Mindfulness, cognition, and long-term meditators: Toward a science of

advanced meditation
Sebastian Ehmann1,2,3, Idil Sezer1,4, Isaac N. Treves1,5,6, John D.E. Gabrieli5,6, Matthew D.
Sacchet1,*
1
Meditation Research Program, Department of Psychiatry, Massachusetts General Hospital, Harvard Medical School,
Boston, MA, USA
2
Department of Psychology, University of Arizona, Tucson, AZ, USA
3
Center for Consciousness Studies, University of Arizona, Tucson, AZ, USA
4
Paris Brain Institute, Sorbonne University/CNRS/INSERM, Paris, France
5
Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, 43 Vassar Street, Cambridge,
Massachusetts, 02139, United States of America
6
McGovern Institute for Brain Research, Massachusetts Institute of Technology, 43 Vassar Street, Cambridge,
Massachusetts, 02139, United States of America

Abstract

Mindfulness meditation is a form of mental training rooted in ancient wisdom traditions and is
focused on cultivating a non-judgmental stance toward present-moment awareness. Here, we
synthesize cognitive-behavioral effects in long-term meditators (LTMs) resulting from diverse
and prolonged meditation practices. Preliminary evidence suggests that LTMs exhibit increased
cognitive-sensory integration and decoupling of affective processes, as demonstrated in enhanced
interoceptive awareness, reduced negative affective pain perception, and more rational decision-
making. Additionally, LTMs may experience more emotional neutrality, self-boundary
dissolution, and less normative self-awareness. Neuroimaging findings include increased bottom-
up activation, particularly within the salience network (interoception, pain, affect), and reduced
connectivity between the executive (dorsolateral prefrontal cortex) and salience (dorsal anterior
cingulate cortex) networks (reduced pain). Research also displayed reduced amygdala activation
to fear (reduced negative affect), increased temporoparietal junction activation (pre-reflective
experiential processes, empathy), and altered midline default-mode network activation, which is
associated with emotional neutrality and pre-reflective experiential processes, such as non-
ordinary states of consciousness. Methodological limitations, specifically heterogeneous predictor
variables, restrict the interpretation of trait effects, temporal dynamics in cognitive processing,
and the unique influences of meditative activities. These limitations indicate the need for a unified
research framework and a systematic neurophenomenological investigation of advanced
meditation—through the study of unfolding states, stages, and endpoints in meditative
development. In summary, LTMs display a distinct neurophenomenological gestalt of
mindfulness, wherein meditative expertise is reflected in altered general brain processing,
potentially enhanced cognitive integration, increased cognitive flexibility and self-regulation, and
heightened non-dual awareness—signifying a potentially important form of embodied cognition.

Keywords
mindfulness; meditation; long-term meditators; advanced meditation; cognition; emotion regulation;
awareness.
*
Corresponding author address: Meditation Research Program, Department of Psychiatry, Massachusetts General Hospital, Harvard Medical School,
Boston, MA, USA, [email protected], (M. D. Sacchet).

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1. Introduction

1.1. Meditation and Mindfulness

Meditation is a fundamental capacity of the human mind and has been practiced for thousands

of years in various contemplative branches of Hinduism, Buddhism, and other traditions. The

roots of mindfulness meditation can be traced back to the Satipatthana, one of the prominent

Buddhist meditation systems (Bodhi, 2011). It is a part of the Pāli Canon, the oldest Buddhist

scriptures to survive time entirely. To practice according to the Satipatthana, two factors are

emphasized, sati and sampajañña, translating from Pali, the liturgical language of Theravada

Buddhism, into mindfulness and clear comprehension (Sharf, 2014). In Buddhism, mindfulness is

understood as ‘lucid awareness,’ which, if cultivated with clear comprehension, contributes to the

development of insight into the ‘nature of how things are’ (Bodhi, 2011). The intention behind

diligently practicing mindfulness meditation is to cultivate the core mental faculties involved in

insight (Sugunasiri, 2008), gradually reducing unwholesome mental actions and, in turn, suffering

(Ingram, 2018). These soteriological origins of mindfulness meditation paved the way for the

development of modern scientific definitions that extracted mindfulness from its traditional

context. Another notable definition describes mindfulness as the intentional and nonjudgmental

act of paying attention in the present moment (Kabat-Zinn, 1994). Furthermore, Kabat-Zinn

developed a treatment method known as Mindfulness-Based Stress Reduction (MBSR), now one

of the most popular approaches to training in modern secular mindfulness (Kabat-Zinn, 2013).

Since then, numerous mindfulness-based programs have been developed to address clinical

conditions, including Mindfulness-Based Cognitive Therapy (MBCT; Segal et al., 2013),

Mindfulness-Based Relapse Prevention (MBRP; Witkiewitz et al., 2013), and Mindfulness-

Oriented Recovery Enhancement (MORE; Garland, 2013). Additionally, Dialectal Behavioral

Therapy (DBT) and Acceptance and Commitment Therapy (ACT) are also based on mindfulness

and have been shown to be effective interventions for borderline personality disorder, chronic

pain, tinnitus, and possibly depression, psychotic symptoms, obsessive-compulsive disorder,

anxiety, substance abuse, and stress (Ost, 2014; Panos et al., 2014). Collectively, there is strong

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evidence that mindfulness practices can be used to treat substance abuse (Goldberg et al., 2018;

Priddy et al., 2018), depression (Goldberg et al., 2018; Hofmann et al., 2010; Hoge et al., 2023;

Khoury et al., 2013; Piet & Hougaard, 2011; Strauss et al., 2023), anxiety (Hofmann et al., 2010;

Khoury et al., 2013), stress (Khoury et al., 2013), pain (Goldberg et al., 2018), post-traumatic

stress (PTSD; Boyd et al., 2018), attention-deficit/hyperactivation disorder (ADHD; Poissant et

al., 2019), eating disorder (Wanden-Berghe et al., 2011), and smoking (Goldberg et al., 2018).

Mindfulness has also become popular outside of the clinic, as practices are now widely used for

alleviating emotional distress and increasing overall well-being (Pepping et al., 2016). As the

evidence for transdiagnostic clinical benefits of mindfulness meditation has emerged, there is

renewed interest in mindfulness mechanisms, specifically as they relate to more advanced

practitioners (Sacchet et al., 2024). In this review, we consider behavioral changes and

neurocognitive mechanisms, leveraging a base of evidence from long-term meditators (LTMs).

To set the stage for a synthesis of evidence, we first discuss cognition and how it is theoretically

related to mindfulness meditation. Second, we explore the study of LTMs as prime examples of

how repeated states of mindfulness lead to meaningful trait differences. Lastly, we introduce a

distinction between advanced meditators and LTMs, which provides new explanatory frameworks

for the deep end of cognitive meditative development and its concomitant brain mechanisms.

1.2. Cognition and Mindfulness

Cognition is a multifaceted construct comprised of various domains, including sensation,

perception, motor skills, attention, memory, executive functioning, processing speed, and

language (Harvey, 2019). Its evolutionary development has been closely tied to the expansion and

structural organization of ‘association cortices,’ which have likely been subject to strong selective

pressures during recent hominid evolution (Yeo et al., 2011). These cortices form the bulk of the

cerebral cortex, and the substantial expansion of the parietal region is proposed as a pivotal factor

in molding the distinctive cognitive abilities distinguishing humans from other primates (Bruner

et al., 2014; Bruner & Lozano, 2015). Psychophysiological conceptualizations of cognition have

evolved from signal computations at network nodes implemented by specific neurons and circuits

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(Sherringtonian view) to distributed computations and transformations within representational

spaces by neural populations, making specific neuron details secondary (Hopfieldian view;

Barack & Krakauer, 2021; Hopfield, 1982; Hopfield, 1984; Hopfield & Tank, 1986; Sherrington,

1906). According to Bayne and colleagues, the overarching characteristics of cognition entail:

1) The systematic recombination of representation to generate concepts and models.

2) A spectrum of complexity ranging from logical reasoning to adaptive information

management, incorporating perceptual inferences and flexible thinking

3) The processing of sensory information to direct behavior through intricate neural

mechanisms. (Bayne et al., 2019)

The Free Energy Principle (FEP), a well-established overarching brain theory, reflects these

characteristics, positing that cognitive agents perceive their environments through a predictive

process, which refines future state predictions by minimizing prediction errors through neural

computations (Friston, 2010; Bubic et al., 2010). In this process, top-down predictions of sensory

states are reconciled with incoming bottom-up sensory data. The cognitive agent can either update

internal models constituted of higher-level priors, such as the self (Friston, 2018), or act on the

inferences by extending cognition beyond internal processes onto the external environment

(‘active inference’; Constant et al., 2022). In mindfulness meditation, mitigated physical

activation and the voluntary modulation of attention increases present-moment awareness by

reducing the weighting of higher-order mental predictions in favor of bottom-up sensory

information flow (Lutz et al., 2019). Growing evidence suggests that the brain hierarchically

organizes in progressive levels of abstraction, with higher-order processes being more extended in

time (Friston, 2008; Taylor et al., 2015). Meditation is thus hypothesized to gradually minimize

prolonged temporal processing (Laukkonen & Slagter, 2021), with different practices enabling

more effective minimization and, consequently, deconstruction of habitual self-processes (Dahl et

al., 2015; Friston, 2018; Laukkonen & Slagter, 2021). Other prominent cognitive theories, such as

enactive theory, were directly influenced by contemplative mindfulness meditation traditions and

aimed to bridge the gap between the first-person experience of living cognition and its third-

person objective investigation (Varela et al., 1992). Enaction distinguishes itself from previous

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theories by defining cognition as a non-representational, deeply embodied, and relational process

that enacts a world of significance through constant sense-making (Thompson, 2007; Varela et

al., 1992). Despite claims that the FEP framework integrates enaction, enactive scholars argue

that key concepts like sense-making, agency, and environmental relations are misunderstood and

incompatible with FEP (Di Paolo et al., 2022).

The growing prominence of mindfulness-based practices has shifted the scientific focus from

whether mindfulness is effective to exploring its mental and neural mechanisms. Initial

contemplative research into the underlying mechanisms posited 1) intention, 2) attention, and 3)

attitude, collectively affording 4) re-perception as crucial factors of mindfulness (Shapiro et al.,

2006). Subsequent work advocates a self-processing lens, in which mindfulness meditation leads

to enhanced self-awareness, self-regulation, and self-transcendence by improving constitutive

cognitive processes and associated neural substrates (Vago & Silbersweig, 2012). These

improvements include enhanced meta-cognition and attention regulation, body awareness,

emotion regulation, and shifts in self-perspectives (Coffey et al., 2010; Dorjee, 2016; Hölzel et al.,

2011; Tang et al., 2015; Vago & Silbersweig, 2012). These changes are thought to result in two

transdiagnostic key meta-mechanisms: 1) a shift toward a more embodied self, and 2) increased

dynamism in self-patterns due to enhanced cognitive flexibility (Christoff et al., 2011; Giommi et

al., 2023; Moore & Malinowski, 2009) Notably, the mindfulness mechanisms behind these

changes may be similar for both non-meditators and meditators (Burzler et al., 2019).

1.3. The Study of Long-term Meditators

Due to their extensive mental training, LTMs are a valuable group for studying altered

cognitive functions as related to meditation. As this population may have the highest level of

development in their specific training domains, they, in theory, allow for the most precise insight

and strongest effect in neurocognitive changes. This could enable neuroscientists to more readily

identify the effects of mental training on neurocognitive processes, such as investigating process-

specific learning (Slagter et al., 2011). However, it is important to note that individuals motivated

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to engage in long-term meditation may inherently differ in cognitive and brain characteristics

from the general population, potentially limiting the generalizability of findings.

Sustainable long-term changes have been hypothesized to occur through a natural progression

from momentary cognitive states to persistent cognitive traits and may be dependent on the

expertise of the meditator (Goleman & Davidson, 2017). States and traits differ in their temporal

scale and locus of origination: states are brief and influenced mainly by external circumstances,

while traits are persistent, durable, and internally caused (Chaplin et al., 1988). During

mindfulness meditation, the practitioner acutely experiences deeper mindfulness, which, with

continuous effort, may thus lead to changes in trait mindfulness (Atasoy et al., 2023; Bauer et al.,

2019; Kiken et al., 2015). Trait mindfulness describes the dispositional mindful aptitude of a

person and is usually measured through self-report questionnaires (Baer et al., 2004; Baer et al.,

2008; MacKillop & Anderson, 2007; Walach et al., 2006). Studies show that changes in state

mindfulness predict trait mindfulness changes after interventions (Kiken et al., 2015), and that

meditation experience strongly correlates with dispositional mindfulness scores (Vinchurkar et al.,

2014). However, trait mindfulness may not relate to attentional performance in cognitive tasks

(Quickel et al., 2014), and some trait assessments fail to distinguish between experienced

Buddhist monks and college students (Christopher et al., 2009).

Generally, mindfulness-based programs and continuous personal practice may provide

positive effects on well-being; however, these effects may be limited in both their duration and

magnitude. Persistent deep psychological transformations may only be acquired with extensive,

consistent practice. Various traditions using mindfulness meditation in a soteriological framework

provide evidence for these types of drastic changes through first-person accounts and emerging

scientific findings (Berkovich-Ohana et al., 2013, 2017; Dor-Ziderman et al., 2016; Goleman &

Davidson, 2017; Hagerty et al., 2013). A notable conceptual distinction may, therefore, lie

between LTMs and what we refer to as advanced meditators. The latter are characterized not just

by extensive practice experience—the quantifiable time spent meditating—but by a higher level

of meditative development, marked by access to deeper phenomenological states and traits.

Although they are likely correlated in some instances, practice experience and meditative

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development may diverge and even follow non-linear dynamics (Galante et al., 2023; Sacchet et

al., 2024). Indeed, we propose the study of advanced meditation, states, stages, and traits that

unfold with mastery and time (Sacchet et al., 2024). Advanced meditation includes meditative

development or the developmental trajectory of states and stages that unfold, and also meditative

endpoints that are landmark outcomes of advanced meditation (Galante et al., 2023; Sacchet et al.,

2024). Our recent research within this domain has uncovered new connections between changes

in consciousness observed in advanced meditation states and related neural mechanisms

(Chowdhury et al., 2023; Ganesan et al., 2024; Yang et al., 2023, 2024; van Lutterveld et al.,

2024; Wright et al., 2024). Researching advanced meditators may thus give insight into the

process of meditative development, which promises to inform new clinical and non-clinical

approaches to fostering well-being and treating psychiatric illness.

1.3.1. Classifying Mindfulness and its Practices and States

A major challenge in the scientific study of mindfulness meditation, especially when

considering advanced states and stages, has been effectively categorizing different meditation

techniques. Previous research proposed a “phenomenological matrix” comprising three key

dimensions: 1) object orientation, 2) de-reification, and 3) meta-awareness (Lutz et al., 2015). We

recently introduced a broader, more comprehensive framework leveraging an ‘activation-based

phenomenological classification system,’ which methodically integrates meditative practices with

phenomenology. This approach not only provides a more comprehensive and precise

understanding of the meditator's behaviors and subjective experiences (Sparby & Sacchet, 2022)

but also significantly contributes to clarifying the multidimensional nature of mindfulness, its

application in clinical interventions, and the potential reasons for adverse outcomes in specific

contexts. In our framework, we identify four core activities of meditative techniques, namely, 1)

focusing, 2) releasing, 3) imagining, and 4) moving in relation to an object of meditation (which

includes fields of experiencing). Furthermore, these activities unify into the meditative activities

of observing, producing, and awareness, ultimately unified in the meta-activation of awareness of

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awareness. Our framework is the first to comprehensively classify the various meditation

techniques based on empirical descriptions (Matko et al., 2021) and contemplative traditions

(Anālayo, 2020a)—and thus represents progress in reducing semantic ambiguity in the definition

and meaning of mindfulness and its empirical consequences.

The second integral feature of classifying meditation techniques encompasses the objects of

meditation, usually separated into the different senses: 1) thought, 2) sound, 3) sight, 4) taste, 5)

smell, and 6) bodily sensations. They can be considered the content of one’s awareness, while the

meditative activation describes how one relates to the content. For example, In Theravadan

Buddhist samatha meditation, meditators focus on a specific object, like the breath at the nostrils

(Tiwari, 1988). With increasing expertise, meditators may reach a contentless state, referred to as

a ‘pure consciousness event’ (Woods et al., 2020; Woods et al., 2022), marked by a decrease in

typical subject-object distinctions, which has been termed self-transcendence or unity (Aron et al.,

1992). Meditators can attain various peak states, contingent on their practice and object. For

instance, concentration-focused practices like samatha can lead to samādhi, signifying ‘the state

of being firmly fixed’ and characterized by mental singularity (Sraman, 2002). In so-called pure

consciousness events, the individual still experiences a slight subject-object dichotomy, as

consciousness has become the meditative object. The last step to unification is total self-

awareness, where awareness is aware of itself, completely dissolving the subject/object

dichotomy into non-duality (Josipovic, 2021; Sparby & Sacchet, 2022). ‘Non-dual awareness’

characterizes sustained and non-propositional meta-awareness, not centered on a specific object

but on the intentional relationship itself (Dunne et al., 2019; Lutz et al., 2007).

To enable a more nuanced understanding of mindfulness, one comprehensive system

describes it as a practice-based skill acquisition process encompassing three distinct but

interrelated domains: 1) concentration, 2) sensory clarity, and 3) equanimity (Young, 2016). The

goal is to enhance baseline abilities in these areas. Combined with Sparby & Sacchet’s (2022)

model, which accommodates meditative activities and related phenomenology, this allows for the

systematic investigation of mindfulness meditation states and stages. It aligns with recent

contemplative models integrating cognitive-affective neuroscience, psychological processes, and

8
phenomenology. Dorjee (2016) suggests that continuous mindfulness practice leads to de-

reification of mental phenomena and higher existential awareness. The Mindfulness-to-Meaning

theory expands on this, proposing a spiral model where mindfulness disrupts negative mental

frames, promoting positive affect and self-transcendence, eventually leading to non-dual

awareness (Garland et al., 2015; Garland & Fredrickson, 2019). These self-transcendent states

hold substantial clinical value and have already been applied in studies on addiction and pain

(Hanley et al., 2018; Parisi et al., 2022; Sacchet et al., 2024). The following section addresses the

need for a unified framework to support systematic scientific exploration of LTMs.

1.4. An Empirical Framework for Mindfulness Research

Since 2006, contemplative science has grown exponentially, with over 16,000 scientific

articles published since 1966 (Baminiwatta & Solangaarachchi, 2021). This growth underscores

the crucial need for a unifying framework for empirical investigation. Challenges include

inconsistencies in defining mindfulness and limitations in scientific methods and research designs

(Van Dam et al., 2018), as well as the need to address advanced meditative states and stages

(Galante et al., 2023). Semantic inconsistencies and construct ambiguity have led to divergent

mindfulness self-report instruments, affecting definitions and interpretations (Bergomi et al.,

2013; Grossman & Van Dam, 2011). Additionally, neglecting advanced meditation has resulted in

overlooking unique benefits and adverse events associated with meditative development (Cebolla

et al., 2017; Galante et al., 2023; Goldberg et al., 2020; Lindahl et al., 2017; Sacchet et al., 2024;

Schlosser et al., 2019).

To address these limitations, we recently established a framework for the empirical

investigation of mindfulness and, in particular, meditative development (Galante et al., 2023).

This model emphasized the need for more longitudinal, interdisciplinary, transparent, and

ontologically agnostic research. It also advocated for integrating contemplative scholastic work,

such as traditional stages of insight (Grabovac, 2015), with context-sensitive multi-method

research methodologies to confront the growing body of research indicating that mindfulness can

cause adverse or challenging effects (Aizik-Reebs et al., 2021; Lambert et al., 2021), especially if

9
practiced alone, in secular contexts, and without developmental frameworks (Cebolla et al., 2017;

Schlosser et al., 2019). Additionally, collaborating directly with meditation practitioners could

enhance researchers’ understanding of how to best study meditation, as it enables them to

systematically assess factors such as practitioners’ meditation experience and history, including

specific practicing methods (Matko et al., 2021), experiential patterns, interactions between

meditation effects and individual meditator characteristics, meditation goals, and the cultural and

social environments of practice (Sparby & Sacchet, 2022).

In summary, both introduced models (Galante et al., 2023; Sparby & Sacchet, 2022) enable a

more precise understanding of altered cognitive processes by allowing the capturing of perceptual

adaptations and connecting those to unique behavioral patterns and brain network alterations—a

crucial step in advancing the scientific progress of neurophenomenological mindfulness

meditation research.

1.5. The Current Review

In the following, we present a comprehensive review of the relations between LTMs,

behavior, and cognition, along with a selective overview of related neuroscience research.

Relevant studies of LTMs were identified by requiring an average of at least 1,500 hours of

meditation within the respective group. This work aspires to connect emerging theoretical

frameworks on mindfulness measurement, meditative development, and cognitive mechanisms to

understand the phenotype of higher-end practitioners. Subsequently, it uses this framework to

synthesize the empirical findings of this population. The review is separated into four main

sections encompassing different but interpenetrating cognitive functions, namely, perception,

emotional processing, higher-order functions, and non-ordinary states of consciousness. Sample

tasks from each section are represented in Figure 1. In the discussion, we will address

methodological limitations, summarize and contextualize the observed behavioral trait effects,

and synthesize findings within an overarching dynamical self-processing model.

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Figure 1

Sample cognitive-behavioral tasks employed within the different domains.

Note. Five exemplary tasks used in the study: (a) Cardiac interoceptive awareness was assessed by having
participants mark where they felt their heartbeat on a body map, generating proportional maps based on
these responses (Khalsa et al., 2020). (b) Graded thermal stimulation induced moderate pain, which
participants rated on a point scale (Lutz et al., 2013; Grant & Rainville, 2009; Grant et al., 2010, 2011;
Perlman et al., 2010; Zorn et al., 2020). (c) Participants viewed emotional pictures and rated their
subsequent emotional responses (Chen et al., 2018; Taylor et al., 2011). (d) The Ultimatum Game assessed
economic decision-making, with participants deciding whether to accept or reject various offers from
human and computer partners (Kirk et al., 2018). (e) In the time production task, participants pressed a
button to match target durations, evaluating their time estimation ability with eyes closed (Berkovich-
Ohana et al., 2011).

2. Cognitive Functions in Long-term Meditators

2.1. Somatosensory Perception

Attention involves the process of focusing on relevant information, and perception is the

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cognitive process by which sensory information is interpreted and transformed into a

representation that captures the context of the scene (Albright, 2015). This transformative process

of perception involves neuronal computation that assigns semantic interpretations to incoming

signals, leveraging contextual information to enhance the understanding and meaning of sensory

input. Consequently, perception can be consistently broadened and enriched through systematic

conceptual development (Newen, 2017). This aligns with a predictive processing account, in

which moment-to-moment conscious perception is shaped through the constant updating of

higher-level priors or through action, while also accommodating higher-level features like

emotions and person impressions (Binder et al., 2017; Kahl & Kopp, 2018). There is some

resonance between neuroscientific definitions and Buddhist understanding of perception. In Pali,

Saññā (perception) denotes the discernment of objects from sense signals (Wright et al., 2023). It

constitutes an integral element of the Buddhist doctrine of ‘The Five Aggregates of Clinging,’

which elucidates how people turn neutral sense information into experiences of suffering and

distress. Consequently, changes in ordinary perception are at the core of Buddhist soteriological

frameworks and are viewed as a pivotal mechanism within mindfulness practices (Shapiro et al.,

2006). Substantial perceptual alterations, including those experienced in advanced meditation,

might predominantly manifest in adept meditators or under intense practice regimens, an aspect

that has garnered limited scientific exploration. Qualitative investigations involving accomplished

Burmese meditators revealed four modifications in perception: 1) heightened perceptual

sensitivity, 2) diminished subject-object dichotomous perception, 3) recognition of interconnected

perceptual processes, and 4) nonconceptual perception (Full et al., 2013). These shifts toward

non-conceptual cognition are suggested to facilitate insights into a deeper reality (Bodhi, 2011).

However, it is important to distinguish this from accessing an ‘ultimate reality’ (Kant, 1787/1998)

or overcoming human perceptual illusions (Carbon, 2014). Instead, it represents a progression

toward a less biased mental state, characterized by the disentanglement from habitual cognitive

discriminations (i.e., biased attentional sampling), thus allowing more resources for direct sensory

discrimination (Thompson, 2023). Indeed, intensive meditation training has been linked to

enhanced perceptual discrimination, afforded by increased vigilance and perceptual sensitivity,

12
leading to reduced cognitive load and increased sustained attention (MacLean et al., 2010).

Numerous studies have examined perceptual changes in LTMS in both clinical and non-clinical

contexts. We will next describe two studies on interoceptive awareness and six studies on pain

perception.

2.1.1. Interoceptive Awareness

Recent studies proposed a neurocognitive hierarchy of self-processing, in which body-

environment information follows a gradual integration from interoceptive processing to mental

self-processing (Qin et al., 2020). For example, research has demonstrated that interoceptive

signals can modulate the perception of the external world (exteroception), as experienced through

body surfaces (somatosensory perception; Al et al., 2021). Interoceptive awareness is the

conscious perception of bodily signals that collectively create a physiological sense of self,

encompassing sensations like breathing, heartbeat, hunger cues, and the physical manifestations

linked to emotions (Barrett et al., 2004; Cameron, 2001; Craig, 2002; Vaitl, 1996). A critical

neural substrate for interoception is the insula (Qin et al., 2020), with other implicated regions

including the anterior cingulate cortex (ACC), posterior cingulate cortex (PCC), and medial

prefrontal cortex (mPFC; Treves et al., 2024) and temporoparietal junction (TPJ; Hölzel et al.,

2011).

The cultivation of interoceptive awareness may be a foundational mechanism in vipassanā or

mindfulness meditation with body sensations often used as the objects of meditation in various

practice traditions. For instance, a practice in the vipassanā tradition guides meditators to

methodically scan their bodies, attentively observing bodily sensations without judgment or the

intent to modify them. Within the kāyagatāsati sutta, mindfulness is cultivated by practicing body

awareness with several exercises, such as noting postural or movement differences, or meditating

upon the bodies’ constituents (Anālayo, 2020a; Anālayo, 2020b). In the yogic traditions, asanas

guide the practitioners’ bodies and awareness into specific postures to create relaxation for further

meditative practices (Iyengar, 1995). Thus far, mindfulness meditation has been shown to

influence the insula and TPJ (Gibson, 2019; Haase et al., 2016; Hölzel et al., 2011; Todd & Aspel,

13
2022). Research indicates that increased thickness of the insular cortex may mediate the

heightened body awareness observed in mindful individuals (Friedel et al., 2015; Treves et al.,

2019, 2024). Furthermore, the observe facet of the FFMQ self-report questionnaire may

specifically be sensitive to interoceptive changes in mindful individuals (Hölzel et al., 2011). In

this context, mindfulness meditation has been suggested as a potential intervention to assist with

affective and psychosomatic disorders that encompass interoceptive distortions within their

underlying pathology (Farb et al., 2015).

To test the effects of meditation on interoception, Fox and colleagues enrolled 38 meditators

with an average of 11 years and 2,051 hours of meditation experience, spanning from 1 to 15,000

hours (Fox et al., 2012). The researchers divided the sample into four quartiles, with the bottom

and top quartiles representing novice (average meditation experience = 28 hours, SD = 24 hours)

and LTMs (average meditation experience = 7231 hours, SD = 4410 hours), respectively.

Interoceptive accuracy was assessed by correlating meditators' subjective ratings of bodily

sensations during a body-scan meditation with objective measures of tactile sensitivity.

Participants rated sensations across 20 body regions, which were compared to a composite

Somatic Sensitivity Rank (SSR). The SSR combined psychophysical two-point discrimination

thresholds and cortical representation in the primary somatosensory cortex, derived from literature

and validated through neurosurgical and neuroimaging data. The correlation between subjective

ratings and SSR quantified participants’ ability to accurately perceive and report bodily

sensations. Participants were familiar with the body-scan meditation type, but their experience

varied widely and represented only a fraction of their overall meditation practice (less than 10%).

Results included a large difference in trait introspective accuracy between long-term and beginner

meditators. Furthermore, total body-scan meditation experience, meditation experience, and

average practice amount per month predicted introspective accuracy. There was a dose-response

relationship between meditation experience and interoception. Notably, the significance of this

relationship remained even after accounting for the participants’ body-scan meditation experience

in their overall meditation practice. Nevertheless, this study design precluded the exploration of

the relationship between subjective and objective measures of somatosensory awareness,

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underscoring the need for future research to examine changes in subjective intero-exteroceptive

awareness and brain topography in LTMs.

Conflicting evidence for increases in interoceptive awareness was reported by Khalsa and

colleagues for cardiac interoceptive accuracy during rest (Khalsa et al., 2008) and physiological

arousal (Khalsa et al., 2020). In their first study, participants performed a heartbeat detection task

that involved presenting auditory tones to participants. These tones were either simultaneous or

non-simultaneous with the participants’ actual heartbeats. Participants had to judge whether each

tone coincided with their perceived heartbeat sensation, without being allowed to take their

pulse. To address the methodological limitations of their previous studies, in the second study,

participants received blinded infusions of isoproterenol—a beta-adrenergic drug similar to

adrenaline—at various doses and saline while rating their internal body sensations in real-time

using a dial, followed by retrospective ratings of physical sensations and emotional experiences.

Interoceptive detection accuracy was assessed by calculating cross-correlations between

participants’ dial ratings and their actual heart rate changes during each infusion, with higher

correlations indicating greater accuracy in perceiving bodily sensations. This approach aimed to

enhance physiological arousal by stimulating the sympathetic nervous system and intensifying

internal sensations, potentially counteracting the low detection rates observed in quiescent resting

state scenarios. Cardiac spatial interoceptive awareness was assessed using drawings on a two-

dimensional paper manikin, where participants indicated areas of felt heartbeat sensations. These

drawings were digitized and converted into proportional body maps for specific doses, with each

pixel value representing the proportion of participants reporting sensation in that area. While not

explicitly stated, accuracy was likely inferred from the consistency and anatomical relevance of

reported sensations across participants. Other assessments included self-reported ratings and

electrocardiogram (EKG). The researchers recruited 15 LTMs and 15 matched non-meditators.

The meditators averaged 10.8 years and 4,947 hours of meditation experience, with 19 days

completed in a retreat setting. Eleven meditators practiced vipassanā, while four engaged in

Kundalini meditation, which combines active imagination, breathing techniques, mantras, and

passive observation and release, suggesting familiarity with interoceptive meditations for most

15
meditators. No between-group differences were found for cardiac interoceptive awareness. These

outcomes were further validated through a subsequent meta-analysis, which explored cardiac

interoceptive awareness between meditators and non-meditators from various traditions and

indicated that meditation may not substantially impact cardiac interoceptive awareness,

consolidating the researchers' conclusions.

In summary, preliminary research might indicate improved interoceptive awareness in LTMs

compared to non-meditating controls across various meditation practices, including body-scan

meditation. Nevertheless, these effects are not seen in cardiac interoceptive awareness,

underscoring the need for further investigation to establish the broader applicability of these

findings. Additionally, more longitudinal studies directly comparing different meditation

techniques are warranted.

2.1.2. Pain Perception

Pain is a distressing sensory and emotional phenomenon intricately intertwined with various

cognitive domains, sociocultural influences, and biological determinants (Binder et al., 2011;

Khera et al., 2021; Symbaluk et al., 1997; Ossipov et al., 2010) and processed in the ‘pain matrix,’

including the anterior cingulate cortex (ACC), insula, thalamus, primary (S1) and second

somatosensory (S2) cortices, amygdala, periaqueductal gray, and prefrontal cortices (PFC; Ingvar,

1999; Rainville, 2002; Tracey & Mantyh, 2007). Recent resting-state functional connectivity

(rsFC) research has identified a distinct pain network comprising the dorsolateral prefrontal cortex

(dlPFC), anterior insula, thalamus, precuneus, and ACC (Lee et al., 2019)—which strongly

overlaps with the salience network (SN) due to the high salience of painful stimuli (Borsook et al.,

2013). Pain perception is complex, as sensory pain, a physiological response to harmful stimuli

that can be characterized and localized, is reciprocally intertwined with a cognitive-affective

dimension, which reflects the emotional and motivational appraisal of the stimuli (Grant, 2014;

Price, 2002). Sensory-discriminative aspects of pain involve the somatosensory cortices,

thalamus, and posterior insula (Coghill et al., 1999), while affective-motivational components are

processed in the dorsal ACC and anterior insula (Rainville et al., 1997), and cognitive

16
components are localized in the PFC (Strigo et al., 2002). Compared to a neuroscientific

perspective, Buddhism conceives of pain within the broader context of suffering, wherein sensory

experiences are modulated by approach and avoidance tendencies through the grasping of more

pleasant goal states (Thompson, 2023; Waikakul & Waikakul, 2016). Meditative training aims to

reduce grasping by transforming cognitive processes to re-perceive sensory input less

distressingly (Waikakul & Waikakul, 2016). Functionally, meditation introduces a divide

between the sensory experience and its cognitive interpretation through acceptance, non-

judgment, and decentering, often encapsulated within the overarching construct of equanimity

(Desbordes et al., 2014; Soler et al., 2021). Contemplative neuroscience has demonstrated that

mindfulness meditation-induced pain relief is associated with increased activation in sensory and

effective core pain areas such as the ACC, S2, and insula, reduced activation in the thalamus,

amygdala, PCC, periaqueductal gray, and hippocampus (Tang et al., 2015; Wipplinger et al.,

2023), as well as enhanced cortico-thalamic regulation (Riegner et al., 2023) and increased

within-SN connectivity between the anterior insula and dorsal ACC (Sezer et al., 2022).

Crucially, research thus far has provided preliminary evidence that pain reductions between

novice and LTMs differ, with experts more effectively reducing pain unpleasantness by

deactivating appraisal-related pain regions, such as the PFC, whereas novices may rely on

cognitive control strategies to reduce pain, as they are unable to selectively modulate affective

and sensory domains (Grant, 2014; Wipplinger et al., 2023).

To assess the effects of meditation on pain perception, Grant and colleagues assembled a

cohort comprising 19 long-term Zen meditators and 20 age- and gender-matched controls (Grant

et al., 2010). The meditators averaged 6,404 hours of mindfulness meditation experience, ranging

from 1,229 to 45,000 hours. Employing computer-controlled thermal stimulation, the researchers

induced a moderate pain level, prompting both groups to convey their discomfort on a 10-point

pain rating scale. The outcomes revealed a significant decrease in pain sensitivity among Zen

meditators, as indicated by a substantial two-degree Celsius difference during the hot stimulation

for the same pain magnitude. These results find support in previous and subsequent studies

employing similar research methodologies, consistently observing reduced baseline pain

17
sensitivity and analgesic effects in Zen meditators (Grant & Rainville, 2009; Grant et al., 2011).

Notably, in their earlier research, a focused observing practice heightened pain sensations among

control participants but had no impact on LTMs. Conversely, an observe and release practice

diminished pain in the Zen group without affecting the control group (Grant & Rainville, 2009).

In their earlier work, meditation hours successfully predicted reductions in pain sensitivity (Grant

& Rainville, 2009). However, this effect was not replicated in their subsequent study (Grant et al.,

2010), even though half of the sample was the same. It is still unclear whether there were

additional overlaps in their most recent study (Grant et al., 2011).

To understand the effect of different meditation practices on pain, Perlman and colleagues

enrolled 9 LTMs and 10 novices (Perlman et al., 2010). The LTMs practiced Tibetan Buddhism

and ranged in experience from 10,000 to 45,000 hours. Each subject participated in 8 blocks of 4

trials of focused attention (FA) or open presence (OP) meditation. Open-presence (OP) meditation

is a non-dual and advanced form of open monitoring (OM), and thus, similarly, a release and

observing meditation. Whereas in OM, the monitoring of the present-moment content of

experience might still contain a duality between meditator and content, OP emphasizes a

completely transparent mind in which content is just reflexively aware of itself without a reified

subject (Josipovic, 2010, 2019, 2021; Perlman et al., 2010). FA meditation instructions prompted

the subjects to use a small object other than their pain as an attentional focal point. The

participants were instructed to practice the respective technique for 45 seconds before a graded

thermal pain stimulus was induced. Both groups continued their meditation throughout the task

and were probed regularly for pain ratings. A significant interaction effect was observed between

practice type and rating type within the groups. This interaction revealed a notable decrease in

pain sensitivity and unpleasantness among LTMs practicing OP meditation, with the latter being

the primary contributing factor.

Applying a comparable research design, the team validated their results by replicating the

study in a group of 14 extensively trained Tibetan Buddhist practitioners (Lutz et al., 2013). These

individuals had accrued a minimum of 10,000 hours and an average of 27,000 hours of experience

in OP meditation. This group was then compared to a matched control group of 14 individuals

18
without prior meditation experience. Analysis revealed almost identical pain intensities between

groups but diminished unpleasantness in the LTMs. The supplementary analysis uncovered an

interaction effect between the practice and rating type (intensity and unpleasantness) for the

experts but not the control group, showing lower unpleasantness in the expert OP compared to the

FA group. Still, the expert OP generally outperformed the expert FA group by demonstrating both

reduced pain intensity and unpleasantness ratings.

Consistent with previous findings, Zorn et al. found identical reductions in unpleasantness but

not intensity after an observe and release (Kagyu) practice in 27 experienced Tibetan Buddhist

meditators (average meditation experience = 41,357 hours; Zorn et al., 2020). In this cross-

sectional study, the researchers implemented a design with the intent to enhance cognitive-

affective aspects of pain through modulating pain anticipation and stimulus length. The LTMs

also observed a significant decrease in pain catastrophizing compared to the novices. This

reduction in catastrophizing was found to predict lowered sensory-affective connectivity during

extended pain stimuli and decreased pain intensity during brief exposures. Subsequent analyses

unveiled heightened cognitive defusion—an experiential distancing from thoughts and

emotions—as a central psychological mechanism driving the pain-reducing effects by meditation

(Zorn et al., 2021). This finding corroborates the ideas introduced earlier in this section, which

explain a reduced negative cognitive-affective appraisal in LTMs compared to beginners.

Neuro-mechanistic investigations into the pain processing of LTMs corroborate the previous

findings and point to a mindfulness-induced dis-association of the emotional-cognitive-evaluative

and sensory-discriminant components of pain (Grant et al., 2011; Grant, 2014; Thompson, 2023;

Wipplinger et al., 2023). This is reflected in increased pain-related activation in ACC, thalamus,

and insula, and decreased pain-related activation in prefrontal cortices, amygdala, and the

hypothalamus in LTMs compared to matched naïve controls, with more meditation experience

correlating with reductions in these areas and the most significant reductions being displayed in

prefrontal regions (dlPFC and the mPFC/orbitofrontal cortex (OFC; Grant et al., 2011)).

Additionally, earlier research found greater gray matter in pain-related regions (right dACC and

bilateral S2) in LTMs compared to naïve controls. Significant correlations were observed between

19
the number of years of meditation and grey matter thickness in the ACC, as well as between

meditation hours and thickness in the lower leg area and right-hand areas of S1 (Grant et al.,

2010). These findings suggest a generally increased somatosensory perception (S1, S2) and an

enhanced capacity to process affective pain components. Despite also finding significant negative

associations between pain sensitivity and cortical thickness of various pain-related regions (right

dACC, S2, insula, hippocampus), no interaction effect with the groups was observed, suggesting a

general relationship between cortical thickness and pain sensitivity (Grant et al., 2010). Brain

activation correlations indicate a decoupling of executive and pain-related brain regions during

heat-induced pain, specifically between the right dlPFC and dACC, which strongly predicted

reduced pain sensitivity in LTMs (Grant et al., 2011). The enhancement in the sensory-

discriminative aspects of pain processing has been suggested not to signify heightened pain

perception, but rather an enhanced capability to identify potentially threatening, salient bodily

stimuli (Legrain et al., 2011). Nevertheless, the cognitive/sensory information integration

component of pain may still be heightened in meditators, with a selective decoupling of emotional

appraisal, suggesting an increased differential integration of sensory/cognitive and affective pain

components, potentially leading to a heightened but altered pain perception.

Converging results correlated reduced pain unpleasantness in LTMs with enhanced activation

in the salience network (SN), specifically the dorsal anterior insula and anterior mid-cingulate

cortex (aMCC), during the processing of painful heat stimuli. Baseline testing showed the

opposite pattern, with decreased SN and amygdala activation, potentially demonstrating reduced

pain anticipation and negative affect and increased acceptance and openness during painful

sensory stimuli (Lutz et al., 2013). Meditation hours were significantly negatively associated with

baseline left anterior insula activation, even after controlling for age. This unique neural signature

was also associated with reduced activation over time in the amygdala and pain-processing areas

before pain-onset, demonstrating a decreased temporal slope of brain activation in the posterior

and mid insula, S2, and mid-cingulate cortex (MCC) in LTMs compared to novices. A similar

pattern was observed during pain episodes in the posterior and mid insula and S2 (Lutz et al.,

2013). Amygdala hyperactivation is evidenced to play a major role in pain processing,

20
particularly regarding the emotional-affective component of pain (Veinante et al., 2013). This key

region receives bottom-up sensory pain information from the brainstem as well as polymodal

information from the thalamus (Veinante et al., 2013). Its activation is additionally inhibited by

mPFC (Neugebauer, 2015). Pathological overactivation of the amygdala and decreased afferent

inhibition from mPFC are linked to pain-related disorders, notably physiological pain in chronic

pain and emotional pain in MDD patients (Thompson et al., 2017). Results are in line with an

advanced implicit reappraisal of painful stimuli in the context of long-term meditation, suggesting

overall improved self-regulation of pain by attenuating its emotional impact rather than its

sensory intensity. This indicates that meditators may have learned to decouple the monitoring of

aversive stimulation from the processes that lead to it being labeled or experienced as pain.

In summary, preliminary evidence from several studies suggests that LTMs engaging in

observing and releasing techniques consistently demonstrate altered pain perception, achieved by

primarily diminishing the unpleasantness of the stimulus and, in some instances, the intensity as

well (Grant & Rainville, 2009; Grant et al., 2010; Grant et al., 2011; Lutz et al., 2013; Perlman et

al., 2010; Zorn et al., 2020). These changes are linked to cognitive decentering from the painful

stimulus, resulting In a separation between the sensory and emotional components of the pain

experience. Conversely to popular models of cognitive control, LTMs’ improvements in pain

relief are not based on increased coupling between executive and pain-processing areas (Grant et

al., 2011) but instead rely on enhanced bottom-up functional integration in key nodes shared

between the SN and pain areas, such as the dACC, anterior insula, and amygdala, regions

involved in attentional control and monitoring of salient stimuli (Bush, 2011; Grant, 2014; Lutz et

al., 2013; Seeley et al., 2007; Sezer et al., 2022). This suggests that LTMs have increased

integration of sensory and cognitive aspects of pain, enabling more accurate and efficient

identification of painful stimuli, while their emotional response to pain, such as labeling it as

‘bad’ and creating an emotional reaction, is reduced. In the following section, we will explicate

eight studies on emotional processing to further extend the gestalt changes introduced in this

section.

21
2.2. Affective Processing and Long-term Meditators

In mindfulness meditation research, attention-based practices are predominant; however, there

is growing interest in other practices, including loving-kindness meditation, due to their potential

to profoundly influence emotional subsystems. Indeed, the regulation of emotion, both through

reappraisal and extinction processes, has been proposed as a central mechanism in meditation

(Hölzel et al., 2011; Wheeler et al., 2017). Emotion regulation encompasses the conscious and

unconscious alterations of emotions by processes before and during which the emotion occurs

(Gross, 1998). For example, effective attentional deployment on a given salience landscape may

influence the onset of the emotional response (Gross & Thompson, 2007). Neuroanatomically,

meditations’ effects on emotion regulation have been associated with activation in mPFC, limbic

regions, and striatum (Hölzel et al., 2011; Tang et al., 2015), increased coupling between the

dorsomedial prefrontal cortex (dmPFC) and rostral anterior cingulate cortex (rACC), and

decreased connectivity between the amygdala and rACC (Sezer et al., 2022). Research on the

neural correlates of trait mindfulness demonstrated a potential association with reduced amygdala

activation and ambiguous findings regarding the impact of trait mindfulness on bottom-up and

top-down emotion regulation mechanisms (Treves et al., 2024).

Utilizing a pre- to post-meditation quasi-experimental design, 12 long-term Canadian Zen

meditators and 10 age- and sex-matched beginner meditators responses to emotional pictures were

compared (Taylor et al., 2011). The meditators averaged 1,709 hours of practice experience,

spanning between 1,000 to 3,000 hours. Analyses revealed reduced emotional intensity ratings in

the meditation condition for negative, neutral, and positive pictures. No differences were found

between the meditators and the controls, suggesting a state-dependent effect.

In a similar research design, emotional word evaluation and implicit affective processing were

investigated using the lexical decision task and valence rating task (Lusnig et al., 2020). The study

included 20 skilled German Zen meditators averaging 1,900 practice hours and a matched control

group. Corroborating the previous results, this study found reduced valence ratings to low arousal

positive and low- and high-arousal negative words in the meditators. However, no differences

were found in the time-constrained lexical decision task, suggesting no influence of meditation on

22
automatic emotional word associations. Importantly, while the meditators engaged in a meditation

session before the tasks, the control group watched an engaging movie, making it challenging to

distinguish between short-term and long-term effects.

The processing of self-related emotional stimuli was assessed in 22 long-term vipassanā

meditators and 22 matched naïve controls using a self-referential word presentation task. During

this task, blocks of adjectives—self-praising, self-critical, negative but not self-critical, and

neutral—were presented, and affect was assessed after each block (Lutz et al., 2016). The

meditators had an average of 5,971 total meditation hours, with 4,861.5 hours specifically in

vipassanā practice, including at least one retreat. Questionnaire results revealed a trend for

enhanced self-compassion and reduced alexithymia (or ‘emotional blindness’) scores, while

demonstrating significantly increased non-activation scores among the meditators. Behaviorally,

although overall affective ratings were not significantly predicted by group, the interaction

between group and condition was significant. Specifically, the difference in affective ratings

between self-related positive and self-related negative adjectives was smaller in long-term

mindfulness practitioners compared to mindfulness-naive participants. This suggests that LTMs

may have a more balanced affective response to self-related stimuli, potentially reflecting greater

emotional regulation or equanimity.

To examine emotional sensitivity, a group of long-term Sant Mat meditators was assessed in

their performance in a color perception task when presented with emotional pictures (Chen et al.,

2018). Sant mat meditation is centered on loving-kindness meditation and encompasses practicing

unwavering compassion as a core focus. The experts (n = 21) meditated on average over 4 hours

per day for the last 13 years, whereas the control group (n = 20) had just started their meditation

practice. Results included diminished general trait and state anxiety for the LTMs that correlated

positively with their years of practice. Furthermore, generally longer reaction times and attenuated

reductions in accuracy during the explicit presentation of emotional pictures indicate lower

hypervigilance, and more neutrality towards explicit emotional stimuli, in the LTMs.

The effect of meditations on affect was studied in 15 LTMs from the Tibetan Nyingma

tradition, known for its emphasis on cultivating altruism, loving-kindness, and compassion

23
(Engen et al., 2015). Averaging 40,000 practice hours, each practitioner completed a compassion

meditation condition and a reappraisal control condition. The outcomes revealed significant

modulation of affect in both practices, with the compassion condition increasing positive affect

compared to all other conditions and mitigating negative affect compared to the passive watching

condition. The reappraisal condition was more effective at reducing negative affect than the

compassion condition, although it did not increase positive affect to the same extent.

Nevertheless, the reappraisal condition still showed more favorable affect modulation than the

passive condition, demonstrating benefits in emotional regulation.

Emotion regulation and its associated neural connectivity patterns may be pivotal in

understanding other areas, such as pain perception, decision-making, and their relationship to

foundational cognitive functions. For example, meditators exhibit increased activation within the

insula, particularly within the right posterior region, while the left anterior insula activation

decreases when faced with an unfair decision (Kirk et al., 2011). Such a pattern of neural

activation suggests a leaning toward greater reliance on interoceptive processing rather than

affective processing during the meditators’ decision-making, as right insula activation has been

associated with increased internal attention (Craig, 2003; Hölzel et al., 2008) and momentary self-

reference (Farb et al., 2007) in meditators. Positioned as a pivotal hub within the limbic network,

the insula is central to emotional processing and empathy, as awareness of internal body

sensations is crucial for their regulation and for caring about others (Hölzel et al., 2011).

Specifically, the right insula is associated with perceptual aspects of empathy, while the left insula

is linked to both perceptual and cognitive evaluative facets of empathy (Fan et al., 2011). In

correspondence with these findings, highly experienced Tibetan meditators, ranging between

10,000 to 50,000 hours of meditation experience, observed increased activation in the right insula

when exposed to negative sounds compared to neutral and positive auditory stimuli and while

practicing compassion meditation (Lutz et al., 2008a). Notably, deeper meditative states increased

insula activation in both groups compared to more superficial meditative blocks. Analyzing the

interaction effects between meditation and rest states in different groups revealed increased

activation in the right TPJ, right posterior superior temporal sulcus (pSTS), amygdala, mPFC,

24
PCC/Precuneus (PCC/Prc), and inferior frontal gyrus (IFG) for experts when exposed to all

sounds. Significant expertise-related effects were observed in PCC/Prc and right pSTS/TPJ

regions, particularly concerning the right lateralization of the latter regions. This suggests that

compared to novices, experts may demonstrate increased social cognition, emotion sharing, and

perspective taking (right pSTS/TPJ; Saxe, 2006; Sommerville & Decety, 2006; Tankersley et al.,

2007), as well as heightened recognition of salient emotional stimuli (right IFG/TPJ; Corbetta &

Shulman, 2002) when presented with emotional human vocalizations during compassion

meditation.

Investigating meditations’ effect on amygdala reactivity, LTMs evidenced increased

activation to explicit happiness, while beginners exhibited the opposite pattern with fear (Chen et

al., 2018). LTMs also displayed reduced amygdala activation to implicit emotional states

regardless of valence, and amygdala activation to explicit fear-mediated meditative effects on

anxiety. In contrast, reduced amygdala activation was found in both the MBSR program group

and the long-term mindfulness meditation cohort to positive images (Kral et al., 2018; see Table 1

for details). Furthermore, only extended practice was associated with reduced amygdala activation

to negative stimuli in the latter study, indicating differing patterns of amygdala responses between

the two investigations. Specifically, retreat hours involving observe and release practices were

associated with decreased activation to negative images. The increased connectivity noted

between the ventromedial prefrontal cortex (vmPFC) and the amygdala within the MBSR group,

but not the experts, implies that higher-level control mechanisms guide initial adjustments in

emotional processing. Over time and with sustained practice, these mechanisms gradually recede

as emotional processing becomes more automatic, mimicking experience-dependent mindfulness-

induced changes in pain (Cooper et al., 2022; Grant, 2014; Hölzel et al., 2011; Kral et al., 2018;

Tang et al., 2015; Wipplinger et al., 2023).

Extending these findings, other research is more ambiguous. For instance, Chen et al. (2018)

found increased functional connectivity between amygdala and ventrolateral PFC for happiness,

and reduced connectivity between insula and medial orbitofrontal cortex (mOFC) for fear,

indicating emotion-dependent corticolimbic connectivity changes in meditators. Another study

25
did not identify a link between improved emotion regulation and higher-order cognitive control

mechanisms; instead, improved emotion regulation was attributed to individuals’ ability to

practice acceptance and maintain presence during emotional states (Taylor et al., 2011).

Conversely, increased activation of frontal and limbic regions was observed during emotional

self-appraisal in long-term vipassanā practitioners, most strongly for positive appraisal (Lutz et

al., 2016). Specifically, enhanced dmPFC activation associated with self-reported reductions in

habitual emotional non-activation. All studies thus revealed distinct neural signatures within their

respective groups. Taylor et al.’s Zen meditators exhibited reduced activation in the default-mode

network (DMN), particularly in the right PCC and right mPFC, compared to attenuated activation

in the left amygdala among their novice participants. As the DMN is associated with self-

referential processing, these findings suggest LTMs’ improved emotion regulation may be due to

altered self-awareness instead of enhanced cortico-limbic control. Indeed, reduced within-DMN

activation has been associated with trait-mindfulness-induced changes in self-awareness (Treves

et al., 2024). We also observed reduced functional connectivity between the dmPFC and posterior

midline nodes (precuneus) despite seeing conflicting findings regarding activation in frontal

midline nodes (cf. Cooper et al., 2022; Lutz et al., 2016; Taylor et al., 2011).

To delineate differences between meditation practices and meditative experiences on affective

processing, practitioners with various degrees of expertise in focused attention (FA) and loving-

kindness meditation (LKM) were recruited (Lee et al., 2012). Long-term FA meditators exhibited

heightened brain activation in the left insula when exposed to happy pictures during meditation,

whereas the long-term LKM group showed increased activation in the left ventral ACC, right

IFG, and right Prc. When confronted with sad pictures during meditation, the long-term FA group

displayed significant activation in the left SFG and right IFG, while the long-term LKM group

showed activation in the left middle frontal gyrus (MFG) and left caudate. These areas have been

associated with reorienting attention to endogenous stimuli (MFG; Japee et al., 2015), working

memory (SFG; Boisgueheneuc et al., 2006), attentional control and response inhibition (IFG;

Swick et al., 2008; Hampshire et al., 2010; Liakakis et al., 2011) and several other higher-order

functions, such as speech, language comprehension, reasoning, and empathy (Li et al., 2013;

26
Liakakis et al., 2011). Consequently, these findings suggest enhanced activation in attentional and

emotional processing regions (insula, SFG, IFG) among FA experts, irrespective of picture

valence. In contrast, LKM experts exhibited increased activation in brain regions associated with

emotion identification (left ventral ACC), regulation (right IFG), and self-referential processing

(Prc) when viewing happy pictures, and heightened activation in regions (left MFG/caudate)

linked to emotion activation and voluntary emotion regulation when viewing sad pictures. These

results imply that LKM practice may promote greater emotional sharing and the ability to

cultivate positive emotions in response to affective stimuli. Increases in positive affect may go

beyond the mere response to external stimuli but may instead reflect an internally generated

positive emotional and motivational state. Indeed, Engen et al. demonstrated a heightened

activation of affiliation, positive affect, and reward processing centers, including the bilateral mid

insula, ventral striatum/nucleus accumbens (VS/NACC), and mOFC, prior to stimulus onset,

perhaps suggesting an overall increase in baseline positive affect in LTMs engaging in

compassion practice (Engen et al., 2015). LKM/compassion practice may thus be particularly

helpful in interpersonal situations in which emotional connection predominates instead of the

mere regulation of affective reactions to stressful situations.

In conclusion, neuroimaging of LTMs reveals significant alterations in cortical, limbic and

basal ganglia regions associated with emotional processing, attention and cognitive control, self-

awareness, social cognition and empathy, reward and motivation, interoception, and language and

speech processing. The precise neuroanatomical configuration underlying these changes remains

to be investigated, as heterogeneity in tasks, meditative experience, and meditative activation

prevented homogeneous neural patterns from emerging. Previous research suggested practice- and

experience-dependent differences in emotion regulation, with long-term practitioners potentially

relying on bottom-up and beginners on top-down regulation strategies (Chiesa et al., 2013). Our

findings partially corroborate improved bottom-up emotion regulation in LTMs (Kral et al., 2018;

Taylor et al., 2011) but also suggest a more complex picture, where frontal coupling with limbic

structures may be both increased or decreased (Chen et al., 2018) depending on the task and/or

meditation employed. Increases in corticolimbic coupling in LTMs may demonstrate different

27
signatures, such as connections between more medial (i.e., mOFC) and subcortical structures

(Engen et al., 2015), indicating that emotion regulation in this population may depend on a unique

mix of bottom-up and top-down strategies (Lutz et al., 2008a; Lutz et al., 2016). The noted

behavioral improvements may, similar to reductions in pain, rely on increases in equanimity, as

we observed enhanced emotional neutrality, acceptance, reduced judgment and negative affect,

with compassion meditation potentially elevating positive affect, social cognition, and mentation.

Nonetheless, additional research is needed to pinpoint the precise impacts of diverse meditation

practices on emotional processing and to uncover the developmental changes that transpire in

LTMs over time.

2.3. Decision-Making—Higher-Order Processes in Long-term Meditators

Higher-order cognition refers to advanced processes beyond basic perceptual, sensory, and

memory processing. These involve higher-level mental operations such as abstraction, analysis,

synthesis, evaluation, planning, and cognitive flexibility, enabling individuals to perform complex

problem-solving, reasoning, and decision-making (Luna et al., 2015; Miller & Cohen, 2001;

Miller & Wallis, 2009; Unsworth et al., 2009). Here, we review the two studies on decision-

making in LTMs.

Contemplative practices may fundamentally alter the behavior of individuals. Previous

research has demonstrated improvements in both social and non-social decision-making among

meditators, likely due to enhanced emotion regulation, empathy, and cognitive control (Sun et al.,

2015). Decision-making is a higher-order cognitive process involving the selection of choices

based on evaluating costs and benefits. Leading cognitive models, such as the ‘emotion-imbued

choice model,’ suggest that emotional processes play both direct and indirect roles in decision-

making, combining rational and non-rational elements (Lerner et al., 2015). Other factors like

cognitive biases, heuristics, information constraints, time limitations, and social influences also

shape decisions, with non-rational choices often upholding social conventions (Boyd et al., 2003).

Key prefrontal regions (dlPFC, ACC, mOFC) are crucial for decision-making (Wallis, 2007),

while subcortical areas (insula, amygdala) and other cortical regions (TPJ, vmPFC, dmPFC, IFG,

28
pSTS) are involved in social decision-making (Sun et al., 2015). Cortical areas are associated with

long-term outcomes, and subcortical regions with short-term outcomes, reflecting a recursive top-

down model (Rosenbloom et al., 2012).

To investigate the impact of meditation-induced trait changes on decision-making, a quasi-

experimental neurobehavioral research design was employed among 26 skilled Buddhist

meditators, averaging 9.5 years of meditation experience, and 40 individuals with no meditation

experience as controls (Kirk et al., 2011). Both groups were assessed for their economic decisions

while playing the Ultimatum game. Participants played 45 rounds of the Ultimatum Game as

responders, facing human partners for 30 rounds and computer partners for 15 rounds, with

predetermined offers varying in fairness. The task assessed their willingness to accept ‘unfair’

offers for prosocial reasons versus rejecting them for fairness. The results unveiled more rational

decision-making tendencies in LTMs compared to the control group, with the former accepting

more than half of the most asymmetrical offers while the latter accepted only a quarter. In social

interactions involving rewards, most individuals tend to evaluate their rewards relative to those of

their peers (Boyd et al., 2003). However, these findings suggest a distinct behavior by the

meditators of accepting even the most unfair offers. Corresponding with the behavioral results, the

meditators exhibited increased activation in the posterior insula, an area central to interoceptive

processing (see Section 2.2. Emotional Processing). Anterior insula activation associated with

affective processing was decreased in the meditators, whereas the opposite was observed for the

controls. Furthermore, additional subanalyses showed increased dlPFC activation in particularly

rational control participants, whereas meditators exhibited increases in the postcentral gyrus,

posterior superior temporal cortex, and parahippocampal gyrus. In addition to the generally

enhanced rational decision-making in meditators, rationality may be mediated through different

neural mechanisms in both groups, with meditators relying on altered somatosensory and

perceptual functioning, such as differing body awareness induced feeling states (postcentral

gyrus; Critchley et al., 2004; Lutz et al., 2008a), perspective taking (posterior superior temporal

cortex; Hampton et al., 2008), and altruism (posterior superior temporal cortex; Tankersley et al.,

2007), and controls relying on cognitive self-control (dlPFC; Hare et al., 2009).

29
 To understand decision-making through the intricate effects of meditation’s attentional

manipulation, recent research utilized the drift-diffusion model (DDM) to computationally model

their reciprocal effects (van Vugt & van den Hurk, 2017). The researchers simulated shifts in

accuracy and response times through the lens of the attentional network task across two distinct

datasets involving seasoned meditators (For dataset one, see: van den Hurk et al., 2010; dataset

two: see Table 1). The attentional network task measures the efficiency of alerting, orienting, and

executive control networks by evaluating participants’ reaction times and accuracy in responding

to visual cues and targets that vary in location and congruence. The DDM represents a

computational framework for understanding decision-making, elucidating how individuals

integrate noisy evidence over time to opt between competing alternatives (Fudenberg et al., 2020).

The evidence accumulates over time towards decision thresholds representing the response

options (e.g., “left” and “right”). Once the evidence crosses a threshold, the corresponding

decision is made. With the DDM, the drift rate represents the speed of evidence accumulation, for

example, influenced by the ability to extract relevant information from a stimulus. Through

systematic manipulations of cues and congruency, researchers adeptly fitted these parameters on

the DDM. Group-independent findings demonstrated a heightened decision threshold, reduced

drift rate (information quality), and drift variability (sustained attention) for incongruent trials,

while informative cues decreased the decision threshold and increased drift rate variability. This

aligns with previous research, which indicates that uncertainty is associated with reduced

readiness for choice-making and enhanced diverse information sampling. Despite that, meditators

exhibited higher decision thresholds compared to non-meditators, suggesting a more deliberate

approach before responding to a stimulus, with this effect being most pronounced during

incongruent trials.

In summary, there is initial evidence that LTMs demonstrated enhanced rational decision-

making and increased decision thresholds, particularly when encountering conflicting

information. This may suggest a tendency to accrue more information before deciding and being

more objective throughout the process.

30
2.4. Non-Ordinary States of Consciousness and Long-term Meditators

Non-ordinary states of consciousness, characterized by alterations in time, body, self, and

space perception, are observed in both clinical and non-clinical contexts and can occur

spontaneously or be induced by practices like meditation, psychedelic substance use, fasting, and

hypnosis (Millière et al., 2018; Timmermann et al., 2023; Wright et al., 2024).

Neurophenomenology, conceived as a functional solution to the ‘hard problem of consciousness,’

involves merging objective brain data with subjective accounts and has gained traction in

exploring various facets of non-ordinary states, including meditation-induced changes to self-

boundaries (Berkovich-Ohana et al., 2020), cessations of consciousness (Chowdhury et al., 2023;

van Lutterveld et al., 2024), and deep concentrative absorptive states (Ganesan et al., 2024;

Sparby & Sacchet, 2024; Yang et al., 2023, 2024). In that vein, recently published work from our

group began to translate conceptual elaborations of traditional texts on meditative states and

stages into scientific language to enable a rigorous interdisciplinary study of meditative

development (Galante et al., 2023). Through systematic phenomenological classification, we

characterized the temporal unfolding of advanced concentrative absorptive states, such as jhānas

(Ganesan et al., 2024; Sparby & Sacchet, 2024; Yang et al., 2023, 2024), as well as meditative

endpoints, such as complete cessations of consciousness (nirodha, Agarwal & Laukkonen, 2024;

Chowdhury et al., 2023; Laukkonen et al., 2023; van Lutterveld et al., 2024). Non-ordinary states

of consciousness are surprisingly common yet vastly understudied (Wright et al., 2024) and thus

might particularly benefit from research in advanced meditation and emergent phenomenology to

better identify and support individuals experiencing potentially adverse effects from those

experiences (Sandilands & Ingram, 2024; Wright et al., 2024). Recent multidimensional

frameworks for the study of self-consciousness (Millière et al., 2018) and general

neurophenomenology (Timmermann et al., 2023) could provide structure to comprehensively

investigate subtle differences and similarities between different modalities (i.e., meditation and

psychedelics) and internal and external contextual factors (i.e., personality and culture). Thus far,

meditation-induced changes in self-consciousness have been linked to increased insula activation,

31
as well as reduced DMN activation and within-network rsFC, particularly in the mPFC and

PCC/Prc (Hölzel et al., 2011; Millière et al., 2018; Tang et al., 2015).

Researchers explored temporal cognition changes in four different meditation groups

compared to age-matched controls utilizing a pre-post meditation design (Berkovich-Ohana et al.,

2011). The meditation groups were separated into three mindfulness-based groups, averaging 894

hours (short-term), 2,570 hours (intermediate-term), and 7,556 hours of meditation experience

(long-term). The fourth group practiced transcendental meditation (averaging 16,310 meditation

hours), a meditation technique that uses the silent repetition of a mantra while remaining in a

focused state. Employing a time production task, participants were instructed to press a button

corresponding to target durations signaled by a sound. The intervals were presented in a random

order, and the participants completed the task with their eyes closed. This method measured their

ability to estimate time by analyzing the relationship between the durations they produced and the

target durations. The researchers observed significantly increased time production in the

mindfulness groups but not the other groups. Meditation experience and the meditative state did

not impact time production. This finding is congruent with the cognitive-timer model, which

suggests that lower arousal and higher attention increase time production—both reliable effects of

mindfulness meditation. Mindfulness meditators might have relied on more momentary core

consciousness based on sensorial processing rather than on memory.

Corroborating this finding, a subsequent neurophenomenological study in 12 long-term

Theravada mindfulness meditators (averaging 11,250 hours of meditation experience) suggests

that experiences of ‘timelessness’ and ‘spacelessness’ are linked to neural network changes in

bodily processing, as evidenced by increased theta-band activation in the right TPJ, PCC/Prc, and

cerebellum (Berkovich-Ohana et al., 2013). Theta-band activation is seen in states of deep

relaxation (i.e., meditation and hypnosis) and has been associated with temporospatial processing,

perhaps by allowing for widespread neuropsychological integration across broad areas (Hasselmo

& Stern, 2014; Hunt, 2007; Vaitl et al., 2005). Moreover, PCC and Prc have been linked to

consciousness (Cavanna & Trimble, 2006), TPJ to first-person multisensorial body processing

(Decety & Lamm, 2007), and cerebellum to altered body awareness (Barmack, 2003). These non-

32
ordinary states of consciousness were discrete from memory and imagination processes associated

with the control “then” and “there” conditions, in which meditators were instructed neither to be

outside of time and space, nor in the present, but localized in the near past or in a specific

location. Notably, neurophenomenologically guided analyses demonstrated decreased right TPJ

and insula activation and increased cerebellum activation in a group of participants more capable

of inducing non-ordinary states of ‘timelessness’ and ‘spacelessness.’

A recent study recruited 22 LTMs to assess their first-person account of time perception and

self-boundaries before and after 20 minutes of meditation compared to a reading control condition

(Gutiérrez et al., 2022). The meditators accumulated 2,994 hours of lifetime practice using

present-moment awareness techniques, as common in Zen, vipassanā, Tibetan Buddhism, and

secularized mindfulness. In the within-subjects design, the meditation condition elicited a

diminishment of bodily boundaries, an elongated sense of time passing, and reduced attention

focused on time when contrasted with the reading condition. To quantify the impact of meditation

on present-moment awareness, a metronome task was implemented, gauging the integration of

successive beats as a surrogate for present-moment duration. However, in contrast to prior

findings, no distinctions emerged between the groups in relation to this task, possibly due to a

misunderstanding on behalf of the participants or trait-based ceiling effects equivocating the

results of both conditions.

These and other findings show that changes in perceived time and self often co-occur in non-

ordinary states of consciousness (Droit-Volet & Dambrun, 2019). Consequently, investigating

each of these phenomena can provide mutual insights into the other. By comprehending the

alterations in perceived time, we can better understand the changes in self-experience and vice

versa.

Exploring the different facets of self-dissolution, Nave and colleagues investigated the

phenomenological fingerprints of 46 LTMs from various traditions after participating in a three-

week meditative training in Theravada Buddhism (Nave et al., 2021). The participants had an

average of 3,832 hours of meditative experience, spanning from 115 to 24,837 hours and

including at least one retreat. Within the context of the meditative technique and the extent of

33
dissolution, six experiential dimensions were examined using microphenomenology and

quantitative analysis: perception of location, agency, first-person perspective, attention, body

sensations, and affective valence. The research uncovered a singular dimension associated with

boundary dissolution, with agency and, to a lesser degree, location driving the dissolution process.

Boundary dissolution was also linked to more positive affect than maintaining body boundaries.

Particularly, practices centered on observation and release, inducing relaxation in attention and

agency, were found to be efficacious in promoting dissolution. According to the authors, this

observation is in harmony with the enactive perspective of the minimal self, which posits its

existence due to foundational sensorimotor and attentional processes. Moreover, total meditation

experience correlated weakly to moderately with the degree of dissolution and the

phenomenological clusters of attention, first-person perspective, and agency, such that more

experience was associated with a more dynamic, broad, and formless attention, a more passive

sense of agency, and enhanced non-dual awareness.

Indeed, prolonged meditation has been documented to induce the phenomenological state

known as ‘effortless awareness.’ This state is characterized by a distinct absence of focused

attention and cognitive effort and is often associated with perceptual alterations, including shifts

in the perception of spatial dimensions. As such, Van Lutterveld and colleagues investigated the

effects of targeted EEG neurofeedback during effortless awareness meditation on meditative

depth by enrolling 16 novice and 16 LTMs from multiple traditions with an average of 6,164

meditation hours (van Lutterveld et al., 2017). Specifically, the authors utilized gamma band

activation in the PCC as neurofeedback for the participants’ meditation, as activation in this brain

area has been inversely correlated with the experience of effortless awareness. Furthermore, the

PCC is a central hub of the DMN, a brain network reliably decreased by meditation (Brewer et al.,

2011). Findings showed a strong correspondence between decreased PCC activation and effortless

awareness in both groups, without any significant difference between groups. These results were

replicated moment-to-moment, and participants could successfully modulate PCC activation

while meditating through momentary feedback. However, no significant differences between

groups were observed.

34
 These findings converge with other research indicating that reduced within-DMN activation,

especially between the mPFC and PCC, may be at the core of trait mindfulness-induced changes

in self-awareness (Treves et al., 2024) and foster a transition from a temporally extended narrative

sense of self to an experiential, dynamic, and de-reified sense of self (Farb et al., 2007; Gallagher

et al., 2023; Tang et al., 2015; Timmermann et al., 2023; Vago & Silbersweig, 2012). This is

further reflected in a study investigating the synergistic effects of psychedelics on meditation

during a Zen retreat, wherein the mPFC and PCC decoupling predicted ego-dissolution during the

retreat (Smigielski et al., 2019) and in a recent longitudinal transcranial-focused ultrasound study,

in which targeting the PCC led to increased mindfulness, alterations in sense of self, time, and

memory recollections, and reductions in functional connectivity within the DMN and between

PCC and dlPFC (Lord et al., 2024). To summarize, LTMs can reliably and voluntarily modulate

dimensions of time and space within their consciousness, with various accompanying degrees of

loss of self-boundaries. These shifts into non-ordinary states of consciousness are based on altered

bodily and self-referential processing with a concomitant deemphasis on memory processes.

Difficulty exists in separating the individual phenomenological patterns, as a large overlap in their

respective dimensions is common. In this context, future research should use precise

phenomenological classification systems that encompass meditative activities and associated

experiential patterns (i.e., the jhānas; see Sparby & Sacchet, 2022, 2024; Yang et al., 2024) while

also employing frameworks, such as the Thin Model (cf. Wright et al., 2023), to generate testable

neuroscientific hypotheses for delineating the similarities and differences between non-ordinary

states of consciousness in LTMs.

3. Discussion, Limitations, and Future Directions

3.1. Methodological Limitations in the Study of Long-term Meditators

To properly discuss the results and patterns observed throughout this review, it is essential to

review the research’s methodological frame and quality. Numerous researchers have raised

concerns about the current state of contemplative neuroscience, noting challenges such as

35
inconsistent research findings and false positives (Kral et al., 2022). These issues may stem from

the broad use of ‘mindfulness’ as an umbrella term, heterogeneity in reporting and study design,

and varying levels of methodological rigor (Sezer et al., 2022; Vago et al., 2019; van Dam et al.,

2018). This situation highlights areas for development, such as improving construct validity in

mindfulness measures, gathering more evidence for ecological validity, and increasing awareness

of potential adverse effects (Van Dam et al., 2018). Consequently, the resulting interpretations are

often highly context-dependent and make generalizability difficult. Moreover, future research may

also require new measurement tools to tease phenomenology from appraisal, better time

conceptualization to understand the meditator’s development, and more extensive samples as well

as rigorous research methodologies through preregistering their methods (Galante et al., 2023).

Due to the complex interplay of these elements, we, in Galante et al. (2023), proposed that

studying them within unified models of mindfulness meditative development will provide a more

comprehensive understanding of the subject. In the following, the methodological limitations of

the reviewed studies of LTMs will be discussed. This will hopefully highlight the future steps

necessary for creating a unified framework for meditation research (Galante et al., 2023).

One research aspect of the unified framework concerns the predictors of meditation research,

such as dose, technique, individual characteristics, phenomenological patterns, and sociocultural

context. The first limitation pertains to meditation dose and technique predictors. First, a lack of

precision in reporting habitual meditation techniques has limited interpretations pertaining to the

trait effects of the LTMs. For example, the classification of ‘Buddhist meditation’ (Kirk et al.,

2011) is too broad to allow for conclusions regarding the impact of specific meditation types on

cognitive functions. Furthermore, whereas some studies defined mindfulness meditation as an

equal balance of samatha (focused observing) and vipassanā (observing and releasing; Kral et al.,

2018; van Vugt & van den Hurk, 2017), and vipassanā itself as mostly encompassing observing

and release practices (Kral et al., 2018), others did not define the meditative activation at all (i.e.,

vipassanā in Khalsa et al., 2008). Other studies ambiguously referred to Zen practice as based on

observing and releasing activities (Grant et al., 2011) or mindfulness meditation (Grant et al.,

2009; Grant et al., 2010; Taylor et al., 2011) or both, equating mindfulness meditation with

36
observing and release activities (Lusnig et al., 2020). These examples illustrate the significant

heterogeneity in defining meditation activities. To investigate potential trait or state-trait

interaction effects, it is crucial to clearly report and distinguish specific habitual meditation

experiences from the practices used in the experimental design. Additionally, total meditation

experience requires systematic quantification, as some researchers only report the total years of

meditation (van den Hurk et al., 2010), or total meditation hours (Lee et al., 2012), or both (Grant

et al., 2010). Other studies failed to report mean scores (Lutz et al., 2008a; Perlman et al., 2010;

van Vugt & van den Hurk, 2017), and most did not describe average daily practice amounts

before engaging in the experiment (Sparby & Sacchet, 2022).

The second observed methodological limitation pertains to the social, cultural, and historical

context of the practice. First, the studies lacked comprehensive reporting of demographic

information, specifically on the race, ethnicity, and culture of the participants, often only

mentioning the country of recruitment. This is concerning for multiple reasons. First, cultural

background significantly influences cognitive processes by shaping the external representations

and tools people use, the content of their thoughts, and their habitual practices, thereby potentially

affecting how information is processed, what cognitive strategies may be employed, and even the

neurological structures of the brain involved (Bender & Beller, 2013; Kitayama & Murata, 2013;

Lao et al., 2013; Nisbett & Miyamoto, 2005). Second, meditative practices are deeply rooted in

centuries-old cultural traditions of Buddhist origin (Bodhi, 2011; Nilsson & Kazemi, 2016; Sharf,

2014; Sugunasiri, 2008). Existing research has revealed concerning findings on the cross-cultural

validity of psychometric measures, exemplified by the observation that American college students

exhibited equivalent mindfulness scores compared to Theravada Buddhist monks (Christopher et

al., 2009). Second, little to no data were collected on advanced phenomenological states, leaving

no information on whether meditators’ experience had reached significant stages of meditative

development and advanced meditation more broadly. Although several studies reported over

10,000 hours of lifetime meditation experience, the absence of first-person data-limited

assessments of meditative mastery, making it difficult to distinguish between LTMs and those

who may also be advanced meditators (Galante et al., 2023; Sacchet et al., 2024; Sparby &

37
Sacchet, 2022; Yang et al., 2024). A comprehensive retrospective phenomenological assessment

may still be beyond the reach of current contemplative science, as it requires systematic efforts to

scientifically evaluate, translate, and map traditional stages of development to contemporary

empirically informed models (Grabovac et al., 2015; Sparby & Sacchet, 2022; Wright et al.,

2023). However, this may be necessary to bridge the gap between behavioral, neural, and

phenomenological observations of different meditative populations, enabling individualized

multivariate statistical approaches that provide a more unified understanding of the relationship

between first-person accounts of well-being or psychological distress and cognitive processes.

Taken together, these methodological limitations extend previous critiques of the literature

(Sezer et al., 2022; Galante et al., 2023; Vago et al., 2018; van Dam et al., 2018) and emphasize

the need for a unified framework in meditation research, particularly for advanced meditation

(Sacchet et al., 2024). In the following section, we will summarize the behavioral trait effects

identified in our review and specifically delineate the differences between meditative activities.

3.2. Behavioral Trait Effects

Although most studies in this review exhibited substantial changes in the cognitive functions

of LTMs, the variable methodological design challenged evaluating true trait effects observed in

this population (refer to Table 1 for further information). Considering this, we will first

schematize and summarize the previous findings and then explicate specific methodological

nuances.

Collectively, LTMs exhibited between-group improvements in interoceptive accuracy (Fox et

al., 2012), albeit not extending to cardiac interoceptive awareness (Khalsa et al., 2020); reductions

in pain sensitivity (Grant & Rainville, 2009; Grant et al., 2010; Grant et al., 2011; Perlman et al.,

2010) and pain unpleasantness (Lutz et al., 2013; Perlman et al., 2010; Zorn et al., 2021);

increased emotional neutrality (Chen et al., 2018; Engen et al., 2015; Lusnig et al., 2020), positive

affect (Engen et al., 2015), and mitigated negative affect (Chen et al., 2018). Concomitant

changes in emotional neutrality and affect may be best explained by the study design, as well as

38
the cognitive tasks and measurements employed, instead of differences in meditative activities.

Although meditation reliably induced increased time production (Berkovich-Ohana et al., 2011),

an elongated sense of time-passing (Gutiérrez et al., 2022), and states of effortless awareness (van

Lutterveld et al., 2017), there were no significant differences between LTMs and beginners,

suggesting activation-dependent state effects. The lack of control groups impedes insight into

persistent phenomenological changes within LTMs, but heterogeneous samples point to an

increased capacity for body- and self-boundary dissolution (Nave et al., 2021) and experiences of

‘Timelessness’ and ‘Spacelessness’ under meditation (Berkovich-Ohana et al., 2013). The

experiences of dissolution are linked to five distinct dimensions: passive agency, non-locality,

non-dual perception, formless attention, and body imperceptibility (Nave et al., 2021). However,

further research is imperative to distinctly delineate the phenomenological alterations attributable

to pure trait effects and those arising from trait-state interactions. Lastly, LTMs’ evidence

increased rational decision-making (Kirk et al., 2011), as well as decision thresholds, particularly

for incongruent cues (van Vugt & van den Hurk, 2017), suggesting heightened objectivity and

deliberateness before acting.

To explore if meditation experience yielded incremental cognitive benefits, associations

between meditative expertise and biobehavioral cognitive outcomes were observed in pain

perception (Grant & Rainville, 2009), anxiety (Chen et al., 2018), non-ordinary states of

consciousness, specifically phenomenological self-boundary dissolution, non-dual awareness,

attentional broadening, and volitional agentic control (Nave et al., 2021), and introspective

accuracy (Fox et al., 2012). One study on pain perception investigated the relationship but lacked

significant correlations (Grant et al., 2010), another found no link between meditation experience

and pain catastrophizing (Zorn et al., 2020), and one directly compared different expertise levels

(Berkovich-Ohana et al., 2011) but did not find trait differences in time production. Increased

brain thickness in pain and emotion-associated regions (ACC, S1; Grant et al., 2010), as well as

decreased activation in pain (dACC, thalamus, insula (specifically left anterior insula; Lutz et al.,

2013)) and PFC areas (dlPFC, med-PFC/OFC; Grant et al., 2011) were found when analyzing

correlations between brain changes and meditation experience. Furthermore, meditation retreat

39
hours predicted reduced amygdala activation to negative pictures (Kral et al., 2018). No

significant link with meditation experience was found in a study investigating brain activation to

emotional pictures (Taylor et al., 2011) and in research exploring the brain effects of compassion

meditation (Lutz et al., 2008a). In summary, although dose-response effects within prolonged

meditation practice seem likely, many studies did not directly assess this relationship, preventing

comprehensive insight into the temporal development of cognitive processes. Future work should

comprehensively quantify meditative predictor variables and systematically analyze dose-

response effects, including potential non-linearities between meditative dose and development

(Cearns & Clark, 2023; Cooper et al., 2022; Galante et al., 2023; Lindström et al., 2023).

Additionally, it should consider other moderating factors when encountering diverging findings.

To understand how meditator characteristics influence cognitive processes, we will next explicate

the effects of various meditation activities on the observed results in LTMs.

The most consistent meditation-specific effects were evident in pain perception, favoring

observing and releasing practices. This practice might be particularly effective in promoting

cognitive decentering, a core mechanism associated with meditation-induced pain reduction (Zorn

et al., 2021). Other mechanistic research conducted with chronic pain patients has identified

decentering as a primary marker of psychological flexibility and the quality of functioning within

that population (McCracken et al., 2013). Qualitative research further supports these findings by

revealing an increase in decentering and reappraisal as meditation experience accumulates (Poletti

et al., 2020). For example, novice practitioners engage in increased experiential avoidance and

perceive pain as purely physical. In contrast, long-term practitioners develop patterns of

reappraisal, openness, and curiosity, ultimately viewing pain as a mental construct that can be

harnessed for altruistic purposes. Overall, research into meditation-based pain regulation suggests

that these effects primarily hinge on perceptual modifications, where enhanced cognitive-sensory

integration and decoupling of affective processes may lead to improved sensory clarity and

equanimity. Cognitive decentering, with its potential to uncouple sensory and affective

components, may facilitate emotional and cognitive control, ultimately enabling a cognitive shift

that allows for the recontextualization of noxious stimuli (Garland et al., 2015; Zeidan et al.,

40
2012; Zorn et al., 2021). Similarly, preliminary evidence points to observing and releasing

practices as particularly efficacious for cultivating experiences of self- and boundary-dissolution,

as they cultivate attentional disengagement and agentic passivity (Nave et al., 2021). This aligns

with the predictive processing hypothesis of meditation research, suggesting that attentional

relaxation or “in-action” reduces precision weights and consequently self-evidencing, ultimately

flattening the predictive hierarchy and giving rise to non-dual awareness, characterized by a

unified field of phenomenological experience of “here” and “now,” in which the dichotomy of

perceiver and perceived has collapsed (Laukkonen & Slagter, 2021; Lutz et al., 2019).

In the final section of the discussion, we will integrate our diverse findings with contemporary

neurophenomenological theories. Using a self-processing perspective, we will explore the

dynamical relations between the various cognitive domains and their neural correlates, and

discuss how LTMs may exemplify an embodied dynamical gestalt of mindfulness.

3.3. Self-Processing and the Brain in Long-term Meditators

Extensive research systematically synthesized brain changes associated with mindfulness and

meditation (Cooper et al., 2022; Fox et al., 2016; Hölzel et al., 2011; Rahrig et al., 2022; Sezer et

al., 2022; Tang et al., 2015; Treves et al., 2024; Vago & Silbersweig, 2012), identifying several

unique neural correlates. Despite evidencing converging findings, a gap remains in consolidating

neural, behavioral, and phenomenological results (Hölzel et al., 2011). Even when disregarding

the vast array of methodological limitations in contemplative research, integrating the various

findings into an overall phenotypic model of meditation is difficult, as the various components are

not distinct but rather highly interrelated processes (Hölzel et al., 2011; Vago & Silbersweig,

2012). As such, a specific behavior may be associated with several neural processes, which, in

turn, dynamically change based on population, as well as meditative activation and practice

experience. To transcend this difficulty, some work relied on a self-processing lens, in which a

model of the self is used as a non-reductive interdisciplinary ground for integration (Cooper et al.,

2022; Vago & Silbersweig, 2012). We will follow in these footsteps by integrating our results

41
 within the pattern theory of self (Gallagher, 2013, 2021, 2024; Gallagher & Daly, 2018; Gallagher

et al., 2023) while using previous syntheses as context.

The pattern theory of self is a non-reductionistic, interdisciplinary enactivist account positing

that the self is a dynamic pattern constituted by a set of factors or processes, which do not have

any strictly necessary conditions but together form a sufficient configuration (Gallagher, 2013,

2021). The self-pattern characterizes a dynamical gestalt, such that the weighting of each

constituent depends on and influences the relations among the other constituents. The theory is

compatible with Buddhist accounts of the self (Gallagher & Daly, 2018; Gallagher et al., 2023)

and offers a robust framework for psychiatry, suggesting that psychopathology can be understood

as a self-disorder (Gallagher, 2024), with each disorder representing distinct and identifiable

patterns. Please see Table 2 for a detailed exposition of the dynamical processes underlying the

self-pattern. Figure 2 summarizes the neurobehavioral findings of our review fractionated by the

various elements of the self-pattern. There are two significant overarching results worth

addressing.

Figure 2

The cognitive neurobehavioral results of long-term meditators contextualized in the pattern theory of
self.

42
Note. The ten circles graphically depict the pattern theory of self, where each self-process dynamically
relates to all other processes, creating an overarching dynamical gestalt (Gallagher, 2021). The central part
of the figure provides a non-comprehensive description of the dynamical gestalt of long-term meditators.
We propose a skill-based comprehensive model of mindfulness as a suitable way to describe changes in
the gestalt and, consequently, the self-pattern, with less transparency in the three factors of mindfulness
indicating enhanced robustness of evidence (Young, 2016). The dynamical gestalt does not causally
influence the pattern; rather, the pattern itself embodies the gestalt through its processes and relations (e.g.,
rational decision-making as a behavioral expression of equanimity). The dynamical gestalt and the
evidence for different self-processes are limited by the scope of the review and do not represent an
absolute depiction of long-term meditators. The boxes summarize the behavioral and neural results for
individual constituents in the self-pattern. The arrows indicate the directionality of change, with grey
arrows denoting preliminary support and dark arrows representing stronger evidence. Small arrows show
the direction of connectivity or activation changes for neural findings. aMCC: anterior midcingulate
cortex; dACC: dorsal anterior cingulate cortex; dmPFC: dorsomedial prefrontal cortex; dlPFC: dorsolateral
prefrontal cortex; IFG: inferior frontal gyrus; mPFC: medial prefrontal cortex; mOFC: medial orbitofrontal
cortex; PCC: posterior cingulate cortex; PCG: precentral gyrus; PHG: parahippocampal gyrus; Prc:
precuneus; pSTC: posterior superior temporal cortex; pSTS: posterior superior temporal sulcus; TPJ:
temporoparietal junction.

First, there is no isolated relationship between neural and behavioral findings; rather, similar

brain regions are associated with multiple distinct behavioral and self-related processes,

cautioning against reverse inference, especially in significantly different populations like this one

43
(Jack et al., 2019). Second, despite no one-to-one coherence between neural and behavioral

activations, results arguably converged toward an embodied dynamical configuration of

mindfulness. From this vantage point, different investigative and explanatory levels enable a

comprehensive and non-reductionistic viewpoint of long-term meditation, as the evidence is still

incomplete and convergence must be sought through triangulation. Accordingly, when practicing

meditation, initiating changes to the self as a cohesive entity may influence the individual

elements that constitute the self. Similarly, modifying these individual elements can adjust the

overall self-pattern by reassigning weights and values to the components of this dynamic process

(Gallagher, 2021). To understand the dynamical gestalt of an LTM, it may be helpful to look at

recent systematic classifications of mindfulness, as extended practice may shape the whole pattern

based on its core tenets. The model depicted in Figure 2 provides a non-comprehensive

characterization of the dynamical configuration of LTMs based on our reviewed evidence and

Young’s mindfulness model (Young, 2016). Crucially, the different elements of Young’s

mindfulness model are embedded and embodied within the self-pattern, rather than emerging

from it as the figure might suggest. For example, rational decision-making is one expression of

equanimity, rather than separate from it. Consequently, the dynamical gestalt is the pattern and its

relations.

Supporting previous research that found a link between body awareness and mindfulness

(Treves et al., 2019), our study found evidence of increased sensory clarity. This clarity influenced

bodily, pre-reflective experiential, affective, behavioral/action-related, and social intersubjective

processes, and it may be particularly linked to increased activation in brain regions within the SN,

such as the insula and dACC, as well as the right TPJ, thalamus, and aMCC (see Figure 2; Grant

et al., 2011; Kirk et al., 2011; Lutz et al., 2013). Increased insula activation is a common finding

in meditation research (Cooper et al., 2022; Fox et al., 2016; Hölzel et al., 2011; Sezer et al.,

2022; Tang et al., 2015; Vago & Silbersweig, 2012; Young et al. 2018) and associated with

increased body awareness (Treves et al., 2024), although more heterogeneity is evident when

delineating nuances. For example, decreased activation within the anterior insula but increased

activation within the posterior insula suggests heightened interoceptive but decreased affective

44
processing (Kirk et al., 2011), and enhanced lateralization to the right indicates stronger

perceptual than cognitive empathy (Fan et al., 2011; Lutz et al., 2008a). As such, the insula may

be posited as an integral hub for the clear comprehension of sensations and gaining differential

insight into different layers of self-processing (Bodhi, 2011; Sharf, 2014). However, more

research is needed to clearly differentiate meditation effects on interoceptive accuracy compared

to perceptual acuity of sensory information and their relationship to insula activation. Studies on

perceptual processing in LTMs evidenced reduced perceptual habituation and increased

perceptual sensitivity to sensorial information, emphasizing mitigated top-down and increased

bottom-up processing (Antonova et al., 2015; Berkovich-Ohana et al., 2013; Fucci et al., 2018;

Kasamatsu & Hirai, 1966). Generally, sensory clarity may directly enhance interoceptive

awareness by improving sensory detection, discrimination, and penetration (Young, 2016). This

enhancement allows for a sequential effect in which lower-level sensory information is better

understood and integrated within higher levels of self-processing, paralleling the previously

proposed frontal-parietal control network, which shows strong overlap in key brain regions

(dACC, aMCC, TPJ; Vago & Silbersweig, 2012). For example, enhanced sensory clarity can help

separate cognitive-evaluative and sensory-discriminant experiences, allowing for the

decomposition of congealed sensory-affective phenomena into their components. This is further

demonstrated by LTMs’ increased activation within the ACC, thalamus, insula, and

somatosensory cortices and decreased activation in the PFC, hypothalamus, and amygdala (Grant

et al., 2011; Lutz et al., 2013), in which aforementioned brain areas (e.g., ACC, insula, thalamus)

are necessary but not sufficient for this deconstruction. Consequently, rational decision-making,

emotion recognition, pain unpleasantness, and emotional neutrality may significantly depend on

clearly delineating sensory information but are also modulated by other relevant self-processes.

Stronger evidence was found for increases in equanimity, as demonstrated by changes in

bodily, pre-reflective experiential, affective, behavioral/action-related, and social intersubjective

processes. Specifically, LTMs displayed reduced pain sensitivity and unpleasantness, enhanced

emotional neutrality, more rational decision-making, and improved empathy. Equanimity may be

defined as a state of internal balance, in which there is no suppression, nor grasping toward

45
internal or external sensory experiences (Young, 2016). Despite disagreement on whether

equanimity is enclosed within or separate from mindfulness, previous research proposed

equanimity as one of the key effects of mindfulness meditation (Eberth et al., 2019) as it is

critically related to emotion regulation and thus well-being (Desbordes et al., 2014). Indeed,

besides changes in sensory processing, improved emotion regulation emerged as the most

dominant result in our review and similarly extended to the other earlier noted self-processes.

Previous research proposed corticolimbic decoupling at the root of emotion regulation in

meditation, consequently extending to favorable changes in other areas, such as pain perception

(Sezer et al., 2022). Besides changes in insula activation, we observed intrinsic activation changes

in affective and pain-related areas, such as the mOFC, aMCC, and limbic regions (amygdala and

VS). Critically, PFC activation was ambiguous, with some research demonstrating increased

activation primarily in the initial stages of emotion regulation (Kral et al., 2018) without showing

any long-term increases (Young et al., 2018). These results, however, may be too simplistic, as

reduced within-DMN activation (Taylor et al., 2011), as well as increased dmPFC and decreased

within-DMN connectivity (Lutz et al., 2016) suggest alternative mechanisms driving

improvements in emotion regulation. The latter finding points to the essentiality of brain regions

involved in attentional control and higher-order executive functions in emotion regulation.

Attention has been discussed as the core cognitive mechanism underlying the benefits of

mindfulness meditation (Malinowski, 2013), with attentional control from continuous practice

being associated with increased rsFC between the dlPFC and PCC when not distinguishing for

experience levels (Sezer et al., 2022) and enhanced surface area in prefrontal and parietal regions

(Treves et al., 2024). However, attention’s role in emotion regulation is more complex and

depends on the meditative practice (Lee et al., 2012). In fact, different meditative activities

facilitate emotion regulation through different neural substrates, which converge in overlapping

brain regions (Fox et al., 2016; Vago & Silbersweig, 2012). For instance, experts in attention-

based practices engage attentional- and emotion-processing areas, whereas loving-kindness

practitioners evidence heightened activation of emotion, affiliation, reward, and self-referential

processing areas (Engen et al., 2015; Lee et al., 2012). Similarly, results from Zen experts

46
uncovered altered mental self-processing as a core component of their emotion regulation (Taylor

et al., 2011). Altered mental self-processing is most closely associated with acute changes in self-

awareness, specifically the experience of the narrative self. Advanced meditators heightened

aptitude to dissolve the narrative self and experience states of “timelessness” and “spacelessness”

may reflect improved voluntary control over crucial DMN regions, such as the PCC (Berkovich-

Ohana et al., 2013). Indeed, reduced PCC activation is one of the most consistent neural findings

in meditators (Cooper et al., 2022), whereas mPFC changes are more ambiguous and may depend

on meditative expertise and/or location of frontal midline activation (ventral/dorsal; Lutz et al.,

2016). Collectively, these results suggest that cognitive functions like emotion regulation

dynamically depend on and influence other self-processes, where altering any constituent could

modulate the entire pattern. For instance, midline DMN activation may be most closely related to

narrative self-referential processes (Northoff et al., 2006), but changing these processes can also

improve emotion regulation (Gallagher, 2024). Consequently, one may detect neural changes

typically associated with narrative self-awareness alongside enhanced behavioral emotional

neutrality during an emotional processing task (Taylor et al., 2011), or observe variable activation

of attentional and self-processing areas depending on the meditative practice employed when

exposed to emotional pictures (Engen et al., 2015; Lee et al., 2012). This insight might be

particularly beneficial if harnessed within precision medicine, as targeting one specific process or

neural correlate associated with the process may provide the necessary control to explore what

else “wiggles” in the dynamic configuration of the pattern (Gallagher, 2024). For example,

previous research using fMRI neurofeedback to target and reduce self-referential DMN processes

observed significant reductions in symptoms associated with other parts of the self-pattern (Bauer

et al., 2020; Zhang et al., 2023). Neuromodulation methods may also extend to non-clinical

applications, such that targeting regions associated with mindful gestalt changes either through

direct modulation (Abellaneda-Pérez, et al., 2024; Lord et al., 2024) or neurofeedback (van

Lutterveld et al., 2017) could be leveraged to deepen meditative practice, perhaps accelerating the

speed of meditative development.

47
 Our results converge with Cooper et al.’s (2022) proposed ‘Topographical Reorganization of

Meditation,’ in which they map the transformation of three spatially nested self-processing layers

onto different stages of meditation experience. Self-processing can be conceptually and

topographically divided into interoceptive, exteroceptive, and mental constituents (Qin et al.,

2020). Interoceptive processing involves physiological interaction with the environment,

exteroceptive processing includes proprioceptive and affective perception, and mental self-

processing encompasses cognitive aspects of self-representation. Lower-level intero-exteroceptive

processing is naturally non-dual, while higher-level mental processing introduces dichotomous

self-object perception. Increased meditation shifts non-dual intero-exteroceptive processing to the

foreground, revealing brain-body-environment interconnectedness, and relegates self-referential

mental processing, making non-dual perception explicit (Thompson & Varela, 2001; Northoff &

Zilio, 2022). Our results similarly suggest improved lower-level intero-exteroceptive self-

processing, subsequently impacting higher-order processes. This also aligns with a review by

Vago and Silbersweig (2012), as our findings observed increased activation in networks

associated with the non-conscious experiential enactive self (limbic regions, thalamus, and

posterior insula) and the conscious experiential phenomenological self (anterior insula, S1, pSTS,

and Prc). Additionally, reduced activation was noted in areas related to the narrative self-

reflective self (mPFC and PCC). Crucially, in Cooper’s model, low meditation proficiency

associates with high DMN and low FPN activation. With ongoing practice, this relationship

reverses and ultimately equalizes to co-active FPN and DMN in long-term meditators. The

authors hypothesize a mediating function of the SN between DMN and FPN, with other research

showing an inverted ‘U’ relationship between SN activation and meditation proficiency

(Brefczynski-Lewis et al., 2007; Cooper et al., 2022; Rahrig et al., 2022). Despite finding some

evidence for increased SN activation (see also fronto-parietal control network; Vago &

Silbersweig, 2012), we could not confirm or deny this temporally sensitive reorganization of the

brain. Due to space constraints, we decided against reviewing the broad attentional literature on

LTMs. Consequently, confirming these topographical changes and evaluating dispositional

increases in attentional concentration (see Figure 2) are beyond the scope of this review.

48
 To summarize, meditations’ cognitive changes may reflect a new neurophenomenological

configuration of self-processing in which a specific cognitive-behavioral outcome may not only

represent changes within a particular brain network but also extend to other networks and regions.

Different mindfulness practices then afford the ability to primarily modulate one or several of

these constituents, such as attention, emotions, or self-consciousness, and provide general

flexibility to transition between different self-networks (Gallagher et al., 2023; Vago &

Silbersweig, 2012). Preliminary evidence suggests a dispositional gestalt of mindfulness

embodied in LTMs, with our review supporting neurobehavioral equanimity and possibly sensory

clarity embedded in the self-pattern. This parallels predictive processing accounts of meditation,

in which the meditative activation offers the learning opportunity to dynamically disengage from

top-down habitual sensory predictions and re-orient to bottom-up sensory information (Lutz et al.,

2019), improving access to the more embodied cognitive-affective aspects of the self (Christoff et

al., 2011; Laukkonen & Slagter, 2021). Increases in subjective well-being may then emerge from

the suspension of habitual avoidant mental actions, allowing previously repressed content to

surface and be acknowledged, thereby enhancing self-understanding and disrupting habitual

processes that maintain negative affect (Deane et al., 2024). Consequently, the meditators

pragmatically acquire increased cognitive flexibility and epistemic access to the cause and effects

of their mental processes, which is purported in Buddhist psychology as essential for gaining

insight into “No-Self,” meaning seeing the self as a dynamic process lacking a substantive

existence and not wrongly identifying with individual components (i.e., the five aggregates; Lutz

et al., 2019; Vago & Silbersweig, 2012). Furthermore, inflexibility within self-processing,

especially when cemented through rigid top-down patterns, might represent a fundamental

transdiagnostic element in psychopathology that meditative practices can alleviate (Gallagher et

al., 2023; Gallagher, 2024; Giommi et al., 2023).

Corroborating the idea of the meditative neurophenomenological gestalt, preliminary research

investigating the functional connectome signature of meditative expertise suggests a relaxation of

the functional gradient hierarchy in LTMs, as well as an integration of major brain

networks; associated with cognitive-affective decentring (Czajko et al., 2023). Gradients are

49
characteristic spatial topographies of brain organization, indexing structural-functional-cognitive

relationships, and linked with cognitive-behavioral developmental changes (Bernhardt et al.,

2022; Dong et al., 2021; Huntenburg et al., 2018). Decentring has been proposed as critical for

non-conceptual cognition (i.e., non-dual awareness) and cognitive-affective reappraisal (Garland

& Fredrickson, 2019; Thompson, 2021). Czajko et al. hypothesized that meditative expertise may

reflect a fundamental shift in embodied cognition, which is described by the degree of enhanced

integration of sensorimotor, attentional, affective networks. Diminished functional segregation

through the compression of the functional gradient between sensory and transmodal brain regions,

thereby increasing structured communication between these areas, is also observed in acute

psychedelic-induced states (Erritzoe et al., 2024). These states exhibit significant overlap with the

neurophenomenology of meditative states, particularly regarding changes in various degrees and

dimensions of self-consciousness (Millière et al., 2018).

In line with prior models, the cognitive improvements from meditative expertise may arise not

from enhanced cognitive control but rather from alterations in the brain’s default activation

patterns related to cognitive processes (Cooper et al., 2022; Tang et al., 2015). Persistent

perceptual shifts toward non-dual awareness and concomitant changes in default cognitive

processing, such as less repression and enhanced attentional engagement with equanimity and

sensory clarity, might be the most distinguishing factor between advanced and beginner

meditators. Short-term practice is sufficient to reliably induce pragmatically and clinically

relevant changes in cognitive functions (Ainsworth et al., 2013; Tang et al., 2007; Zeidan et al.,

2010a; Zeidan et al., 2010b); however, more practice may be necessary to gain access to

fundamental changes in perceptual (Cooper et al., 2022; Josipovic, 2010; Josipovic, 2021), social

(Singleton et al., 2021; Trautwein et al., 2016; Williams et al., 2016), and emotional processing

(Sahdra et al., 2011) as described by ancient soteriological traditions.

To fully understand the long-term effects of meditation on biobehavioral cognitive changes,

future research should prioritize: 1) employing a unified framework to assess predictors of

meditative development and mastery (Galante et al., 2013), 2) systematically gathering

phenomenological data and its associated practices (Sparby & Sacchet, 2022), 3) translating and

50
 testing experience patterns from different stages of meditative development into neuroscientific

hypotheses (Wright et al., 2023); and 4) investigating topological and topographical changes

associated in cognitive functions across timescales to illuminate the interrelated, dynamic, and

nested nature of cognitive and meditative development.

4. Summary and Conclusion

Here, we aggregated and synthesized the biobehavioral cognitive outcomes of LTMs and

some of their neural correlates. Findings indicate preliminary evidence for cognitive alterations in

perception, emotional processing, normative self-awareness, and higher-order functions.

Specifically, long-term practitioners show increased interoceptive awareness, rational decision-

making, emotional neutrality, and self-body-boundary dissolution, as well as reduced pain

perception and potentially normative self-awareness. Notable correlations between meditative

practice experience and cognitive enhancements suggest a potential biobehavioral dose-response

relationship, however, methodological limitations prevent a more definite conclusion.

To address these issues, unitive meditative frameworks, including the systematic assessment

of predictor variables and the sociocultural context, as well as a clear delineation of acute (state)

and habitual (trait) variables in the research design, are necessary to reduce the inconsistencies in

meditation research findings and improve interpretability (Galante et al., 2023). Furthermore,

research should attempt to close the gap between phenomenological reports and brain correlates

by utilizing interdisciplinary communication and models, such as the Thin Model (Wright et al.,

2023). This collaboration will aid the understanding of adverse events associated with meditation

training (cf. Lindahl et al., 2017; Lindahl et al., 2020) and improve evidence-based approaches to

meditation for overall well-being or psychopathologies. Lastly, more LTMs evidence a unique

neurophenomenological configuration of mindfulness, characterized by altered default processing

of affective, attentional, sensorimotor, and self-referential brain areas, as well as potentially

increased cognitive integration. Progressive phenomenological shifts from implicit to explicit

non-dual awareness may represent a new fingerprint of embodied cognition in which top-down

51
mental self-processing recedes and more bottom-up experiential self-processing moves to the

foreground. More specifically, meditative expertise in self-regulation was encoded within the

changes of bodily, pre-reflective experiential, affective, behavioral/action-related, and

social/intersubjective self-processes and their dynamical relations, perhaps reflecting increased

embodied equanimity and sensory clarity. Future research in contemplative neuroscience should

test and expand comprehensive neurophenomenological models, such as the ‘Topographical

Reorganization of Meditation,’ by investigating the topographical and temporal changes in the

context of different cognitive functions and in various meditative populations. Ultimately, the

field must adopt a more comprehensive framework of advanced meditation that meaningfully

integrates both development and phenomenology, moving toward a science of embodied mastery

rather than relying solely on 'long-term practice' as measured by hours completed (Sacchet et al.,

2024). Despite these concerns, preliminary results provide consistent functional,

phenomenological, and neural associations, suggesting promise in closing the methodological gap

within meditation research.

52
Acknowledgments
Declaration of competing interests

Dr. Sacchet and the Meditation Research Program are supported by the National Institute of Mental Health
(Project Number R01MH125850), Dimension Giving Fund, and Tan Teo Charitable Foundation.

Glossary

ACC anterior cingulate cortex

aMCC anterior mid-cingulate

ACC anterior cingulate cortex

dACC dorsal anterior cingulate cortex

DDM drift diffusion model

DMN default mode network; associated with self-referential thought,

autobiographical memory, and mental time travel

dmPFC dorsomedial prefrontal cortex

dlPFC dorsolateral prefrontal cortex

FPN frontoparietal network

IFG inferior frontal gyrus

IPS intraparietal sulcus

LKM loving-kindness meditation

OFC orbitofrontal cortex

PCC posterior cingulate cortex

PFC prefrontal cortex

Prc precuneus

pSTS posterior superior temporal sulcus

MCC mid-cingulate cortex

MFG middle frontal gyrus

mOFC medial orbitofrontal cortex

mPFC medial prefrontal cortex

53
Mindfulness meditation mindfulness meditation comprises samatha and vipassanā, both

observing type practices, with the former based on focusing attention

and the latter a combination of focusing and releasing activities

Nyingma Tradition a specific tradition rooted in Tibetan Buddhism that is known for its

cultivation of compassion, loving-kindness, and altruism.

rACC rostral anterior cingulate cortex

rsFC resting state functional connectivity

S1 primary somatosensory cortex

S2 Secondary somatosensory cortex

Sant Mat mediation Sant mat meditation is centered on loving-kindness meditation and

encompasses practicing unwavering compassion as a core focus.

SFG superior frontal gyrus

SN salience network

Tai Chi Chuan Tai chi chuan is a form of self-defense training that emphasizes

purposeful movement, making it consistent with the concept of an

active form of meditative practice

Theravada Buddhism Theravada or Tibetan Buddhism practices include observing and

releasing and focused observing activities as well as compassion/LKM

(metta) practices

TPJ temporoparietal junction

Transcendental Transcendental meditation is an active meditation technique using the

meditation silent repetition of a mantra while remaining in a focused state

vmPFC ventromedial prefrontal cortex

VS ventral striatum

Vipassanā a mediation type utilizing focused observing, and release and observe

practices.

54
Declaration of generative AI and AI-assisted technologies in the writing process

During the preparation of this work the authors used Chat GPT-4 (OpenAI, 2023) in order to improve the clarity
of the written content. After using this tool/service, the authors reviewed and edited the content as needed and
take full responsibility for the content of the publication.

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 Table 1

Biobehavioral cognitive studies of meditation. Study design, assessment type, practice experience,
meditation type, and summary of major findings of each article. N = number of participants.

Article Study design Assessment Participants Meditation Meditation Findings

experience type
Perception
Fox et al., 2012 Quasi- Objective Practitioners 2,051 hours, Vipassana, Experts
experimental Psychophysical and (n = 38, F = SD = 3,600, 1 body-scan Introspective
cross-sectional Cortical Measures 19) – 15,000 meditation accuracy:
design of Tactile hours; 11.0 ↗ MED: r = .31 – .46
(practitioners) Sensitivity, years, SD = ↗ body-scan
Subjective Measure 10.3 years. meditation: r = .41 –
of Sensitivity body-scan .64
meditation:
154 hours, Beginners
SD = 322, 0 – Introspective
1,643 hours accuracy:
↘ MED: r = –.01 – –
.16
↗ body-scan
meditation: r = .06 –
.18

General
Expertise (hours)
predicting intro.
accuracy:
↗ MED: r = .37 – .48
↗ body-scan
meditation: r = .32 –
.36
↗ MED controlled for
body-scan meditation:
r = .22 – .36

Khalsa et al., Double-blind Body Map by Meditation- 4,947 hours, Vipassana, Between-group
2020 RCT (naïve vs. Drawing, EKG, naïve (n = 15, SD = 6,251; Kundalini differences
experienced) Self-Report F = 5), Experts 10.8 years, Cardiac interoceptive
(n = 15, F = 5) SD = 10.8; awareness:
Retreat: 19 No significant
days, SD = 14 differences found.

Grant et al., Quasi- Computer- Meditation- 14.4 years, Zen: trained in Experts
2010 experimental Controlled Thermal naïve (n = 20, SD = 8.39, 2 mindfulness Pain sensitivity:
repeated- Pain Sensitivity F = 5), Experts – 30 years; meditation ↘ moderate pain
measures design Test (n = 19, F = 4) 6,404 hours, stimulus: p = .002
(naïve vs. SD = 8,522, Expertise predicting
experienced) 1,229 – cortical thickness:
45,000 hours ↗ meditation hours–
S1: r = .69, p < .05
↗ meditation years–
ACC: r = .59, p < .05

Grant & Quasi- Computer- Meditation- 6,247 hours, Zen: trained in Experts
Rainville, 2009 experimental Controlled Thermal naïve (n = 13, SD = 11,789 mindfulness Pain sensitivity:
repeated- Pain Sensitivity F = 5), Experts meditation ↘ moderate pain
measures design, Test (n = 13, F = 5) stimulus: p = .01
counterbalanced ↘ observing and
(naïve vs. release: p = .02
experienced)
Controls
Pain sensitivity:
↗ focused observing:
p < .001

General
Correlations with pain
sensitivity:

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Article Study design Assessment Participants Meditation Meditation Findings
experience type
↘ meditation
experience: r = –.82, p
< .01

Grant et al., Quasi- Computer- Meditation- Not reported Zen Experts

2011 experimental Controlled Thermal naïve (n = 13, Pain sensitivity:
repeated- Pain Sensitivity F = 4), Experts ↘ moderate pain
measures design Test, fMRI (n = 13, F = 4) stimulus: p = .01
(naïve vs. Brain activation
experienced) group-differences:
↘ dlPFC, mPFC/OFC,
amygdala,
hippocampus, IFG,
MFG
↗ dACC, insula,
thalamus
Brain activation
predicting pain
sensitivity:
↘ dACC-dlPFC
connectivity: r = –.71,
p < .001
Brain activation
correlating with
meditation
experience:
↘ years – bilateral
insula: r = -.60 – –.65,
p < .05
↘ years – bilateral
dACC: r = –.69, p <
.01
↘ years–left dlPFC: r
= –.80, p < .001
↘ right mPFC/OFC: r
= –.64, p < .05
↘ hours–thalamus: r =
–.56 – –.62, p < .05

Perlman et al., Quasi- Computer- Meditation- 10,000 – Tibetan Experts

2010 experimental Controlled Thermal naïve (n = 10, 45,000 hours Buddhism: Pain sensitivity:
repeated- Pain Sensitivity F = 5), Experts Kagyu and ↘ observing and
measures design, Test (n = 9, F = 4) Nyingma release: p = .011
counterbalanced Pain unpleasantness:
(naïve vs. ↘ observing and
experienced) release: p = .013

Lutz et al., 2013 Quasi- Computer- Meditation- 27,000 hours, Tibetan Experts
experimental Controlled Thermal naïve (n = 14, SD = 12,500 Buddhism: Pain unpleasantness:
repeated- Pain Sensitivity F = 9), Experts Kagyu and ↘ p = .001
measures design, Test, fMRI (n = 14, F = 9) Nyingma ↘ observing and
counterbalanced release: p < .001
(naïve vs.
experienced) Brain activation
baseline group-
differences:
↘ dorsal anterior
insula, aMCC,
amygdala
Brain activation
during pain group-
differences:
↗ dorsal anterior
insula, aMCC
Baseline brain
activation correlating
with meditation
experience:
↘ left anterior insula: r
= –.63, p < .05

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Article Study design Assessment Participants Meditation Meditation Findings
experience type
Neural habituation
before pain-stimulus:
↗ amygdala, right
posterior insula, right
S2, right mid insula,
MCC
Neural habituation
before pain-stimulus:
↗ right posterior
insula, right S2, right
mid insula

Zorn et al., 2020 Quasi- Computer- Meditation Experts: Tibetan Pain unpleasantness
experimental Controlled Thermal beginner (n = 41,357 hours, Buddhism: Experts:
repeated- Pain Sensitivity 14, F = 9), SD = 17,999, Kagyu and ↘ observing and
measures design, Test, Pain Experts (n = 13,110 – Nyingma release: p = .021
counterbalanced Catastrophizing 14, F = 9) 94535 hours Beginner:
(beginner vs. Scale Beginners: ↘ observing and
experienced) 19.4 hours, release: p = .018
SD = 12.9,
2.2 – 49.2
hours

Emotional
processing
Taylor et al., Quasi- Emotional Intensity Meditation 1,709 hours, Zen General
2011 experimental Ratings beginner (n = SD = 694, Emotional intensity:
repeated- 10, F = 4), 1,000 – 3,000 ↘ all pictures,
measures design Practitioners hours, meditative state: p <
(beginner vs. (n = 12, F = 7) excluding one .05
experienced) outlier of
45,000 hours Experts
Brain activation:
↘ right mPFC: p <
.005
↘ right PCC: p <.005

Beginner
Brain activation:
↘ left amygdala: p <
.005

Lusnig et al., Quasi- Lexical Decision Mediation 7.9 years, SD Zen Mediators
2020 experimental Task, Valence naïve (n = 20, = 4.9, 0.5 – Valence ratings post
repeated- Rating Task F = 9), 28 years; mediation:
measures design Practitioners 1,900 hours ↘ positive, low
(naive vs. (n = 20, F = 9) arousal: p < .001
experienced) ↘ negative, low
arousal: p < .001
↘ negative, high
arousal: p < .001

Chen et al., 2018 Quasi- State and Trait Mediation 12.95 years, Sant Mat: Mediators
experimental Anxiety, Color naïve (n = 20, SD = 6.1, 4 – Loving- State and Trait
repeated- Identification Task, F = 12), 26 years; 4 kindness anxiety:
measures design, Emotion Detection Experts (n = hours daily meditation ↘ state: p < .001
counterbalanced Task, fMRI 21, F = 14) ↘ trait: p = .001
(naive vs. ↘ state anxiety
experienced) correlating with
mediation experience:
r = –.48, p = .001
↘ trait anxiety
correlating with
mediation experience:
r = –.48, p = .001
Color Identification
Task:
↗ RT independent of
attention and emotion:
p = .001
↗ RT, explicit: p < .05

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Article Study design Assessment Participants Meditation Meditation Findings
experience type
↗ accuracy, explicit: p
< .05
Brain activation,
explicit happy:
↗ Amygdala: p < .005
Amygdala activation
mediating reductions
in Anxiety:
↘ fear, explicit: p <
.05

Controls
Brain activation,
explicit fearful:
↗ Amygdala: p < .005
Brain activation,
implicit fearful:
↗ Amygdala: p < .005
Brain activation,
implicit happy:
↗ Amygdala: p < .005

Engen et al., Quasi- Visual Analogue Experts (n = 40,000 hours, Tibetan Compassion
2015 experimental Rating of Affect, 15, F = 5) SD = 9,000, Buddhism: meditation
repeated- Emotion 10,000 – Nyingma Positive affect:
measures Regulation Task, 62,000 hours tradition ↗ compared to all
crossover design, fMRI other conditions: p <
counterbalanced .01
(experienced) Negative affect:
↘ compared to watch-
negative: p < .001
Brain activation,
before stimulus:
↗ mOFC: p < .001
↗ VS/NACC: p < .001
↗ bilateral mid insula:
p < .001

Reappraisal
condition
Positive affect:
↗ compared to watch-
negative: p < .01
Negative affect:
↘ compared to watch-
negative: p < .001
↘ compared to
compassion: p < .01

Lutz et al., Quasi- Human Emotional Mediation 10,000 – Tibetan Experts

2008a experimental Vocalizations, naïve (n = 15, 50,000 hours Buddhism: Negative sounds vs.
repeated- fMRI F = 2), Experts Nyingmapa and others while
measures design (n = 15, F = 2) Kagyupa meditating:
(naive vs. ↗ right insula: p < .05
experienced) Good vs. poor
mediation blocks:
↗ right insula: p < .05
Mediation vs. rest:
↗ right TPJ: p < .001
↗ right pSTS: p < .001
↗ Amygdala: p < .001
↗ PCC/Prc: p < .001
↗ IFG: p < .001
↗ mPFC: p < .001

Kral et al., 2018 Quasi- Automatic Emotion MBSR (n = 9,081 hours, Mindfulness Experts
experimental, Regulation Task, 32, F = 22), 1,439 – meditation, Brain activation:
cross-sectional fMRI Experts (n = 32,612 hours some Loving- ↗ right amygdala, positive: p
longitudinal 30, F = 16) kindness = .001
comparison meditation Lifetime retreat hours–
(experienced vs. observing and release:
vs. MBSR)

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Article Study design Assessment Participants Meditation Meditation Findings
experience type
↘ Amygdala, negative: p =
.02

MBSR
Brain activation:
↗ right amygdala, positive: p
= .01
↗ amygdala-vmPFC
connectivity, positive: p =
.01
↗ amygdala-vmPFC
connectivity, negative: p =
.001

Lutz et al., 2016 Quasi- Self-referential Meditation 5971 hours, Vipassana Experts
experimental Word Processing naïve (n = 22, 506 – 18,805 Questionnaires:
repeated- Task, Five-Facet F = 8), Experts hours. ↗ self-compassion: p < .1
measures design Mindfulness (n = 22, F = Vipassana: ↘ alexithymia: p < .1
(naive vs. Questionnaire, 10) 4861.5 hours, ↗ non-activation: p = .002
experienced) Toronto 281 – 18,325 Affect:
Alexithymia Scale, hours. ↘ affective difference
Self-Compassion between self-praise and self-
Scale, fMRI criticism: p < .05
Brain activation for self-
appraisal conditions:
↗ dmPFC
↘ dmPFC-Prc and dmPFC-
occipital connectivity,
positive
Psychophysiology for self-
appraisal conditions:
↗ dmPFC correlating with
non-activation: r =.49, p =
.03
Non-Ordinary
States of
Consciousness
Berkovich- Quasi- Time Production Meditation ST MM (894 Mindfulness Mindfulness
Ohana et al., experimental Task naïve (n = 12) hours, SD = meditation, meditators
2011 repeated vs. 450), IT MM Transcendental Time duration:
measures design Mindfulness (2,570 hours, meditation ↗ compared to
(naïve vs. meditators (n SD = 471), control, pre med.: p <
experienced) = 36, 12 per LT MM .005
group); (7,556 hours, ↗ compared to
Meditation- SD = 502), control, post med.: p <
naïve (n = 9) TM (16,310 .005
vs. TM (n = hours, SD =
10) 11,970)

Berkovich- Quasi- Specific Meditation Experts (n = 16.5 years, Theravada Experts

Ohana et al., experimental Task, MEG, 12, F = 3) SD = 7.9; 9 – Buddhism: Brain activation,
2013 repeated Phenomenological 34 years; mindfulness Spacelessness/
measures design Analysis 11,225 hours, meditation Timelessness:
(experienced) SD = 9,909, ↗ PCC: p < .05
1,290 – ↗ right TPJ: p < .05
29,290 hours ↗ cerebellum: p < .05

Gutiérrez et al., Quasi- Self-Report Experts (n = 19.2 years, Mindfulness, Experts

2022 experimental Questionnaires, 22, F = 12) SD = 14.83; Vipassana, Zen, Body-boundaries:
repeated Metronome Task 2,994 hours, Tibetan ↘ during meditation: p
measures design, SD = 3,213; Buddhism = .003
counterbalanced 4.0 hours Time-passage:
(experienced) weekly, SD = ↗ during meditation: p
2.08 = .014
Attention to time:
↘ during meditation: p
= .003

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Article Study design Assessment Participants Meditation Meditation Findings
experience type
Nave et al., 2021 Quasi- Interviews, State Experts (n = 3,832 hours, Theravada Experts
experimental and Trait Anxiety 49, F = 19) SD = 4,845, Buddhism Degree of dissolution:
repeated Questionnaire, 5 115 – 24,837 ↗ observing and
measures design, Dimensions- hours release: p < .001
counterbalanced Altered States of ↗ correlation with
(experienced) Consciousness lifetime meditation
Questionnaire, hours: r = .52, p <
Dissociative .001
Experience First-person
Questionnaire perspective:
↗ correlation with
lifetime meditation
hours: r = .49, p < .01
Attention:
↗ correlation with
lifetime meditation
hours: r = .44, p < .01
Agency:
↗ correlation with
lifetime meditation
hours: r = .65, p <
.001

Van Lutterveld Quasi- Effortless Meditation 6,164 hours, Heterogenous, General

et al., 2017 experimental Awareness beginner (n = 1,527 – mostly Effortless awareness:
repeated Neurofeedback 16, F = 5), 50,978 hours Theravada/ ↘ PCC activation: p <
measures Task, EEG Practitioners Vajrayana .0025, extended to
(beginner vs. (n = 16, F = 6) moment-to-moment
experienced) correspondence
Volitional PCC
control:
↗ effortless awareness
meditation: p < .0005

Decision-
making
Kirk et al., 2011 Quasi- Post Ultimatum Meditation- 9.5 years, SD Buddhist Experts
experimental Game Interview, naïve (n = 40, = 7.8 meditation Behavioral:
repeated fMRI F = 21), ↗ asymmetric offers:
measures design Experts (n = p < .01
(beginner vs. 26, F = 10) Brain activation:
experienced) ↗ posterior insula,
unfair: p < .001
↘ anterior insula,
unfair: p < .001
↗ left PCG, rational: p
< .001
↗ left pSTC, rational:
p < .001
↗ bilateral PHG,
rational: p < .001

Controls
Brain activation:
↗ anterior insula,
unfair: p < .001
↗ bilateral dlPFC,
rational: p < .001

Van Vugt & van Quasi- Attentional Meditation 0 – 27 years Vipassana Meditators
den Hurk, 2017 experimental, Network Task, naïve (n = 24, Decision threshold:
cross-sectional Drift Diffusion F = 17), ↗ compared to
longitudinal Model Practitioners controls: p < .05
comparison (n = 24, F = ↗ incongruent: p <
(experienced vs. 17) .001
mediation naïve
vs. MBSR vs. General
active control) Decision threshold:
↗ incongruent trials: p
< .001
↘ main effect of cues:
p < .001

86
Article Study design Assessment Participants Meditation Meditation Findings
experience type
Drift rate:
↘ incongruent trials: p
< .001
Drift variability:
↗ time-space cue: p <
.05
↘ incongruent trials: p
< .05

Note. ACC: anterior cingulate cortex; aMCC: anterior midcingulate cortex; dACC: dorsal anterior
cingulate cortex; MFG: middle frontal gyrus; PHG: parahippocampal gyrus; SFG: superior frontal
gyrus; dlPFC: dorsolateral prefrontal cortex; IFG: inferior frontal gyrus; MCC: midcingulate cortex;
mOFC: medial orbitofrontal cortex; mPFC: medial prefrontal cortex; PCC: posterior cingulate cortex;
PCC/Prc: posterior cingulate cortex/precuneus; S1: primary somatosensory cortex; S2: secondary
somatosensory cortex; VS/NACC: ventral striatum/nucleus accumbens; TPJ: temporoparietal
junction.

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 Table 2

Dynamical processes of the self-pattern.

Elements of the pattern Brief description

Bodily processes Including bio-systemic processes related to motoric, autonomic, endocrine, enteric, immune, interoceptive
functions, supporting homeostasis and a basic distinction between self and non-self.
Prereflective experiental Pre-reflective self-awareness is a structural feature of consciousness constrained by bodily factors; it includes
processes a sense of ownership or mineness, and a sense of agency for intentional action. These processes form what is
sometimes called the minimal self (Gallagher, 2000; Gallagher & Zahavi, 2020; Gallagher & Zahavi, 2021).
Affective processes Including factors ranging from basic bodily affects (e.g., hunger, fatigue) to typical emotion patterns,
existential feelings, and moods (Newen et al., 2015; Ratcliffe, 2008).
Behavioral/action-related Our actions and habitual behaviors which contribute significantly to our self-identity and character
processes (Dewey, 1922; Verplanken & Sui, 2019).
Social/intersubjective Ranging from a basic capacity for attuning to others (de Waal, 2003; Reddy, 2008; Rochat, 2011;
processes Trevarthen, 1979) to a more developed consciousness of self as distinct from others (Mead, 1913;
Sartre, 1969; Taylor, 1989).
Cognitive and Standard theories of personal identity highlight psychological continuity and memory (e.g.,
psychological Processes Shoemaker, 2011); self-related cognitive processes include concepts, beliefs, cognitive dispositions, and
personality traits.
Reflective processes “The ability to consciously reflect on one’s experiences and actions, closely related to notions of autonomy
and moral personhood, including the capacity to evaluate and form second- order volitions about one’s
desires” (Gallagher, 2021, p. 129; see Frankfurt, 1988; Taylor, 1989).
Narrative processes Narrative self-interpretation recursively reflects other processes in the self-pattern. Theories of self-narrative
may include strong claims about how narratives constitute the self (Dennett, 1991; Ricoeur, 1992;
Schechtman, 2011).
Ecological processes “Our embodied-situated actions engage with (and sometimes incorporate) artifacts, instruments, bits and
structures of the environment in ways that define us and scaffold our identities. Situations shape who we are,
and affordances define our possibilities” (Gallagher, 2021, p. 129).
Normative processes Including social and cultural features that are expressed in value-determining norms that define oughts,
obligations and expectations. Self-identity or the sense of who one is, is shaped by everything that comes
along with one’s profession, one’s religion, social status, the various roles involved in marriage, in parenting,
in friendship, as well as constraints imposed by gender, race, and economic circumstances, for better or worse.

Note. Adapted from Gallagher 2021, p. 128.

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