Toward a Refined Mindfulness Model Related to Consciousness and Based on Event-Related Potentials

Abstract

Neuroimaging, behavioral, and self-report evidence suggests that there are four main cognitive mechanisms that support mindfulness: (a) self-regulation of attention, (b) improved body awareness, (c) improved emotion regulation, and (d) change in perspective on the self. In this article, we discuss these mechanisms on the basis of the event-related potential (ERP). We reviewed the ERP literature related to mindfulness and examined a data set of 29 articles. Our findings show that the neural features of mindfulness are consistently associated with the self-regulation of attention and, in most cases, reduced reactivity to emotional stimuli and improved cognitive control. On the other hand, there appear to be no studies of body awareness. We link these electrophysiological findings to models of consciousness and introduce a unified, mechanistic mindfulness model. The main idea in this refined model is that mindfulness decreases the threshold of conscious access. We end with several working hypotheses that could direct future mindfulness research and clarify our results.

Keywords

mindfulness event-related potential consciousness threshold self-regulation of attention emotion regulation cognitive control body awareness

Over the past three decades, researchers have highlighted the benefits of mindfulness on health and well-being, and the number of studies on the topic has increased. A search for the term mindfulness in the PubMed database found more than 5,800 references, the first dating from 1982. A search restricted to 2010 to 2018 found more than 5,300 references, which reflects the substantial increase in interest in recent years. Most researchers have investigated the impact of mindfulness meditation (MM) on clinical populations, drawing on programs such as mindfulness-based stress reduction (MBSR) or mindfulness-based cognitive therapy (MBCT) intended to reduce psychological distress and disease-related physical symptoms such as pain (Kabat-Zinn, 1990; Segal, Williams, & Teasdale, 2012). These studies have consistently reported an improvement in health (with decreased psychological and physical symptoms) and an increase in well-being (Chiesa & Serretti, 2011; Fjorback, Arendt, Ørnbøl, Fink, & Walach, 2011; Goldberg et al., 2018). If the answer to the question “Does mindfulness work?” is clearly positive, the answer to the question “How does mindfulness work?” remains at the frontier of science.

Therefore, in this article, we discuss potential mindfulness mechanisms. In particular, we propose a mindfulness model that focuses on conscious access to information. The starting point is Hölzel’s model, which is based on neuroimaging, behavioral, and self-reported evidence. The latter model describes four main cognitive mechanisms through which mindfulness may work: (a) the self-regulation of attention, (b) change in perspective on the self, (c) enhanced body awareness, and (d) improved emotion regulation (Hölzel et al., 2011).

The aim of our work was to refine this initial model and, in particular, to articulate these four mechanisms with respect to each other. Our refined model suggests that mindfulness (a) lowers the threshold of conscious access as a function of attentional mechanisms (self-regulation of attention in Hölzel’s model) and (b) facilitates the conscious processing of information that comes from within (body awareness and self-awareness) and outside the body (world awareness). The enhanced body awareness and change in perspective on the self found in Hölzel’s model may result from improved conscious processing of information from within, whereas better emotion regulation may be due to improved conscious processing of self-awareness and world awareness.

Finally, we evaluate our mindfulness model with respect to the current event-related potential (ERP) and mindfulness literature.

The Significance of Studies of Event-Related Potential

There are two ways to investigate mindfulness mechanisms: ERP and neuroimaging. In this study, we used ERP because the temporal resolution of electroencephalography renders the method particularly suitable for investigating the very early cognitive mechanisms (from 100 to 600 ms after information onset) involved in the processing of incoming information (Gosseries et al., 2008). Moreover, conscious access to information seems to occur in a similar temporal window of around 270 to 300 ms (Gaillard et al., 2009).

ERP is computed by recording electrical brain activity locked to a specific event (e.g., the presentation of a stimulus or the participant’s motor response). It reflects activity in ensembles of cortical neurons that has a fixed temporal relation to a specific event. Thus, ERP provides information about the participant’s cognitive response to a specific event.

Most ERP-based studies of mindfulness have investigated healthy populations, notably MM practitioners and individuals with dispositional mindfulness (i.e., people who are naturally mindful and do not have a specific MM practice). In theory, this focus on healthy populations makes it possible to investigate mindfulness mechanisms in an, a priori, unaltered brain.

Potential Connections Between Mindfulness Models and Consciousness Models

Developing a joint mindfulness/consciousness model makes sense given that mindfulness and consciousness (within the meaning of conscious awareness) seem to be closely linked. For example, neuroimaging data suggest that some of the brain areas involved in MM and consciousness (notably the anterior cingulate cortex, the insula, the posterior cingulate cortex, some regions of the prefrontal cortex, and the thalamus) could overlap (Manuello, Vercelli, Nani, Costa, & Cauda, 2016).

Models of consciousness

Models of consciousness suggest that the conscious visual perception of a stimulus involves two main processes. The first is bottom-up: the propagation of sensory signals through the visual hierarchy; the second is top-down: attention amplification by late, higher-level integrative processes. Sensory information is continuously processed in an unconscious manner, whereas conscious access is thought to start when attention amplifies a given piece of information, which allows it to access a network of high-level brain regions that are broadly interconnected by long-range connections (Baars, 1993; Dehaene, 2011; Dehaene & Changeux, 2011; De Lafuente & Romo, 2006). This so-called global neuronal workspace integrates incoming information into the current, conscious context, where it becomes available to other neural processes. Conscious access to incoming sensory evidence has been linked to a “decision to engage” the global workspace (Dehaene, Izard, Spelke, & Pica, 2008; Shadlen & Kiani, 2011).

According to these theoretical models, an elevated consciousness threshold could result from a bottom-up perceptual impairment (the object remains subliminal) or a lack of top-down attentional amplification (the object remains preconscious; Dehaene, Changeux, Naccache, Sackur, & Sergent, 2006), both of which prevent conscious access to information (Berkovitch, Del Cul, Maheu, & Dehaene, 2018). Conversely, it seems reasonable to assume that attentional amplification could result in a lower consciousness threshold. Given that mindfulness aims to enhance attentional skills (notably the self-regulation of attention, which includes focused attention), we suggest that the consciousness threshold can be adjusted by modulating the process of attention amplification, as described in theoretical models of conscious processing (Dehaene & Changeux, 2003, 2011). By promoting attentional skills, mindfulness could lower the consciousness threshold and consequently facilitate world awareness (awareness of the external world), self-awareness (awareness of thoughts and emotions), and body awareness (interoception). In other words, mindfulness could help to overcome the situation in which information from the world, the self, and the body remains preconscious by facilitating the conscious processing of this information.

In the next section, we link the mechanisms presented in Hölzel’s mindfulness model (Hölzel et al., 2011) to our refined model.

Self-regulation of attention

The practice of mindfulness aims to enhance the self-regulation of attention. The latter relies on two cognitive abilities: focused attention or open monitoring (Lutz, Jha, Dunne, & Saron, 2015). Focused attention enables individuals to deliberately access an extensive sensory experience and is primarily based on breathing. Open monitoring enhances the ability of individuals to adopt a nonjudgmental attitude to anything that they experience (Lutz, Slagter, Dunne, & Davidson, 2008; Malinowski, 2013). We suggest that the self-regulation of attention per se could support the top-down process of attentional amplification, as described in models of consciousness.

Change in perspective on the self

As Hölzel et al. (2011) stated, change in perspective on the self is difficult to define. The authors described the self as “being the one who inhabits the body, being the one who is thinking the thoughts, being the one experiencing emotions, and being the agent of actions” (p. 547). It can be considered to emerge from a specific brain activity that produces thoughts, emotions, and actions.

The perception of a self leads to the development of meta-awareness. Meta-awareness, in turn, contributes to cognitive control by orienting attention toward the contents of conscious experience and associated processes (Hölzel et al., 2011; Schooler, 2002).

In mindfulness, change in perspective on the self could result from focused attention on self-activities. By bringing the content of experience (i.e., thoughts and emotions) to consciousness, mindful functioning could improve cognitive control through the selection of the most relevant information. Such information could contribute to resolving conflicts, leading to action, and response inhibition or adjustment.

Enhanced body awareness

Mindfulness practice relies on training the mind to pay sustained attention to the body experience, primarily the breath, and deliberately returning attention to it whenever distracted (Lutz et al., 2015). Body awareness refers to the ability to feel engaged by information coming from the body and to notice subtle changes (Mehling et al., 2009). Such information informs individuals about both their internal physiological state (interoception) and their body in relation to space and movement (exteroception; Craig, 2002; Mehling et al., 2009; Valenzuela-Moguillansky, Reyes-Reyes, & Gaete, 2017).

The enhanced body awareness resulting from mindfulness could be due to focusing the attention on the body, which facilitates conscious access to increasingly subtle physical sensations. Body awareness has been found to increase after mindfulness practice and is associated with dispositional mindfulness (Carmody & Baer, 2008; Farb et al., 2015; Hanley, Mehling, & Garland, 2017; Mehling et al., 2012; Treves, Tello, Davidson, & Goldberg, 2019).

Improved emotion regulation

Emotion regulation could be defined as the ability to modify the intensity and duration of emotional responses in the context of meeting goals and managing arousal (Thompson, 1994). It may also extend to the ability to modify the emotion’s significance given that cognitive strategies can alter how emotional stimuli are attended to and interpreted (Dennis & Hajcak, 2009).

In the context of mindfulness, it is thought to rely on two cognitive strategies: “positive reappraisal” and “extinction” (Hölzel et al., 2011, p. 543). Reappraisal is a cognitive strategy that seeks to reinterpret an emotional situation in a different way (Hajcak & Nieuwenhuis, 2006), and it assumes there is conscious access to emotional content. Extinction refers to an active effort to limit internal engagement when exposed to negative emotions, which reduces avoidance and leads to the reversal of stimulus-response conditioning (Chambers, Gullone, & Allen, 2009; Garland, Gaylord, & Fredrickson, 2011; Hölzel et al., 2011). Mindfulness has been associated with decreased reactivity to both negative and positive emotions—in other words, improved emotion regulation (Guendelman, Medeiros, & Rampes, 2017). The amplification of attention and the subsequent facilitation of conscious access may explain this ability to deal with emotions in a nonjudgmental manner.

Method

Our literature review followed Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines (Moher, Liberati, Tetzlaff, & Altman, 2009). The initial data set consisted of the Scopus, PubMed, Google Scholar, and APA PsychNET online databases. Search criteria were “mindfulness” AND “event related potential” in (a) the article’s title, abstract, and keywords in Scopus; (b) the article’s title and abstract in PubMed; (c) the article’s title in Google Scholar; and (d) the abstract in APA PsychNET. We considered publications from 1990 (the year Kabat-Zinn’s book on mindfulness was published) to October 1, 2018. Regarding participants, interventions, comparisons, outcomes, and study design (PICOS) characteristics, the key criteria were adults (for participants), mindfulness (for interventions), and ERPs (for outcomes). All articles were published in English.

Inclusion criteria were (a) that the article directly addressed the topic of mindfulness or an equivalent meditation practice (i.e., Vipassana, Zen, or Buddhist meditation), (b) ERPs were recorded in relation to mindfulness, and (c) the study examined a sample of adults. Exclusion criteria were theoretical articles, commentaries, or editorials.

The following variables were extracted from each article: (a) mindfulness components (i.e., MM intervention, mindfulness-based clinical program, MM practice or dispositional mindfulness); (b) participants’ experience of their MM practice (duration and frequency); (c) population features (healthy or clinical); (d) the experimental design (mindfulness and control groups), measurement time (before and after intervention), and experimental tasks; and (e) ERP outcomes (amplitude, latency).

The main measurements were differences in ERP amplitude among either (a) mindfulness (MM intervention, mindfulness-based clinical program, or healthy meditators) and control (control or nonmeditator groups) conditions, (b) levels of dispositional mindfulness, or (c) mindfulness-meditative and mind-wandering conditions.

Results and Discussion

Researchers of ERP studies of mindfulness have investigated cognitive control, the regulation of attention, and emotion regulation. Our review did not find any ERP studies that reported an investigation of the body-awareness mechanism. For the sake of clarity, we have combined the presentation and discussion of our results with respect to each mechanism separately.

Search results

Our review initially identified 68 citations (39 from the Scopus database, 23 from PubMed, five from Google Scholar, and one from APA PsychNET). After the removal of duplicates, 55 articles were considered relevant. A review of the abstracts led to the exclusion of 26 articles. Nine were not research articles, four were not related to mindfulness, five did not record ERP in relation to mindfulness, five were not carried out in an adult population, and three were not published. The final data set consisted of 29 articles (Fig. 1).

Fig. 1.

Flow diagram showing preferred reporting items included in the review.

General description of ERP studies

Eleven articles reported the investigations of the self-regulation of attention and were based on 11 independent samples (N = 357; age range = 17–80 years; Table 1). Twelve articles reported investigations of cognitive control and were based on 12 independent samples (N = 545; age range = 17–80 years; Table 2). It should be noted that in four studies, both the self-regulation of attention and cognitive control were investigated (Atchley et al., 2016; Malinowski, Moore, Mead, & Gruber, 2017; Moore, Gruber, Derose, & Malinowski, 2012; Norris, Creem, Hendler, & Kober, 2018). Finally, authors of 10 articles focused on emotion regulation in 10 independent samples (N = 411; age range = 18–70 years; Table 3).

Table 1.

Summary of Event-Related Potential (ERP) Studies Focusing on the Self-Regulation of Attention

Study	Mindfulness component	Experience	Experimental design	Task	ERP	Findings
Cahn and Polich (2009)	Vipassana meditation	20 years 30 min/day	Meditative state vs. mind-wandering	Auditory oddball task	P3a	P3a amplitude was lower for meditative state relative to mind-wandering state. The reduction in P3a amplitude in meditative state was positively correlated to the frequency of MM practice.
Delgado-Pastor et al. (2013)	Vipassana meditation	7.5 years 15 hr/week	Pre/post meditation, meditative state vs. mind-wandering	Auditory oddball task	P3b	P3b amplitude was greater both after meditation than before, and in meditative state compared with mind-wandering state.
Atchley et al. (2016)	Tibetan or Zen or Buddhist meditation	Experienced meditators: 22.6 years; novice meditators: 2.4 years; naive meditators	Meditators vs. control	Auditory oddball task + breathing counting task	P3b	P3b amplitude was higher for meditators than naive meditators in the auditory task. In contrast, when instructed to ignore tones, P3b amplitude was lower for meditators than naive meditators.
Lakey et al. (2011)	MM induction (6 min)	—	MM induction vs. control	Visual attentional task	P3b	P3b amplitude was higher in the MM induction group compared with the control group.
Moore et al. (2012)	MM training (16 weeks)	—	MM training vs. waiting list, pre/post intervention	Stroop task	P3b	After intervention, a relative decrease in P3b amplitude was found in the MM training group in incongruent trials. In contrast, it increased in the waiting list group in the same trials.
Malinowski et al. (2017)	MM training (8 weeks)	—	MM training vs. active control	Stroop task	P3b	In a population of healthy older adults, P3b amplitude was not modulated by intervention, both in the MM training group and in the control group.
Howells et al. (2012)	MBCT (8 weeks)	—	Pre/post intervention	Go/no-go task	P3b	In a population of bipolar disorder patients, the MBCT intervention attenuated P3b amplitude in response to a task-irrelevant stimulus.
Slagter et al. (2007)	Vipassana meditation retreat (3-month retreat)	Experienced meditators: 10 hr/day; novice meditators: 1 hr/week	Pre/post intervention	Attentional-blink task	P3b	After intervention, P3b amplitude was lower in experienced meditators in response to irrelevant stimuli in Time 1. No effect was observed for novice meditators.
Norris et al. (2018)	Brief MM (10 min)	—	MM training vs. active control	Attention network task	P3b	No effect of the intervention on P3b amplitude was observed.
Smart et al. (2016)	MM training (8 weeks)	—	MM training vs. active control, healthy vs. clinical older adults	Go/no-go task	P3b	In a population of older adults with subjective cognitive decline, P3b amplitude increased, after intervention in the MM training group, whereas it remained unchanged in the control group. No effect of MM training was observed in healthy older adults.
Bostanov et al. (2012)	MBCT (8 weeks)	—	MBCT vs. waiting list, pre/post intervention	Original task	CNV	In a population of recurrently depressed patients, CNV amplitude increased after intervention in the MBCT group, whereas it remained unchanged in the waiting list group.

Note: MM = mindfulness meditation; MBCT = mindfulness-based cognitive therapy; CNV = contingent negative variation.

Table 2.

Summary of Event-Related Potential (ERP) Studies Focusing on Cognitive Control

Study	Mindfulness component	Experience	Experimental design	Task	ERP	Findings
Teper and Inzlicht (2012)	Vipassana	3.19 years	Meditators vs. control	Stroop task	ERN, Pe	Higher ERN amplitude was observed for meditators compared with control subjects in incorrect trials. Experience of meditation was correlated with ERN amplitude. In contrast, Pe amplitude was not higher in meditators compared with control subjects and was not correlated to meditation experience.
Schoenberg et al. (2014)	MBCT (12 weeks)	—	MBCT vs. waiting list, pre/post intervention	Go/no-go task	ERN, N2, Pe, No-go P3	In a population of ADHD patients, Pe amplitude and no-go P3 components increased after intervention in the MBCT group, whereas it remained unchanged in the waiting list group. ERN and N2 amplitudes were not modulated in either group.
Moore et al. (2012)	MM training (16 weeks)	—	MM training vs. waiting list, pre/post intervention	Stroop task	N2	After intervention, N2 amplitude increased in the MM training group in incongruent trials, whereas it decreased in the waiting list group.
Smart and Segalowitz (2017)	MM training (8 weeks)	—	MM training vs. active control	Eriksen flanker task	ERN, Pe	In a population of older adults with subjective cognitive decline, ERN amplitude increased after intervention in the MM training group, whereas it remained unchanged in the control group. Pe amplitude was not modulated by the intervention in either group.
Eichel and Stahl (2016)	DM	—	Mindful vs. nonmindful	Simon task	ERN	ERN amplitude was greater for detected compared with undetected errors, but only for mindful individuals.
Norris et al. (2018)	Brief MM (10 min)	—	MM training vs. active control	Attention network task	N2	After intervention, N2 was higher in individuals with lower neuroticism scores in the MM training group, whereas no significant change was observed in the control group. No effect was observed in individuals with higher neuroticism scores.
Malinowski et al. (2017)	MM training (8 weeks)	—	MM training vs. active control, Pre/post intervention	Stroop task	N2	N2 amplitude increased in the MM training group after intervention, whereas no significant change was observed in the control group.
Quaglia et al. (2016)	DM	—	Mindful vs. nonmindful	Go/no-go task	N2, No-go P3	Both N2 and no-go P3 amplitudes were higher in mindful compared with nonmindful individuals.
Atchley et al. (2016)	Tibetan, Zen, or Buddhist meditation	Experienced meditators: 22.6 years; novice meditators: 2.4 years; naive meditators	Meditators vs. control	Auditory oddball task + breathing counting task	N2	N2 amplitude was higher in meditators than naive meditators in the auditory task. In contrast, when instructed to ignore tones, N2 amplitude was lower in meditators than naive meditators.
Larson et al. (2013)	MM induction (15 min)	—	MM induction vs. active control	Eriksen flanker task	ERN, Pe	ERN amplitude did not change following the intervention in either mindfulness or control groups. In contrast, postintervention Pe amplitude was smaller in the MM group than in control group.
Saunders et al. (2016)	MM induction (13 min)	—	Emotion-focused vs. thought-focused	Go/no-go task	ERN, Pe	Emotion-focused participants showed enhanced ERN amplitude after intervention, whereas thought-focused participants did not. In contrast, Pe amplitude was unaffected by the intervention.
Bing-Canar et al. (2016)	MM induction (15 min)	—	MM induction vs. active control	Stroop task	ERN, Pe	Postintervention ERN and Pe amplitudes did not differ between MM and control groups.

Note: ERN = error-related negativity; Pe = positivity error; MBCT = mindfulness-based cognitive therapy; ADHD = attention-deficit/hyperactivity disorder; DM = dispositional mindfulness.

Table 3.

Summary of Event-Related Potential (ERP) Studies Focusing on Emotional Regulation

Study	Mindfulness component	Experience	Experimental design	Task	ERP	Findings
Brown et al. (2013)	DM	—	Mindful vs. nonmindful	Affective picture viewing task	LPP	More mindful individuals showed lower LPP amplitude in response to both unpleasant and pleasant stimuli, compared with less mindful individuals.
Cosme & Wiens (2015)	DM	—	Mindful vs. nonmindful	Affective picture viewing task	LPP	No significant effect of DM on LPP amplitude was observed.
Ho et al. (2015)	TM in a population of meditators	6.52 years	Mindful vs. nonmindful	Active affective picture viewing task	LPP	No effect of TM on LPP amplitude was observed.
Sobolewski et al. (2011)	Buddhist meditation	At least 5 years and 5 hr/week	Meditators vs. control	Affective picture viewing task	LPP	Meditators showed lower LPP amplitude in response to negative stimuli compared with naive participants.
Lin et al. (2016)	DM	—	Mindful vs. nonmindful	Affective picture viewing task	LPP	DM was associated with a reduction in LPP amplitude in response to negative stimuli. A task-induced state of mindfulness failed to modulate LPP amplitude compared with the control condition. LPP amplitude decreased after intervention in the MM exercise group compared with an active control.
	Task-induced state of mindfulness		Task-induced state of mindfulness vs. control condition
	MM exercise		MM exercise vs. active control
Egan et al. (2017)	DM	—	Mindful vs. nonmindful	Affective picture viewing task	LPP	A task-induced state of mindfulness was associated with greater LPP compared with the control condition. DM did not correlate with LPP amplitude in the two conditions.
Egan et al. (2017)	Task-induced state of mindfulness	—	Task-induced state of mindfulness vs. control condition	Affective picture viewing task	LPP
Garland et al. (2015)	MORE (8 weeks)	—	MORE vs. active control	Affective picture viewing task	LPP	In a population of patients with chronic pain and a history of opioid misuse, an increase in LPP activation in response to natural reward cues was observed for the MORE group relative to the control group.
Uusberg, et al. (2016)	Task-induced state of mindfulness	—	Task-induced state of mindfulness vs. control conditions	Affective picture viewing task	LPP	A task-induced state of mindfulness was associated with greater LPP amplitude in response to negative stimuli than control conditions.
Howells et al. (2014)	MBCT (8 weeks)	—	Pre/post intervention	Visual matching task	N170	After an MBCT intervention, an attenuation in N170 amplitude was observed for bipolar disorder patients.
Dorjee et al. (2015)	DM	—	Mindful vs. nonmindful	Simon task	P600	P600 amplitude was observed to be lower in more mindful subjects compared with less mindful subjects.

Note: DM = dispositional mindfulness; LPP = late positive potential; TM = trait mindfulness; MM = mindfulness meditation; MORE = mindfulness-oriented recovery enhancement; MBCT = mindfulness-based cognitive therapy.

Because four articles were based on overlapping samples (Howells, Ives-Deliperi, Horn, & Stein, 2012; Howells, Laurie Rauch, Ives-Deliperi, Horn, & Stein, 2014; Smart & Segalowitz, 2017; Smart, Segalowitz, Mulligan, Koudys, & Gawryluk, 2016), our data set of 29 articles drew on results from 27 independent samples (N = 1,185; age range = 17–80 years). Participants were either healthy participants (22 independent samples, N = 954; age range = 17–80 years) or patients (five independent samples, N = 231; age range = 18–80 years). In the latter case, patients were either depressed (Bostanov, Keune, Kotchoubey, & Hautzinger, 2012, N = 64), bipolar (Howells et al., 2012; Howells et al., 2014, N = 12), suffering from chronic pain and at risk of opioid abuse (Garland, Froeliger, & Howard, 2015, N = 29), suffering from attention-deficit/hyperactivity disorders (Schoenberg et al., 2014, N = 50), or older adults experiencing subjective cognitive decline (Smart et al., 2016; Smart & Segalowitz, 2017, N = 76).

Our results can be categorized into (a) studies examining change in ERP amplitude in relation to MM practice (MM interventions, mindfulness-based clinical programs, and healthy meditators), (b) studies examining modulation in ERP amplitude as a function of dispositional mindfulness, and (c) studies focusing on the effects of a mindful state in comparison with a mind-wandering state or a control condition (Tables 1 –4).

Table 4.

Synthesis of Articles as a Function of Principal Findings, Cognitive Mechanisms, and ERP

Outcome	Mindful > nonmindful	Mindful = nonmindful	Mindful < nonmindful	Mindful > RS	Mindful < RS	Other
Self-regulation of attention
P3a					Cahn & Polich (2009)
P3b	Atchley et al. (2016; oddball task); Lakey et al. (2011); Smart et al. (2016)	Malinowski et al. (2017); Norris et al. (2018)	Atchley et al. (2016; meditative state); Moore et al. (2012); Howells et al. (2012); Slagter et al. (2007)	Delgado-Pastor et al. (2013)
CNV	Bostanov et al. (2012)
Emotion regulation
LPP	Egan et al. (2017; meditative state); Uusberg et al. (2016)	Cosme & Wiens (2015); Ho, Sun et al. (2015); Lin et al. (2016; meditative state); Egan et al. (2017; DM)	Brown et al. (2013); Sobolewski et al. (2011); Lin et al. (2016; DM and MM intervention)			Garland et al. (2015)
N170			Howells et al. (2014)
P600			Dorjee et al. (2015)
Cognitive control
ERN	Teper & Inzlicht (2012); Smart & Segalowitz (2017)	Schoenberg et al. (2014); Larson et al. (2013); Bing-Canar et al. (2016)	–			Saunders et al. (2016); Eichel & Stahl (2016)
N2	Quaglia et al. (2016); Moore et al. (2012); Atchley et al. (2016; oddball task); Malinowski et al. (2017); Norris et al. (2018)	Schoenberg et al. (2014);	Atchley et al. (2016; meditative state)
Pe	Schoenberg et al. (2014)	Teper & Inzlicht (2012); Bing-Canar et al. (2016); Smart & Segalowitz (2017)	Larson et al. (2013)
no-go P3	Quaglia et al. (2016); Schoenberg et al. (2014)

Note: > = significantly greater than; < = significantly lower than; = indicates no significant differences; RS = resting state; CNV = contingent negative variation; ERN = error related negativity; Pe = positivity error; DM = dispositional mindfulness; MM = mindfulness meditation. All findings refer to ERP amplitude.

Regulation of attention

Results

Two components were investigated in relation to the regulation of attention: P300 and contingent negative variation (CNV).

P300

P300 was the main ERP of interest in studies focused on attention and was reported in 10 of the 11 articles (91%). The late (P3b) component was more frequently investigated (nine of 10 articles) than the early (P3a) component.

P3a reflects automatic attention attraction and can occur in a nonconscious way (Muller-Gass, Macdonald, Schröger, Sculthorpe, & Campbell, 2007). Conversely, P3b, which occurs about 300 to 500 ms after stimulus onset, is thought to be a signature of conscious perception (for a review, see Dehaene & Changeux, 2011). Although the association with consciousness is challenged, P3b is thought to be related to the frontoparietal network that plays a key role in task monitoring and reporting (Koch, Massimini, Boly, & Tononi, 2016).

In most cases, mindfulness was associated with greater P3b amplitude in tasks based on attention to task-relevant stimuli (e.g., tones in an auditory oddball task; Atchley et al., 2016; Delgado-Pastor, Perakakis, Subramanya, Telles, & Vila, 2013; Lakey, Berry, & Sellers, 2011; Smart et al., 2016). This observation was found (a) in experienced meditators at rest (relative to naive meditators; Atchley et al., 2016) and after 30 min of MM practice (Delgado-Pastor et al., 2013), (b) after 8 weeks of MM training in a clinical population with subjective cognitive decline (Smart et al., 2016), and (c) after a very brief mindfulness intervention (6 min) in a healthy population (Lakey et al., 2011). On the other hand, two studies failed to detect a positive effect of mindfulness on the P3b component (Malinowski et al., 2017; Norris et al., 2018). The P3b component was also investigated in response to stimuli irrelevant to the task that were introduced as distractors. In this case, lower P3b amplitude was reported in meditators (Atchley et al., 2016), after a mindfulness intervention (8 weeks, 16 weeks, or 3 months) in a healthy population (Moore et al., 2012; Slagter et al., 2007), and in a clinical population with bipolar disorder (Howells et al., 2012). Finally, the amplitude of the early P3a component was found to be lower in a meditative state than in a mind-wandering state, and this observation was positively correlated with MM practice frequency (Cahn & Polich, 2009).

Contingent negative variation

In a sequence of paired warning-imperative stimuli, CNV amplitude is positively correlated with the subjective probability of the imperative stimuli (Walter, Cooper, Aldridge, McCallum, & Winter, 1964). The CNV is related to response preparation, attention, and arousal level given that its cerebral generators include motor, premotor, and other cortex areas (Tecce, 1972; Tecce & Cattanach, 1993).

Just one article investigated CNV in mindfulness, and its amplitude was increased after an 8-week MBCT intervention in recurrently depressed patients (Bostanov et al., 2012).

Discussion

If P3b is the electrophysiological sign of conscious access, an increase in amplitude should indicate better access to consciousness. The larger P3b amplitude found in mindfulness experiments (Atchley et al., 2016; Delgado-Pastor et al., 2013; Lakey et al., 2011; Smart et al., 2016) supports the hypothesis that mindfulness decreases the threshold of conscious access. However, at the same time, mindfulness was also associated with lower P3b amplitude in response to task-irrelevant stimuli (i.e., a distractor; Atchley et al., 2016; Howells et al., 2012; Moore et al., 2012; Slagter et al., 2007). Finally, task-relevant stimuli were found to elicit higher CNV amplitude in a mindfulness condition (Bostanov et al., 2012).

Taken together, these findings suggest that better conscious access to information in mindfulness (through a process of attentional amplification) could be solely oriented toward goal-relevant information.

In the context of the global neuronal workspace model (Baars, 1993; Dehaene, 2011; Dehaene & Changeux, 2011; De Lafuente & Romo, 2006), conscious access is related to a “decision to engage” the global workspace when faced with incoming information (Dehaene et al., 2008; Shadlen & Kiani, 2011). Bringing information to consciousness could be viewed as a decisional process, a choice between leaving information unconscious or helping it to reach consciousness. It could be argued that when the stimulus is goal-irrelevant, the decision should be to keep it unconscious. The idea of a lower threshold of conscious access is congruent with this hypothesis because attentional amplification helps to keep information under the threshold or bring it above the threshold depending on its relevance to the context. Moreover, an early decision of whether to bring information to consciousness avoids blocking conscious access and prevents information loss as a result of the psychological refractory period¹ (Marti, Sigman, & Dehaene, 2012).

To summarize, attentional amplification (which lowers the conscious access threshold) could be associated with an attentional focus on pertinent information at the expense of nonpertinent information. The upshot of this could be an optimization that seeks to fill the global neural workspace with pertinent information and avoid saturation with irrelevant information, thus enabling more fluent, effortless thinking.

Furthermore, mindfulness is negatively associated with P3a amplitude (Cahn & Polich, 2009). P3a reflects the automatic capture of attention by any new stimulus (stimulus-driven attention) and can occur unconsciously (Muller-Gass et al., 2007; Polich, 2007). We suggest that mindful functioning is characteristic of unbiased information processing that operates via the disengagement of the attentional system to stimulus-driven activation. In other words, the attentional system could be similarly engaged for all stimuli irrespective of their novelty and without any expectations (i.e., an unexpected stimulus could merely be interpreted as a novel one without any implication of automatic attention). This engagement of automatic attention can be compared with the nonjudgmental attitude that contributes to the definition of mindfulness (Kabat-Zinn, 1990, 1994).

To sum up, mindful functioning could allow the unconscious and unbiased processing of incoming information. Attentional amplification may facilitate relevant information reaching the consciousness to the detriment of irrelevant information. This trade-off is illustrated in the finding that P3a and P3b amplitudes oppose each other.

Cognitive control

Results

The 12 studies of cognitive control mainly focused on the following three ERPs: (a) the N2 component (six articles, 50%), (b) error-related negativity (ERN; seven articles, 58%), and (c) the positivity error (five articles, 42%). An additional ERP—the no-go P3 component—was investigated in two articles.

N2

The N2 component peaks between 200 and 350 ms after stimulus onset. Because its amplitude is sensitive to a mismatch between a stimulus and a mental template, it is thought to be a conflict-monitoring index (Bruin, Wijers, & Van Staveren, 2001; Donkers & Van Boxtel, 2004; Nieuwenhuis, Yeung, Van Den Wildenberg, & Ridderinkhof, 2003). It may also be driven by the inhibition of a planned response (for a review, see Folstein & Van Petten, 2008).

In our review, we found that mindfulness was mainly associated with increased N2 amplitude (Atchley et al., 2016; Malinowski et al., 2017; Moore et al., 2012; Norris et al., 2018; Quaglia, Goodman, & Brown, 2016). Specifically, (a) amplitude was positively correlated with dispositional mindfulness (Quaglia et al., 2016), and (b) its amplitude was higher in experienced than naive meditators (Atchley et al., 2016) and (c) higher following MM interventions (10 min, 8 or 16 weeks) in a healthy population (Malinowski et al., 2017; Moore et al., 2012; Norris et al., 2018). Just one study failed to detect a positive effect of mindfulness (Schoenberg et al., 2014).

Error-related negativity and error positivity

ERN (Gehring, Goss, Coles, Meyer, & Donchin, 1993), also known as “error negativity” (Falkenstein, Hohnsbein, Hoormann, & Blanke, 1991), peaks around 100 ms after the electromyographic response. ERN is a well-established index of action monitoring (Holroyd & Coles, 2002). Moreover, recent work has suggested that it reflects an internal comparison between two signals: an unconscious representation of the ongoing action and a conscious representation of the intended action (Dehaene, 2018). ERN is followed by error positivity² (Pe), which occurs about 200 to 500 ms after an error (Falkenstein et al., 1991; Overbeek, Nieuwenhuis, & Ridderinkhof, 2005). Pe amplitude is larger for consciously perceived errors than unperceived errors (Endrass, Reuter, & Kathmann, 2007; Murphy, Robertson, Allen, Hester, & O’Connell, 2012; Nieuwenhuis, Ridderinkhof, Blom, Band, & Kok, 2001). The Pe component, therefore, is believed to reflect error awareness (Steinhauser & Yeung, 2010; Ullsperger, Danielmeier, & Jocham, 2014).

In contrast to N2 findings reported above, just 33% of studies reported higher ERN amplitude in mindfulness. In the same way, just 20% of studies reported higher Pe amplitude in mindfulness (Schoenberg et al., 2014; Smart & Segalowitz, 2017; Teper & Inzlicht, 2012). Although experienced meditators were found to have a higher ERN amplitude than naive meditators, this was not found for the Pe component (Teper & Inzlicht, 2012). A very brief mindfulness intervention (15 min) was found to lead to an increase, a decrease, and no change—depending on the ERP component and the study (Bing-Canar, Pizzuto, & Compton, 2016; Larson, Steffen, & Primosch, 2013; Saunders, Rodrigo, & Inzlicht, 2016). The same finding was observed for MM interventions in clinical populations. A 12-week MBCT intervention in patients with attention-deficit/hyperactivity disorders was found to increase the Pe component without modifying ERN and N2 components (Schoenberg et al., 2014). At the same time, an 8-week MM training program in a population of older adults with subjective cognitive decline increased ERN amplitude without changing the Pe component (Smart & Segalowitz, 2017).

No-go P3

The no-go P3 component emerges in the frontocentral region 300 to 600 ms after stimulus presentation and reflects response inhibition (Falkenstein, Hoormann, Christ, & Hohnsbein, 2000; Salisbury, Griggs, Shenton, & McCarley, 2004). In our review, we found that mindfulness was consistently associated with higher no-go P3 amplitude (Quaglia et al., 2016; Schoenberg et al., 2014).

Discussion

The enhanced cognitive control found in mindfulness is thought to be a consequence of an increase in self-awareness (i.e., increased conscious processing of thoughts and emotions) and world awareness (i.e., better conscious access to information from the external world).

The findings from ERP studies support this hypothesis because N2 amplitude is higher in mindfulness (Atchley et al., 2016; Malinowski et al., 2017; Moore et al., 2012; Norris et al., 2018; Quaglia et al., 2016). However, results based on the ERN component are puzzling. Our model could predict either a decrease or an increase in ERN amplitude in mindfulness, depending on the functional significance. On the one hand, if we adopt the position of better world awareness, consistent with Dehaene’s hypothesis³ regarding ERN functional significance, our model predicts that mindfulness would increase ERN amplitude by improving the conscious representation of the intended action (Dehaene, 2018). On the other hand, mindfulness could decrease ERN amplitude following an emotion-reappraisal strategy (Hölzel et al., 2011) because ERN amplitude is reduced when negative emotions are attenuated through cognitive reappraisal (Hobson, Saunders, Al-Khindi, & Inzlicht, 2014). Concerning the Pe component, although the reviewed evidence is mixed, our model assumes greater error awareness in mindfulness and predicts an increase in Pe amplitude.

Finally, mindfulness could also improve cognitive control through better inhibition, as reflected in the enhanced no-go P3 component (Quaglia et al., 2016; Schoenberg et al., 2014). ERP findings are consistent with behavioral evidence showing improved performance in the Stroop task by experienced meditators relative to naive meditators (Chan & Woollacott, 2007; Moore & Malinowski, 2009) or after an MM intervention (Wenk-Sormaz, 2005).

Emotion regulation

Results

Three components were investigated in relation to emotion regulation: the late positive potential (LPP), N170, and P600.

Late positive potential

The LPP was the main ERP of interest in the reviewed studies (10 articles, 91%). It occurs around 400 to 600 ms after stimulus and refers to the P3b component in situations with emotional stimuli (Schupp, Flaisch, Stockburger, & Junghöfer, 2006). LPP is enhanced in participants watching emotionally arousing pictures as opposed to neutral pictures. Moreover, Foti and Hajcak (2008) demonstrated that LPP amplitude after exposure to unpleasant pictures was reduced when a more neutral interpretation of the picture was given. Therefore, the reduction in LPP after a directed, positive reappraisal may reflect reduced emotional reactivity because of a strategy of emotion regulation (Foti & Hajcak, 2008).

In three ERP studies, mindfulness was associated with lower LPP amplitude in response to negative and positive emotional stimuli (Brown, Goodman, & Inzlicht, 2013; Lin, Fisher, Roberts, & Moser, 2016; Sobolewski, Holt, Kublik, & Wrobel, 2011). More precisely, (a) higher dispositional mindfulness was associated with lower LPP amplitude (Brown et al., 2013; Lin et al., 2016), (b) LPP amplitude was lower in experienced meditators compared with naive meditators (Sobolewski et al., 2011), and (c) MM interventions in a healthy population were associated with a decrease in LPP amplitude (Lin et al., 2016). On the other hand, mindfulness was associated with higher LPP amplitude in two studies (Egan, Hill, & Foti, 2017; Uusberg, Uusberg, Talpsep, & Paaver, 2016), and three studies reported no effect (Cosme & Wiens, 2015; Ho, Sun, Ting, Chan, & Lee, 2015; Lin et al., 2016). Note that two studies found a mixed effect of mindfulness on the LPP component depending on the experimental condition (dispositional mindfulness, meditative state, or MM intervention; Egan et al., 2017; Lin et al., 2016).

N170 and P600

The amplitude of N170 (also known as the “face potential”) and P600 components varies positively with emotional salience and valence, reaching a maximum in response to salient, negative stimuli (Bentin, Allison, Puce, Perez, & McCarthy, 1996; Holt, Lynn, & Kuperberg, 2009; Schupp et al., 2000).

In our review, we found that dispositional mindfulness was negatively associated with P600 amplitude (Dorjee, Lally, Darrall-Rew, & Thierry, 2015), and an MM intervention was found to be associated with a decrease in N170 amplitude, at least in a clinical population with bipolar disorder (Howells et al., 2014).

Discussion

Reduced reactivity to emotion

Our mindfulness model assumes that the extinction strategy (i.e., active limitation of emotional reactivity) could be supported by increased self-awareness.

ERP evidence showed that the intensity of the emotional response is (a) negatively correlated with dispositional mindfulness (Brown et al., 2013; Dorjee et al., 2015; Lin et al., 2016), (b) is lower in experienced meditators than in naive meditators (Sobolewski et al., 2011), and (c) decreases after MM interventions both in healthy (Lin et al., 2016) and clinical populations (Howells et al., 2014). This negative effect of mindfulness on emotional response supports the notion of extinction in mindful functioning. However, the effect was not observed in the case of a task-induced state of mindfulness in which participants were instructed to pay attention to emotional stimuli with a mindful mind-set. In this case, an increase in emotional response (Egan et al., 2017; Uusberg et al., 2016) or a lack of change was reported (Lin et al., 2016). These mixed findings suggest that outcomes could be a function of the experimental context. In particular, it is possible that naive participants mobilize attentional resources to adopt a mindful mind-set when faced with emotional stimulation; this would account for their higher emotional reactivity associated with a task-induced state of mindfulness.

Emotion regulation and consciousness in mindfulness

In mindfulness, attention focuses on whatever is experienced, including emotions and related physical changes (e.g., increased heart rate, vasomotor flush; Hölzel et al., 2011).

Our model suggests that mindfulness supports the conscious processing of emotional significance and its reevaluation on the basis of reappraisal and extinction strategies. Mindfulness could involve dissociation from emotion, defined in terms of physical change, and the significance given to it. We suggest that the reduced emotional reactivity found in mindfulness could reflect an increased ability to dissociate an emotion from its significance. More than a “reappraisal,” this strategy could be seen as an “appraisal” because no significance is initially attributed to the emotion.

The uncoupling of an emotion from its significance could account for the notion of “acceptance” that contributes to the definition of mindfulness (Kabat-Zinn, 1990, 1994). Acceptance can be defined as an objective, nonreactive lens through which momentary experience is viewed. Regardless of the content of the sensory experience, the individual is encouraged to adopt a mental attitude of acceptance and allow all experiences—even diffcult or stressful ones—to arise and pass without further elaboration, evaluation, or reactivity (Lindsay & Creswell, 2018).

Enhanced body awareness in mindfulness

In our model, improved body awareness may arise from bodily information having better access to consciousness. Our review highlighted that no ERP studies of mindfulness have investigated this topic.

Classically, objective measures of body awareness are peripheral (e.g., heartbeat detection, skin conductance arousal, respiratory tracking, joint position sense, etc.; Treves et al., 2019), but some central signals could be of interest. For instance, the heartbeat-evoked potential is an ERP that occurs about 200 to 600 ms after the cardiac R-wave signal (Schandry, Sparrer, & Weitkunat, 1986). Its amplitude is consistently reported to be a good predictor of individual performance in a heartbeat-detection task (Pollatos, Kirsch, & Schandry, 2005; Pollatos & Schandry, 2004; Schandry et al., 1986). Perception awareness can be evaluated using pN1 and pP1, which reflect sensory awareness and awareness of sensorimotor integration, respectively (Perri et al., 2018). Respiratory movements and postural balance are other interesting candidates because these physical signals exhibit periodic oscillations that make them particularly suited to analyzing time-locked brain activity (Winter, 1995; Zelano et al., 2016).

Future studies should probe these physiological markers to investigate the enhanced body-awareness mechanism through which mindfulness may work.

Furthermore, we suggest that enhanced body awareness in mindfulness could play a major role given that effective body-to-brain communication could foster emotion regulation and cognitive control. The literature suggests that bodily information plays a major role in the way individuals regulate their emotions (Damasio, 1996; Dunn et al., 2010; Magalhaes, Oliveira, Pereira, & Menezes, 2018). Here, we suggest that enhanced body awareness could improve cognitive control by contributing to building the self (i.e., thoughts or emotions) and developing meta-awareness. Studies have revealed that just a third of partial errors⁴ are consciously detected (Rochet, Spieser, Casini, Hasbroucq, & Burle, 2014). Our model suggests that mindfulness could improve the conscious perception of partial errors and support their correction before they turn into overt errors.

What is the “half-life” of mindfulness?

Investigation of mindfulness efficacy and its mechanisms raises the following question: “What is the half-life of mindfulness?”—in analogy to the half-life of a drug. In this section, we discuss the short- (time-limited) or long-term (persistent over time) effects of mindfulness on cognition.

Our review of ERP findings showed that neural features of attention can be detected very early in MM practice, after just a few minutes (Lakey et al., 2011). This quick effect is consistent with the early phases of MM practice that specifically address focused attention (Chiesa et al. 2011). But does the effect persist over time? If not, it could be argued that effects reported after long-term interventions (MBSR, MBCT; Fjorback et al., 2011) could merely be the consequence of the participant’s most recent session.

Note that in most of the reviewed studies, the cognitive task was performed immediately after the mindfulness intervention. Furthermore, when assessing experienced meditators, the interval between the cognitive assessment and the most recent meditative session was not controlled. Consequently, it could be argued that the mindfulness effect reported in experienced meditators could be the result of the session undertaken just before the experiment. Therefore, we suggest that future work should control for the timing of the cognitive task with respect to the mindfulness intervention (or practice for experienced meditators). This precaution should provide a better understanding of the short- or long-term characteristics of the effect of mindfulness.

Note that the point discussed above is slightly different to understanding the effect of mindfulness expertise in terms of length of practice (frequency and duration). ERP evidence suggests that long-term mindfulness practice is positively correlated with improvement in cognitive control (Atchley et al., 2016; Teper & Inzlicht, 2012). Long-term practice is also associated with an inverted u-shaped cerebral activation curve in the attentional network (Brefczynski-Lewis, Lutz, Schaefer, Levinson, & Davidson, 2007). These findings suggest that although the development of mindful cognitive abilities is resource-consuming, mindful functioning is beneficial because it becomes more economical with the level of expertise.

Conclusion

In our work, we aimed to develop a refined model of mindfulness in relation to consciousness. This extended model was evaluated using the existing ERP literature. Our model suggests that mindfulness decreases the threshold of conscious access by supporting attention (Fig. 2). Mindfulness, therefore, facilitates the conscious processing of information that comes from within (body awareness and self-awareness) and outside the body (world awareness).

Fig. 2.

Schematic illustration of the mindfulness model and its relation to consciousness.

In our work, we sought to articulate the cognitive mechanisms associated with mindfulness in relation to each other and took the self-regulation of attention as a starting point. It seems that as bodily information has better access to consciousness, body awareness improves. Our model suggests that these first two mechanisms (self-regulation of attention and enhanced body awareness) contribute to better cognitive control and emotion regulation. We argue that effective body-to-brain communication combined with better access to consciousness support the processing of emotion and its appraisal, thus reducing reactivity. Finally, the individual’s representation of the intended action with respect to the context is improved by the conscious processing of information from within and in the external world, thus supporting the control of ongoing action and its outcomes.

Footnotes

Acknowledgements

The opinions or assertions expressed herein are the private views of the authors and are not to be considered official or to reflect the views of the French Military Health Service.

Transparency

Action Editor: Laura A. King

Editor: Laura A. King

ORCID iD

Charles Verdonk

Notes

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