kim, S. H., Jung, Y.-H., Choi, C.-H., Lutz, A., Jha, A., Dunne, J. D., and Saron, C. D. (2015). Investigating
et al. (2013). The effect of meditation on brain structure: cortical thickness the phenomenological matrix of mindfulness-related practices from a
mapping and diffusion tensor imaging. Soc. Cogn. Affect. Neurosci. 8, 27–33. neurocognitive perspective. Am. Psychol. 70, 632–658. doi: 10.1037/a0039585
doi: 10.1093/scan/nss056 Lutz, A., McFarlin, D. R., Perlman, D. M., Salomons, T. V., and Davidson,
Kazdin, A. E. (2007). Mediators and mechanisms of change R. J. (2013). Altered anterior insula activation during anticipation and
in psychotherapy. Annu. Rev. Clin. Psychol. 3, 1–27. experience of painful stimuli in expert meditators. Neuroimage 64, 538–546.
doi: 10.1146/annurev.clinpsy.3.022806.091432 doi: 10.1016/j.neuroimage.2012.09.030
Kazdin, A. E. (2009). Understanding how and why psychotherapy leads to change. Lutz, J., Herwig, U., Opialla, S., Hittmeyer, A., Jäncke, L., Rufer, M., et al.
Psychother. Res. 19, 418–428. doi: 10.1080/10503300802448899 (2013). Mindfulness and emotion regulation-an fMRI study. Soc. Cogn. Affect.
Kirk, U., Brown, K. W., and Downar, J. (2015). Adaptive neural reward Neurosci. 9, 776–785. doi: 10.1093/scan/nst043
processing during anticipation and receipt of monetary rewards in mindfulness Main, M. (1991). “Metacognitive knowledge, metacognitive monitoring, and
meditators. Soc. Cogn. Affect. Neurosci. 10, 752–759. doi: 10.1093/scan/nsu112 singular (coherent) vs. multiple (incoherent) models of attachment: some
Kirk, U., Downar, J., and Montague, P. R. (2011). Interoception drives increased findings and some directions for future research,” in Attachment Across the
rational decision-making in meditators playing the ultimatum game. Front. Life Cycle, eds P. Marris, J. Stevenson-Hinde, and C. Parkes (New York, NY:
Neurosci. 5:49. doi: 10.3389/fnins.2011.00049 Routledge), 127–159.
Kirk, U., and Montague, P. R. (2015). Mindfulness meditation modulates Malinowski, P. (2013). Neural mechanisms of attentional control in mindfulness
reward prediction errors in a passive conditioning task. Front. Psychol. 6:90. meditation. Front. Neurosci. 7:8. doi: 10.3389/fnins.2013.00008
doi: 10.3389/fpsyg.2015.00090 McRae, K., Misra, S., Prasad, A. K., Pereira, S. C., and Gross, J. J. (2012). Bottom-
Kiverstein, J. (2012). The meaning of embodiment. Top. Cogn. Sci. 4, 740–758. up and top-down emotion generation: implications for emotion regulation. Soc.
doi: 10.1111/j.1756-8765.2012.01219.x Cogn. Affect. Neurosci. 7, 253–262. doi: 10.1093/scan/nsq103
Kohn, N., Eickhoff, S. B., Scheller, M., Laird, A. R., Fox, P. T., and Mennin, D., and Farach, F. (2007). Emotion and evolving treatments
Habel, U. (2014). Neural network of cognitive emotion regulation - for adult psychopathology. Clin. Psychol. Sci. Pract. 14, 329–352.
An ALE meta-analysis and MACM analysis. Neuroimage 87, 345–355. doi: 10.1111/j.1468-2850.2007.00094.x
doi: 10.1016/j.neuroimage.2013.11.001 Neumann, R., Förster, J., and Strack, F. (2003). “Motor compatibility: the
Kong, F., Wang, X., Song, Y., and Liu, J. (2016). Brain regions involved bidirectional link between behavior and evaluation,” in The Psychology of
in dispositional mindfulness during resting state and their relation with Evaluation: Affective Processes in Cognition and Emotion, eds J. Musch and K.
well-being. Soc. Neurosci. 11, 331–343. doi: 10.1080/17470919.2015.10 C. Klauer (Mahwah, NJ: Lawrence Erlbaum Associates Inc.), 7–49.
92469 Niedenthal, P. M., Barsalou, L. W., Winkielman, P., Krauth-Gruber, S., and Ric,
Koole, S. L. (2009). The psychology of emotion regulation: an integrative review. F. (2005). Embodiment in attitudes, social perception, and emotion. Pers. Soc.
Cogn. Emot. 23, 4–41. doi: 10.1080/02699930802619031 Psychol. Rev. 9, 184–211. doi: 10.1207/s15327957pspr0903_1

Nock, M. K. (2007). Conceptual and design essentials for evaluating Shafir, T., Taylor, S. F., Atkinson, A. P., Langenecker, S. A., and Zubieta,
mechanisms of change. Alcohol. Clin. Exp. Res. 31, 4s–12s. J.-K. (2013). Emotion regulation through execution, observation,
doi: 10.1111/j.1530-0277.2007.00488.x and imagery of emotional movements. Brain Cogn. 82, 219–227.
Ochsner, K. N., Ray, R. R., Hughes, B., McRae, K., Cooper, J. C., Weber, J., doi: 10.1016/j.bandc.2013.03.001
et al. (2009). Bottom-up and top-down processes in emotion generation: Shapiro, S. L., Carlson, L. E., Astin, J. A., and Freedman, B. (2006). Mechanism of
common and distinct neural mechanisms. Psychol. Sci. 20, 1322–1331. mindfulness. J. Clin. Psychol. 62, 373–386. doi: 10.1002/jclp.20237
doi: 10.1111/j.1467-9280.2009.02459.x Shaw, P., Stringaris, A., Nigg, J., and Leibenluft, E. (2014). Emotion dysregulation
Okon-Singer, H., Hendler, T., Pessoa, L., and Shackman, A. (2015). in attention deficit hyperactivity disorder. Am. J. Psychiatry 171, 276–293.
The neurobiology of emotion–cognition interactions: fundamental doi: 10.1176/appi.ajp.2013.13070966
questions and strategies for future research. Front. Hum. Neurosci. 9:58. Shonin, E., Van Gordon, W., and Griffiths, M. D. (2013). Mindfulness-based
doi: 10.3389/fnhum.2015.00058 interventions : towards mindful clinical integration. Front. Psychol. 4:194.
Opialla, S., Lutz, J., Scherpiet, S., Hittmeyer, A., Jäncke, L., Rufer, M., et al. (2014). doi: 10.3389/fpsyg.2013.00194
Neural circuits of emotion regulation: a comparison of mindfulness-based and Singleton, O., Hölzel, B. K., Vangel, M., Brach, N., Carmody, J., and
cognitive reappraisal strategies. Eur. Arch. Psychiatry Clin. Neurosci. 265, 45–55. Lazar, S. W. (2014). Change in brainstem gray matter concentration
doi: 10.1007/s00406-014-0510-z following a mindfulness-based intervention is correlated with
Parent, A., and Hazrati, L.-N. (1995). Functional anatomy of the basal ganglia. improvement in psychological well-being. Front. Hum. Neurosci. 8:33.
I. The cortico-basal ganglia-thalamo-cortical loop. Brain Res. Rev. 20, 91–127. doi: 10.3389/fnhum.2014.00033
doi: 10.1016/0165-0173(94)00007-C Slaby, J., Paskaleva, A., and Stephan, A. (2013). Enactive emotion and impaired
Parvizi, J. (2009). Corticocentric myopia: old bias in new cognitive sciences. Trends agency in depression. J. Conscious. Stud. 20, 33–55. Retrieved from: http://www.
Cogn. Sci. 13, 354–359. doi: 10.1016/j.tics.2009.04.008 ingentaconnect.com/content/imp/jcs/2013/00000020/F0020007/art00003
Piet, J., and Hougaard, E. (2011). The effect of mindfulness-based cognitive Slonim, J., Kienhuis, M., Di Benedetto, M., and Reece, J. (2015). The
therapy for prevention of relapse in recurrent major depressive disorder: relationships among self-care, dispositional mindfulness, and psychological
a systematic review and meta-analysis. Clin. Psychol. Rev. 31, 1032–1040. distress in medical students. Med. Educ. Online 20, 1–13. doi: 10.3402/meo.v20.
doi: 10.1016/j.cpr.2011.05.002 27924
Posner, M. I., Sheese, B. E., Odludaş, Y., and Tang, Y. (2006). Analyzing Stoffers, J. M., Völlm, B. A., Rücker, G., Timmer, A., Huband, N., and
and shaping human attentional networks. Neural Netw. 19, 1422–1429. Lieb, K. (2012). Psychological therapies for people with borderline
doi: 10.1016/j.neunet.2006.08.004 personality disorder. Cochrane Database Syst. Rev. 15:CD005652.
Premack, D. G., Wooddruff, G. (1978). Does the chimpanzee have a doi: 10.1002/14651858.CD005652.pub2
theory of mind? Behav. Brain Sci. 1, 515–526. doi: 10.1017/S0140525X000 Strack, F., Martin, L., and Stepper, S. (1988). Inhibiting and facilitating conditions
76512 of the human smile: a nonobtrusive test of the facial feedback hypothesis. J.
Prinz, J. J. (2004). Gut Reactions: A Perceptual Theory of Emotion. Oxford: Oxford Pers. Soc. Psychol. 54, 768–777. doi: 10.1037/0022-3514.54.5.768
University Press. Tang, Y., Hölzel, B. K., and Posner, M. I. (2015). The neuroscience of mindfulness
Qin, P., and Northoff, G. (2011). How is our self related to midline meditation. Nat. Rev. Neurosci. 16, 213–225. doi: 10.1038/nrn3916
regions and the default-mode network? Neuroimage. 57, 1221–1233. Taylor, V. A., Grant, J., Daneault, V., Scavone, G., Breton, E., Roffe-Vidal, S.,
doi: 10.1016/j.neuroimage.2011.05.028 et al. (2011). Impact of mindfulness on the neural responses to emotional
Quaglia, J., Brown, K., Lindsay, E., Creswell, J., and Goodman, R. (2015). pictures in experienced and beginner meditators. Neuroimage 57, 1524–1533.
“From conceptualization to operationalization of mindfulness,” in Handbook doi: 10.1016/j.neuroimage.2011.06.001
of Mindfulness: Theory, Research, and Practice, eds K. W. Brown, J. D. Creswell, Thompson, E., and Stapleton, M. (2009). Making sense of sense-making:
and R. M. Ryan (New York, NY: Guilford), 151–170. reflections on enactive and extended mind theories. Topoi 28, 23–30.
Raichle, M. E., and Snyder, A. Z. (2007). A default mode of brain doi: 10.1007/s11245-008-9043-2
function: a brief history of an evolving idea. Neuroimage 37, 1083–1090. Thompson, E., and Varela, F. J. (2001). Radical embodiment: neural
doi: 10.1016/j.neuroimage.2007.02.041 dynamics and consciousness. Trends Cogn. Sci. 5, 418–425.
Rapgay, L., and Bystrisky, A. (2009). Classical mindfulness: an introduction to its doi: 10.1016/S1364-6613(00)01750-2
theory and practice for clinical application. Ann. N. Y. Acad. Sci. 1172, 148–162. Thompson, R. A. (1990). “Emotion and self-regulation,” in Socioemotional
doi: 10.1111/j.1749-6632.2009.04405.x development. Nebraska Symposium on Motivation Vol. 36, ed R. A. Thompson
Roffman, J. L., Gerber, A. J., and Flick, D. (2012). “Neural models of (Lincoln: University of Nebraska Press), 367–467.
psychodynamic concepts and treatments: implications for psychodynamic Tomasino, B., Fregona, S., Skrap, M., and Fabbro, F. (2013). Meditation-
psychotherapy,” in Psychodynamic Psychotherapy Research: Evidence-Based related activations are modulated by the practices needed to obtain it and
Practice and Practice-Based Evidence, eds R. A. Levy, J. S. Ablon, and H. Kächele by the expertise: an ALE meta-analysis study. Front. Hum. Neurosci. 6:346.
(New York, NY: Humana Press; Springer), 193–218. doi: 10.3389/fnhum.2012.00346
Rudrauf, D., Lachaux, J. P., Damasio, A., Baillet, S., Hugueville, L., Martinerie, Trungpa, C. (2001). Glimpses of Abhidharma: From a Seminar on Buddhist
J., et al. (2009). Enter feelings: somatosensory responses following early Psychology. Boston, MA: Shambhala Publications.
stages of visual induction of emotion. Int. J. Psychophysiol. 72, 13–23. Vago, D. R., and Silbersweig, D. A. (2012). Self-awareness, self-regulation,
doi: 10.1016/j.ijpsycho.2008.03.015 and self-transcendence (S-ART): a framework for understanding the
Santarnecchi, E., D’Arista, S., Egiziano, E., Gardi, C., Petrosino, R., Vatti, neurobiological mechanisms of mindfulness. Front. Hum. Neurosci. 6:296.
G., et al. (2014). Interaction between neuroanatomical and psychological doi: 10.3389/fnhum.2012.00296
changes after mindfulness-based training. PLoS ONE 9:e108359. Van Dam, N. T., Hobkirk, A. L., Danoff-Burg, S., and Earleywine, M.
doi: 10.1371/journal.pone.0108359 (2012). Mind your words: positive and negative items create method
Schore, A. N. (2003). “Effect of early relational trauma on affect regulation: effects on the five facet mindfulness questionnaire. Assessment 19, 198–204.
the development of borderline and antisocial personality disorders and a doi: 10.1177/1073191112438743
predisposition to violence,” in Affect Dysregulation and Disorders of the Self, ed Van der Velden, A. M., Kuyken, W., Wattar, U., Crane, C., Pallesen, K.
A. N. Schore (New York, NY: W.W. Norton), 266–306. J., Dahlgaard, J., et al. (2015). A systematic review of mechanisms
Segal, Z. V., Williams, M. G., and Teasdale, J. D. (2002). Mindfulness- Based of change in mindfulness-based cognitive therapy in the treatment
Cognitive Behavior Therapy for Depression: A New Approach to Preventing of recurrent major depressive disorder. Clin. Psychol. Rev. 37, 26–39.
Relapse. New York, NY: Guildford. doi: 10.1016/j.cpr.2015.02.001
Selby, E. A., Fehling, K. B., Panza, E. A., and Kranzler, A. (2016). Rumination, Van Rheenen, T. E., Murray, G., and Rossell, S. L. (2015). Emotion regulation in
mindfulness, and borderline personality disorder symptoms. Mindfulness 7, bipolar disorder: profile and utility in predicting trait mania and depression
228–235. doi: 10.1007/s12671-015-0432-5 propensity. Psychiatry Res. 225, 425–432. doi: 10.1016/j.psychres.2014.12.001

Varela, F. J., Thompson, E., and Rosch, E. (1991). The Embodied Mind: Wilcox, C. E., Pommy, J. M., and Adinoff, B. (2016). Neural circuitry of impaired
Cognitive Science and Human Experience. Cambridge, MA: MIT emotion regulation in substance use disorders. Am. J. Psychiatry. 173, 344–361.
Press. doi: 10.1176/appi.ajp.2015.15060710
Veenstra, L., Schneider, I. K., and Koole, S. L. (2016). Embodied mood regulation: Wupperman, P., Neumann, C. S., and Axelrod, S. R. (2008). Do deficits in
the impact of body posture on mood recovery, negative thoughts, and mood- mindfulness underlie borderline personality features and core difficulties? J.
congruent recall. Cogn. Emot. doi: 10.1080/02699931.2016.1225003. [Epub Pers. Disord. 22, 466–482. doi: 10.1521/pedi.2008.22.5.466
ahead of print]. Zeidan, F., Emerson, N. M., Farris, S. R., Ray, J. N., Jung, Y.,
Vestergaard-Poulsen, P., van Beek, M., Skewes, J., Bjarkam, C. R., Stubberup, McHaffie, J. G., et al. (2015). Mindfulness meditation-based pain
M., Bertelsen, J., et al. (2009). Long-term meditation is associated with relief employs different neural mechanisms than placebo and sham
increased gray matter density in the brain stem. Neuroreport 20, 170–174. mindfulness meditation-induced Analgesia. J. Neurosci. 35 15307–15325.
doi: 10.1097/WNR.0b013e328320012a doi: 10.1523/JNEUROSCI.2542-15.2015
Vrticka, P., and Vuilleumier, P. (2012). Neuroscience of human social Zeidan, F., Martucci, K. T., Kraft, R. A., Gordon, N. S., McHaffie, J.
interactions and adult attachment style. Front. Hum. Neurosci. 6:212. G., and Coghill, R. C. (2011). Brain mechanisms supporting the
doi: 10.3389/fnhum.2012.00212 modulation of pain by mindfulness meditation. J. Neurosci. 31, 5540–8.
Way, B. M., Creswell, J. D., Eisenberger, N. I., and Lieberman, M. D. (2010). doi: 10.1523/jneurosci.5791-10.2011
Dispositional mindfulness and depressive symptomatology: correlations with
limbic and self-referential neural activity during rest. Emotion 10, 12–24. Conflict of Interest Statement: The authors declare that the research was
doi: 10.1037/a0018312 conducted in the absence of any commercial or financial relationships that could
Westbrook, C., Creswell, J. D., Tabibnia, G., Julson, E., Kober, H., and Tindle, be construed as a potential conflict of interest.
H. A. (2013). Mindful attention reduces neural and self-reported cue-induced
craving in smokers. Soc. Cogn. Affect. Neurosci. 8, 73–84. doi: 10.1093/scan/ Copyright © 2017 Guendelman, Medeiros and Rampes. This is an open-access article
nsr076 distributed under the terms of the Creative Commons Attribution License (CC BY).
Whitaker, R. C., Dearth-Wesley, T., Gooze, R. A., Becker, B. D., Gallagher, The use, distribution or reproduction in other forums is permitted, provided the
K. C., and McEwen, B. S. (2014). Adverse childhood experiences, original author(s) or licensor are credited and that the original publication in this
dispositional mindfulness, and adult health. Prev. Med. 67, 147–153. journal is cited, in accordance with accepted academic practice. No use, distribution
doi: 10.1016/j.ypmed.2014.07.029 or reproduction is permitted which does not comply with these terms.

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published: 06 March 2017

Mindfulness and Emotion Regulation:

There is increasing interest in the beneficial clinical effects of mindfulness-based

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TABLE 2 | Psychological and neurocognitive models of mechanisms of MBIs.

Author Type of model Components

Shapiro et al., 2006 Psychological Attention, attitude, intention

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control) based functional magnetic

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control) based functional magnetic

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control) based functional magnetic

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*Finding indicates bottom-up mechanisms.

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REFERENCES Brefczynski-Lewis, J. A, Lutz, A, Schaefer, H. S., Levinson, D. B., and

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