Systematic Review

Neurobiological Changes Induced by Mindfulness and

Meditation: A Systematic Review
Andrea Calderone 1 , Desirée Latella 2, * , Federica Impellizzeri 2 , Paolo de Pasquale 2 , Fausto Famà 3 ,
Angelo Quartarone 2 and Rocco Salvatore Calabrò 2

1 Department of Clinical and Experimental Medicine, University of Messina, Piazza Pugliatti,

98122 Messina, Italy; [email protected]
2 IRCCS Centro Neurolesi Bonino-Pulejo, S.S. 113 Via Palermo, C.da Casazza, 98124 Messina, Italy;
[email protected] (F.I.); [email protected] (P.d.P.);
[email protected] (A.Q.); [email protected] (R.S.C.)
3 Department of Biomedical and Dental Sciences and Morphofunctional Imaging, University Hospital “G.
Martino”—Via Consolare Valeria1, 98125 Messina, Italy; [email protected]
* Correspondence: [email protected]

Abstract: Background and Objectives: Meditation and mindfulness, rooted in ancient traditions,
enhance mental well-being by cultivating awareness and emotional control. It has been shown
to induce neuroplasticity, increase cortical thickness, reduce amygdala reactivity, and improve
brain connectivity and neurotransmitter levels, leading to improved emotional regulation, cognitive
function, and stress resilience. This systematic review will synthesize research on neurobiological
changes associated with mindfulness and meditation practices. Materials and Methods: Studies
were identified from an online search of PubMed, Web of Science, Cochrane Library, and Embase
databases without any search time range. This review has been registered on Open OSF (n) GV2JY.
Results: Mindfulness-Based Stress Reduction (MBSR) enhances brain regions related to emotional
processing and sensory perception, improves psychological outcomes like anxiety and depression,
and exhibits unique mechanisms of pain reduction compared to placebo. Conclusions: This review
highlights that mindfulness, particularly through MBSR, improves emotional regulation and brain
Citation: Calderone, A.; Latella, D.;
structure, reduces anxiety, and enhances stress resilience. Future research should focus on diverse
Impellizzeri, F.; de Pasquale, P.; Famà,
populations and naturalistic settings to better understand and optimize these benefits.
F.; Quartarone, A.; Calabrò, R.S.
Neurobiological Changes Induced by
Keywords: mindfulness; neurobiological correlates; meditation; neurorehabilitation
Mindfulness and Meditation: A
Systematic Review. Biomedicines 2024,
12, 2613. https://doi.org/10.3390/
biomedicines12112613
1. Introduction
Academic Editors: Silvio Ionta and
Meditation is a mental practice aimed at helping each person concentrate his mind
Hideya Kawasaki
and achieve a deep state of relaxation, clear mental conditions, and inner peace. Meditation
Received: 18 September 2024 is rooted deep in ancient spiritual and philosophical traditions, and has a rich history
Revised: 12 November 2024 of propagation spanning thousands of years across various cultures [1]. First adapted
Accepted: 13 November 2024 in religious contexts by Buddhist, Hindu, and Taoist traditions, meditation eventually
Published: 15 November 2024 broke free from these spiritual roots to become a very familiar component of contemporary
mental health and personal development [2]. At the core of meditation lies the cultivation
of a higher state of awareness and control over one’s thoughts and emotions. It trains
the mind to focus on one object, thought, or sensation, which is important for developing
Copyright: © 2024 by the authors.
Licensee MDPI, Basel, Switzerland.
self-regulation and building resilience [3]. One can practice watching thoughts and feelings
This article is an open access article
as they come without being attached to or reacting to them. This creates detachment and
distributed under the terms and calm [4]. Such meditation practiced regularly is likely to increase one’s mental well-being
conditions of the Creative Commons by way of reduction in stress, anxiety, and depression. The non-judgmental awareness
Attribution (CC BY) license (https:// of internal experiences brings about a balanced state of mind and, thus, better mental
creativecommons.org/licenses/by/ health. One is bound to gain emotional stability and develop poises to deal with difficult
4.0/). situations in life [5]. Other than the mental healing benefits, meditation is also practiced for

Biomedicines 2024, 12, 2613. https://doi.org/10.3390/biomedicines12112613 https://www.mdpi.com/journal/biomedicines

Biomedicines 2024, 12, 2613 2 of 21

personal growth, self-realization, and spiritual development. Through this, one can deeply
understand themselves and their places in the world. Therefore, it can help start a journey
toward self-actualization and self-fulfillment [6]. Several practitioners have experienced
deeper insights into things and the feeling of connecting to a larger, all-encompassing
universal consciousness through meditation [7]. Several meditation techniques can be
practiced based on personal preferences and goals: mindfulness meditation, focusing on
current moment awareness, and transcendental meditation involving mantras to ease deep
relaxation. Other forms include Zen meditation, which requires correct body posture
and breathing; loving-kindness meditation cultivates compassion and equanimity toward
oneself and others [8]. Each technique offers unique pathways to mental and emotional
well-being, and understanding these methods can help individuals choose the practice that
best aligns with their personal needs and aspirations [9]. Among the numerous meditation
techniques available that best suit various personal needs and aims, one practice that
has gained much attention is mindfulness meditation. This mental practice emphasizes
that one focuses his or her attention on the present moment, which features an open and
non-judgmental attitude. This practice of mindfulness originated in the ancient traditions
of Buddhism through its roots in contemplation, but today it has spread to wide recognition
and application in modern psychological and medical practice [10]. In general, mindfulness
encourages individuals to witness thoughts, emotions, and physical sensations without
becoming entangled in them [11]. This means the act of allowing whatever arises in the
present moment to come in without judgment or alteration [12]. This is achieved through
meditation processes, where most practitioners sit in silence, focus on breathing or any other
point of focus, and every time the mind begins wandering, it is brought back gently [13].

1.1. Mindfulness and Its Impact on Mental and Physical Health

Mindfulness has been empirically shown to have deep impacts on both mental and
physical health [14]. It reduces stress, anxiety, and depression, but improves well-being and
quality of life [15]. It has been shown that regular mindfulness practice induces structural
changes at the neural level, specifically in cortical thickness in areas related to emotional
regulation and sensory processing. They are usually accompanied by increased functioning
of the mind, which includes reduced anxiety, worry, and depression [16]. Mindfulness
also aids in coping with pain. Studies have shown that mindfulness meditation is signifi-
cantly superior to placebo treatments in reducing both the intensity and unpleasantness
of pain [17]. This effect is believed to be derived from the potential of mindfulness to
change activity within the brain in areas important for pain perception and emotional
processing around the orbitofrontal cortex and anterior cingulate cortex (ACC) [18]. Not
only will some research into mindfulness likely improve psychological and physiological
benefits, but it is also likely to improve social interaction [19,20]. In particular, advanced
brain imaging techniques have shown that mindfulness practitioners increase inter-brain
synchrony during face-to-face interactions. This synchrony is evident at particular brain
wave frequencies and may indicate a high degree of mutual understanding and connection
between people interacting [21]. Deng et al. (2024), in this regard, found that mindfulness
meditation produces strong synchrony in the brains of adolescents, particularly when
they are experiencing multiple emotional states at the same time. The study observed,
through cutting-edge neuroimaging, that meditation increased synchrony in brain regions
typically involved in emotional processing and empathy. This increased synchrony is
related to better emotional regulation and stronger interpersonal bonding [22]. It has also
been applied in a variety of contexts, including addiction treatment and chronic disease
management. For example, mindfulness-based interventions have been found to help
reduce cigarette cravings and consumption in smokers, particularly in women [23,24]. In
the context of chronic diseases, such as multiple sclerosis, mindfulness practices have been
linked to improvements in behavioral outcomes and decreases in inflammatory markers;
however, most studies have failed to find significant changes in physiological measures,
such as cortisol [25–27]. While this has helped to show a reduction in impulsivity and
Biomedicines 2024, 12, 2613 3 of 21

some neural correlates, its effects on these variables are indeed complex and sometimes
even conflicting [28,29]. Long-term meditators may differ from non-meditators in terms
of impulsivity and brain connectivity; however, these findings are not always replicable
across studies [30]. A synthesis of the benefits of mindfulness across different pathologies
is presented in Figure 1. In its MBSR format, mindfulness has developed over the years
into perhaps the best-known and most empirically supported approach to integrating
mindfulness practices into healthcare and therapeutic settings. MBSR is an evidence-based,
structured program for the cultivation of mindfulness through meditation practices and
exercises that aim to develop body awareness. It was developed in 1979 at the University
of Massachusetts Medical School by Dr. Jon Kabat-Zinn. From its conceptualization to
help chronically pain-inflicted patients and those with stress disorders, the breadth of
physical and mental health conditions it currently addresses makes the program one of the
most popular and researched mindfulness interventions in the world [31,32]. The MBSR
program typically runs for eight weeks. For this duration, participants meet once a week
with the group; this group session is about 2.5 h long. Guided mindfulness meditation,
some easy yoga exercises, and body scan exercises are held within the sessions; one allows
time to be directed toward breathing, sensations in the body, thoughts, and emotions
without judgment [33,34]. Supplementing these sessions, participants are asked to commit
to daily mindfulness practice at home, but the recommended duration is approximately
45 min of formal meditation per day [35,36]. One of the central components of MBSR is
the cultivation of a mindful attitude characterized by openness, curiosity, and acceptance
of the present moment—whatever the experience is—whether pleasant, unpleasant, or
neutral. Thus, the attitude adopted places one in a better position to know one’s automatic
responses to difficulties and stressors and to deal with them skillfully, without falling into
impulsive reactions. It has been proven that MBSR is effective in reducing stress symptoms,
anxiety, and depression [37,38]. This program has also been proven efficient in assuaging
chronic pain [39,40]. Many research studies have shown that participants often divulge
greatly improved psychological well-being, like reduced ruminations, enhanced emotional
regulation, and increased resilience [41,42]. In addition, MBSR has been associated with
positive changes in brain structure and function levels, especially regarding attention,
self-mention, and emotion regulation. It has also been tailored for different populations,
including healthcare professionals and students, as well as those in highly stressed 4profes-
Biomedicines 2024, 12, x FOR PEER REVIEW of 22
sions [43]. Clinical situations also use it to augment the conventional treatment of cancer,
cardiovascular diseases, and autoimmune disorders [44].

Figure 1. The benefits of mindfulness across different pathologies.

Figure 1. The benefits of mindfulness across different pathologies.

1.2. Mindfulness and Neuroplasticity: Structural Changes in the Brain

The mindfulness practice fields have been well researched, especially in the last cou-
ple of years, regarding their neurobiological effects. Quite several changes in structure and
function were identified in the brain. The consistent practice of mindfulness meditation
results in neuroplasticity, which brings about observable modifications in different areas
of the brain, associated with managing emotions, focusing, and being conscious of oneself.
Biomedicines 2024, 12, 2613 4 of 21

1.2. Mindfulness and Neuroplasticity: Structural Changes in the Brain

The mindfulness practice fields have been well researched, especially in the last couple
of years, regarding their neurobiological effects. Quite several changes in structure and
function were identified in the brain. The consistent practice of mindfulness meditation
results in neuroplasticity, which brings about observable modifications in different areas of
the brain, associated with managing emotions, focusing, and being conscious of oneself.
It has been demonstrated that increasing the production of brain-derived neurotrophic
factor (BDNF) can support neuroplasticity. A higher amount of BDNF leads to a longer
lifespan, growth of neurons, and synaptic plasticity, enhancing learning and memory. One
of the best revelations that has been well-documented is the increased cortical thickness,
mainly in the prefrontal cortex and anterior cingulate cortex (ACC) [45,46]. Where infor-
mation on executive functions, such as decision-making and problem-solving, is stored,
the prefrontal cortex is involved; moreover, the ACC is involved in critical aspects of
attention and self-regulation [47]. Such structural changes might indicate that mindfulness
strengthens the parts of the brain essential for maintaining control over the physiology of
stress and emotional responses [48]. Another important discovery regards the changes in
the amygdala, a region associated with the processing of emotions, in particular fear and
stress. Specifically, it has been determined that mindfulness can lead to a reduction in size
and reactivity in the amygdala, which is in line with reports of reduced levels of stress and
anxiety [49,50]. This downregulation of the amygdala is also associated with an improved
capacity for the regulation of affective responses, leading to a calmer and more resilient
mind [51]. Mindfulness also appears to connect more robustly and functionally across
different brain regions. Functional magnetic resonance imaging (fMRI) studies have shown
increased connectivity between the prefrontal cortex and the default mode network (DMN),
which is a network of brain areas in self-referential thoughts and mind-wandering [52,53].
In meditators, activity in the DMN is lowered during meditation, which has been connected
to a lesser degree of ruminations and mind-wandering, typical contributors to anxiety and
depression [54,55].

1.3. Mindfulness and Neurochemical Balance: GABA, Serotonin, and BDNF

Mindfulness techniques affect neurotransmitter systems in the brain. For example,
individuals who regularly practice meditation have been found to have higher levels
of GABA. Since the neurotransmitter GABA works to minimize neural activity, more
significant levels of this chemical will help reduce anxiety in an individual while at the
same time improving his or her mood [56,57]. Evidence of higher levels of serotonin
production has also been associated with mindfulness. Serotonin functions in the body to
aid in regulating a person’s mood and general well-being feelings [58,59]. On the other side,
higher levels of BDNF have been related to increased cognitive functioning and emotional
resilience [60–62]. Lastly, the practice of mindfulness is associated with lower cortisol levels,
the major hormone related to stress, a finding that is already by itself enough to recommend
a decrease in bodily stress response [63–65]. This reduction in cortisol, coupled with the
above-mentioned neurobiological alterations, brings out the deep impact of mindfulness
on the brain and the body, thereby impacting total mental health and well-being. These
findings all point to one thing: that mindfulness does not only improve psychological
health but also changes the brain in general and in a long-lasting way, supporting better
emotional regulation, cognitive function, and resilience against stress [66–68]. A summary
of neurobiological and neurotransmitter changes due to mindfulness is shown in Figure 2.
 the above-mentioned neurobiological alterations, brings out the deep impact of mindful-
ness on the brain and the body, thereby impacting total mental health and well-being.
These findings all point to one thing: that mindfulness does not only improve psycholog-
ical health but also changes the brain in general and in a long-lasting way, supporting
Biomedicines 2024, 12, 2613 better emotional regulation, cognitive function, and resilience against stress [66–68]. A
5 of 21
summary of neurobiological and neurotransmitter changes due to mindfulness is shown
in Figure 2.

Figure 2. The neurobiological and neurotransmitter changes following mindfulness.

Figure 2. The neurobiological and neurotransmitter changes following mindfulness.
1.4.
1.4.Mindfulness
MindfulnessTechniques
Techniques and and Their
TheirApplications
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engaging By
in specific phrases and mental images, individuals develop a mindset of love and com-
engaging in specific phrases and mental images, individuals develop a mindset of love
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ers.
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promotes the acceptance of physical sensations. Through the comprehension of these di-
tions and promotes the acceptance of physical sensations. Through the comprehension
verse methods,
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their individual tastes and objectives. Every approach provides different ways to improve
that suits their individual tastes and objectives. Every approach provides different ways
to improve mental and emotional health, underscoring the significance of identifying the
most suitable option for individual requirements and goals.
This systematic review will synthesize research on the neurobiological changes associ-
ated with mindfulness and meditation practices.

2. Materials and Methods

2.1. Search Strategy
A comprehensive literature search was performed using PubMed, Web of Science,
Cochrane Library, and Embase databases, employing the keywords (All Fields: “Mindful-
ness”) AND (All Fields: “Neurobiological correlates”), without any specific search time
range. The PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-Analyses)
flow diagram was utilized to outline the process (identification, screening, eligibility, and
inclusion) for selecting relevant studies, as illustrated in Figure 3. The titles and abstracts
of the database searches were independently reviewed. Articles were evaluated for their
 ness”) AND (All Fields: “Neurobiological correlates”), without any specific search ti
range. The PRISMA (Preferred Reporting Items for Systematic Reviews and Meta-An
yses) flow diagram was utilized to outline the process (identification, screening, eligibili
Biomedicines 2024, 12, 2613
and inclusion) for selecting relevant studies, as illustrated in Figure 3. The titles and
6 of 21
stracts of the database searches were independently reviewed. Articles were evaluated
their relevance based on predefined inclusion criteria. Titles and abstracts that met th
criteria were
relevance basedfully reviewed.
on predefined Multiple
inclusion expert
criteria. teams
Titles independently
and abstracts selected
that met these articles, an
criteria
lyzedfully
were thereviewed.
data to Multiple
minimize bias,
expert andindependently
teams discussed discrepancies
selected articles,until a consensus
analyzed the w
reached. This review has been registered on Open OSF (n) GV2JY.
data to minimize bias, and discussed discrepancies until a consensus was reached. This
review has been registered on Open OSF (n) GV2JY.

Figure 3. PRISMA 2020 flow diagram of evaluated studies.

Figure 3. PRISMA 2020 flow diagram of evaluated studies.
2.2. PICO Evaluation
2.2. PICO Evaluation
We applied the PICO (Population, Intervention, Comparison, Outcome) model to
create the search terms. The population included people participating in different mind-
Weand
fulness applied the PICO
meditation (Population,
practices, includingIntervention, Comparison,
MBSR, performed across bothOutcome) model to c
clinical and
ate the search
nonclinical terms. The
populations. populationoutcomes
Neurobiological included people
will participating
be measured based onin different mindf
neuroimag-
ness and meditation practices, including MBSR, performed across both clinical and no
ing techniques such as fMRI and magnetic resonance imaging (MRI), electrophysiological
clinical populations.
techniques Neurobiological outcomes
such as electroencephalogram (EEG), andwill bebiological
other measured basedlike
markers on neuroimagi
neu-
techniques such as fMRI and magnetic resonance imaging (MRI), electrophysiologi
rotransmitter levels and cortical thickness. The comparison would consist of individuals
not engaging in mindfulness, those receiving standard care, or alternative therapeutic
techniques such as electroencephalogram (EEG), and other biological markers like neu
interventions. The outcome includes neurobiological changes associated with mindful-
transmitter
ness measured levels andneuroimaging
through cortical thickness. The comparison
techniques, wouldassessments,
electrophysiological consist of individuals
and n
other biological markers. Key outcomes include changes in brain structure and function,
neuroplasticity, and alterations in stress-related biomarkers.

2.3. Inclusion Criteria

A study was included if it described or examined the neurobiological changes asso-
ciated with mindfulness and meditation practices. Only articles written in English were
included in this study. Additionally, studies that described or investigated the functional
assessments of these patients were included. We only included studies conducted in human
populations and published in English that met the following criteria: (i) original or protocol
studies of any kind, and (ii) articles that detail the neurobiological changes associated with
mindfulness and meditation practices.
Biomedicines 2024, 12, 2613 7 of 21

2.4. Exclusion Criteria

A study was excluded if it lacked data or information regarding the neurobiological
changes associated with mindfulness and meditation practices. Systematic, integrated, or
narrative reviews were also excluded; however, their reference lists were reviewed and
included when relevant. Additionally, any articles written in languages other than English
were excluded.

3. Results
In total, 151 articles were found: 40 articles were removed due to duplication after
screening, 0 articles were excluded because they were not published in English, and
83 articles were excluded based on title and abstract screening. Finally, 19 articles were
removed after screening for inadequate and untraceable study designs (Figure 3). Nine
research articles met the inclusion criteria and were therefore included in the review. The
studies are summarized in Table 1. The studies discussed in this review analyzed the
neurobiological changes associated with mindfulness and meditation practices. Nine
articles explored the diverse impacts of mindfulness practices on brain activity [69–77].

3.1. The Diverse Impact of Mindfulness Practices on Brain Activity

Researchers have investigated the effects of MBSR on neural and psychological out-
comes, a rather broad field with a focus on changes in brain structure, emotional regulation,
and pain perception. These studies have been conducted using randomized controlled
trials and observational designs and have provided insights into how MBSR enhances
the brain areas generally responsible for emotional processing, sensory perception, and
cognitive control. In the first randomized clinical trial, neuroanatomical changes and
associated changes in psychological variables related to an 8-week MBSR course were
compared between 23 meditation-naïve individuals and a waiting list control group of
comparable age and sex. The cortical thickness of the right insula and somatosensory cortex
was significantly increased in the MBSR trainees. State anxiety, worry, depression, and
alexithymia paralleled these increases with a significant decrease. This was associated with
greater thickness of the right insula and decreased alexithymia [69]. In one study, 75 healthy
volunteers were randomized to one of four interventions: mindfulness meditation versus
placebo conditioning versus sham mindfulness meditation versus book-listening control.
The current study found that all cognitive manipulations significantly reduced pain inten-
sity and unpleasantness relative to the control group. In particular, mindfulness meditation
decreased both pain intensity and unpleasantness significantly more than placebo analgesia
or sham mindfulness meditation. On fMRI, the mindfulness meditation group showed
increased activity in brain regions of a priori interest associated with the cognitive modula-
tion of pain, specifically in the orbitofrontal and anterior cingulate cortices. In contrast, the
neural mechanisms of placebo analgesia were found in sensory processing regions and the
dorsolateral prefrontal cortex. This is the first evidence to date that mindfulness meditation
engages a distinctive mechanism of pain reduction, independent of placebo treatment [70].
Biomedicines 2024, 12, 2613 8 of 21

Table 1. Summary of studies included in the research.

Treatment Outcomes
Author Aim Study Design Sample Size Main Findings Statistical Analyses
Period Measures
The MBSR participants experienced Multivariate pattern
To investigate the
a notable rise in cortical thickness in classification approaches
neuroanatomical and
the right insula and somatosensory were used to identify
psychological changes in
cortex. There was also a notable clusters of brain regions
individuals before and
BDI-II, TAS-20, reduction in psychological indices responsive to MBSR
Santamecchi after an 8-week MBSR Longitudinal 23 meditation-
8 weeks. PSWQ, STAI, related to worry, state anxiety, training. Correlational
et al., 2014 [69] training program, focusing Study. naïve subjects.
MAAS. depression, and alexithymia. analyses were conducted
on both cortical and
Additionally, a correlation was to link neuroanatomical
subcortical brain changes
found between the increase in right changes with
as well as multiple
insula thickness and the decrease in psychological
psychological dimensions.
alexithymia levels. improvements.
Mindfulness meditation
Statistical comparisons of
significantly reduced pain intensity
pain intensity and
and unpleasantness more than
unpleasantness ratings,
To determine whether the placebo analgesia and sham
with significant results
analgesic mechanisms of 75 healthy mindfulness meditation. This pain
Zeidan et al., Randomized indicated by p-values less
mindfulness meditation 4 days. human fMRI. relief was linked to activation in
2015 [70] Controlled Trial. than 0.05. Neural activity
are distinct from those of volunteers. brain regions associated with
associated with each
the placebo effect. cognitive pain modulation, such as
intervention was analyzed
the orbitofrontal, subgenual anterior
through functional
cingulate, and anterior
neuroimaging.
insular cortex.
To evaluate whether MT
Correlational analyses
delivered via a
67 participants, The MT group showed a significant were used to link changes
smartphone app could
with 33 in the correlation between reduced PCC in PCC reactivity with
Janes et al., reduce PCC reactivity to Randomized
One Month. MT group and fMRI. reactivity to smoking cues and a smoking outcomes, with
2019 [71] smoking cues and if Controlled Trial.
34 in the decrease in cigarette consumption, significance levels reported
changes in PCC reactivity
control group. particularly in women. for overall and
were associated with
sex-specific correlations.
smoking reduction.
Biomedicines 2024, 12, 2613 9 of 21

Table 1. Cont.

Treatment Outcomes
Author Aim Study Design Sample Size Main Findings Statistical Analyses
Period Measures
The eight-week mindfulness
intervention did not reduce
To evaluate whether an impulsivity or alter neural correlates
Analyses compared
eight-week mindfulness 105 meditation of impulsivity.
impulsivity and
intervention could reduce naive Long-term meditators reported
Korponay et al., Randomized Eight neurobiological metrics
impulsivity and affect participants and BIS-11. lower attentional impulsivity but
2019 [72] Controlled Trial. weeks. and assessed changes in
related cognitive and 31 long-term higher motor and non-planning
these metrics following the
neural correlates compared meditators. impulsivity compared to
mindfulness intervention.
to control groups. meditation-naïve participants and
exhibited different
neurobiological profiles.
The MBSR program was associated
Statistical analyses
with significant improvements in
involved comparing pre-
various behavioral outcomes and an
and post-MBSR
To explore the enlargement of the right
intervention measures,
psychological, biological, Prospective hippocampus head. The CTRA
Hemond et al., 23 patients MRI, BIPS, including MRI structural
and neuroarchitecture Observational 8 weeks. profile revealed that higher
2024 [73] with MS. DASS-21. changes, behavioral
changes linked to a MBSR Study. inflammatory gene expression
outcomes, and
program in MS patients. correlated with worse anxiety,
inflammatory markers as
depression, stress, and loneliness,
indexed by the
alongside lower eudaimonic
CTRA profile.
well-being.
The MBSR group showed significant
To determine if an improvements in back pain and
Statistical analyses
abbreviated 4-week MBSR somatic-affective depression
compared pre- and
course improves symptoms, along with increased
Pilot pseudoran- post-intervention
Braden et al., symptoms in chronic back regional frontal lobe hemodynamic
domized 4 weeks. 23 participants. BDI-II. measures within and
2016 [74] pain patients and to activity related to emotional
controlled trial. between groups, assessing
examine the neural and awareness. Both groups saw
changes in self-reported
behavioral correlates of improvements in total depression
symptoms and fMRI data.
MBSR treatment. symptoms, but the MBSR group had
unique benefits in specific areas.
Biomedicines 2024, 12, 2613 10 of 21

Table 1. Cont.

Treatment Outcomes
Author Aim Study Design Sample Size Main Findings Statistical Analyses
Period Measures
BA47 showed noticeably higher
levels of activity when participants
repeated a meditation sound in
To assess whether brain The statistical analyses
comparison to when they
activation during relaxed used software for data
Four advanced performed concentrative
Davanger et al., focusing on a meditation Not Specifi- preprocessing and
Clinical Study. male fMRI. meditation-style cognitive tasks.
2010 [75] sound could be cated. modeling the expected
practitioners. Additionally, meditation-specific
distinguished from similar, hemodynamic response
brain activation increased in
concentrative control tasks. with a canonical function.
strength over continuous
meditation bouts rather than
habituating over time.
There was
no
treatment
period
involved, as
the study
HA was negatively correlated with
To explore the neural basis was cross-
sgACC/vmPFC glucose
of personality, particularly sectional in
metabolism, while ST was positively
examining the interplay nature, Region of interest and
Hakamata et al., Observational 140 healthy correlated. Individuals with high
between character and focusing on TCI, PET. whole brain analyses were
2013 [76] Study. adults. HA and high ST had
temperament and their a single conducted.
sgACC/vmPFC glucose metabolism
effects on the sgACC measure-
comparable to those with low scores
and vmPFC. ment point
on both traits.
of brain
glucose
metabolism
and
personality
traits.
Biomedicines 2024, 12, 2613 11 of 21

Table 1. Cont.

Treatment Outcomes
Author Aim Study Design Sample Size Main Findings Statistical Analyses
Period Measures
The study did not replicate previous
To investigate the neural
379 individuals laboratory-based findings linking
correlates of trait mindful
for individual trait mindfulness to individual Pearson correlation
awareness during
brain response brain responses. Nevertheless, it coefficients to assess the
naturalistic social
Naturalistic analysis and was discovered that self-reported relationship between
Chen et al., interactions, focusing on
observational 62 dyads for the EEG. mindfulness was linked to higher mindful awareness scores
2022 [77] how mindfulness affects
design. dyadic levels of dyadic inter-brain and inter-brain coupling
both individual brain
inter-brain synchrony in theta and beta across different
activity and inter-brain
synchrony frequencies (specifically around frequency bins.
synchrony during
analysis. 5–8 Hz and 26–27 Hz) during
face-to-face interactions.
in-person communication.
Legend: Mindfulness-based stress reduction (MBSR), multiple sclerosis (MS), Conserved Transcriptional Response to Adversity (CTRA), Magnetic resonance imaging (MRI), Brief
Inventory of Perceived Stress (BIPS), Depression, Anxiety, and Stress Scale (DASS-21), electrocardiographic (ECG), electroencephalographic (EEG), Beck Depression Inventory-II
(BDI-II), Toronto Alexithymia Scale (TAS-20), Penn State Worry Questionnaire (PSWQ), State-Trait Anxiety Inventory (STAI), Mindful Attention Awareness Scale (MAAS), Functional
Magnetic resonance imaging (fMRI), Bilateral areas of the inferior frontal gyrus (BA47), mindfulness training (MT), posterior cingulate cortex (PCC), Barratt Impulsiveness Scale (BIS-11),
subgenual anterior cingulate cortex (sgACC), ventromedial prefrontal cortex (vmPFC), Temperament and Character Inventory (TCI), positron emission tomography (PET), Harm
avoidance (HA), self-transcendence (ST).
Biomedicines 2024, 12, 2613 12 of 21

3.2. Neurobiological Outcomes of Mindfulness Interventions

In another randomized controlled trial of 67 smokers, 33 were randomized to the
mindfulness training (MT) group and 34 to the active control group. App-delivered MT
significantly reduced posterior cingulate cortex reactivity to smoking cues relative to the
control. This decrease was related to a decrease in cigarette consumption that was not
engendered in the control group for the MT participants. This effect was specifically sex-
related; in women, the reduction in posterior cingulate cortex reactivity was related to
reduced smoking, whereas no such association was observed in men [71]. In an eight-week
mindfulness intervention with 60 participants, mindfulness practice had no significant
effects on impulsivity, as operationalized by the go/no-go task and the Barratt Impulsive-
ness Scale, or on neural correlates of impulsivity (frontostriatal gray matter, functional
connectivity, and dopamine levels) relative to active or wait-list controls. In the go/no-go
task, long-term meditators did not differ from meditation-naïve participants; however, they
showed less attention, and more motor and non-planning impulsivity. Moreover, long-
term meditators had less gray matter in the striatum and higher cortico-striatal-thalamic
connectivity, together with a lower spontaneous eye-blink rate, than meditation-naïve
participants [72]. In an observational study of 23 patients with multiple sclerosis, an MBSR
program improved several behavioral outcomes and increased the size of the right hip-
pocampus. This study also showed that greater inflammatory gene expression, indexed
by the conserved transcriptional response to adversity profile, was associated with worse
patient-reported anxiety, depression, stress, loneliness, and lower eudaimonic well-being.
In contrast, hair cortisol did not differ significantly before and after the MBSR program [73].
Braden et al. conducted a nonrandomized trial with a sample of 12 subjects who received
a shorter, 4-week MBSR course compared to 11 controls (reading material about reduc-
tion of stress) and found significant improvements in overall depression in both groups.
Overall, the MBSR group was the only group able to achieve significant reductions in
back pain and somatic-affective depression symptoms. The MBSR team also demonstrated
significant boosts in frontal lobe blood flow linked to better detection of emotional shifts.
Another inference drawn was that the shortened MBSR course was good for pain relief
and emotional awareness, but the 8-week traditional MBSR course might be necessary
to see wider benefits in anxiety and cognitive aspects of depression [74]. In a study of
long-term meditators of Acem meditation, fMRI was used to explore brain activation at
rest and during the presentation of a meditation sound. Compared with the concentrative
control tasks, it was found that the meditation sound evoked remarkably higher activation
of the bilateral areas in the inferior frontal gyrus. This was not a decreasing activation
over time; on the contrary, activation increased with the length of the meditation sessions.
The findings show that meditation with relaxed focus is associated with parts of unique
activation in the prefrontal cortex, distinguishing it from other cognitive tasks, and increas-
ing knowledge about the neurobiological mechanisms underlying meditation [75]. In a
group of 140 healthy adults, the researchers examined the association between mindful
awareness and brain activity in an extremely localized manner within the prefrontal cortex
and subgenual ACC. Brain glucose metabolism can be used to determine how personality
influences brain function. These findings reverse the effects of temperament on metabolism
in this brain area, which is related to anxiety and vulnerability. In contrast, a personality
trait called self-transcendence, which is associated with spirituality and intuitiveness, has
the opposite effect on the same area of the brain. Furthermore, an analysis of the whole
brain revealed that high spirituality is significantly associated with higher metabolism in
that area of the brain [76]. One last article investigated the neural correlates of mindfulness
in the context of social interactions; they recorded brain activity using a portable EEG while
individuals were engaged in face-to-face conversations as part of an exhibit in a museum.
While failing to replicate earlier work associating specific mindfulness with brain responses
produced in a laboratory environment, they showed that self-reported mindful awareness
was associated with increased inter-brain synchrony between interacting individuals. This
synchrony was visible in both theta and beta frequency bands. This study: therefore,
Biomedicines 2024, 12, 2613 13 of 21

underlines the need for the study of mindfulness under more naturalistic social conditions,
where although individual brain responses may not agree with previous findings on social
traits, the potential of mindfulness in driving inter-brain synchrony when interacting with
others is evident. This further cautions the use of commercially available EEG systems
in research related to the neural correlates of social traits [77]. In summary, MBRS can
enhance brain areas related to emotional processing and sensory perception and improve
psychological outcomes like anxiety and depression.

3.3. The Complexities of Mindfulness: Emotional Regulation and Impulsivity Outcomes

In addition, MBSR can increase the cortical thickness. Mindfulness has dissimilar
neural mechanisms of pain reduction in comparison with placebo treatments for pain
reduction. Mindfulness training also helps reduce smoking cue reactivity, particularly
in women, and enhances emotional regulation. However, its effect on impulsivity is still
unclear, as long-term meditators both provide evidence and fail to provide evidence of its
effects on impulsivity and neural connectivity. It is also the case that mindfulness practice
gives rise to better emotional awareness and social interaction, as shown by enhanced
inter-brain synchrony during naturalistic interactions.

4. Discussion
This systematic review investigated neurobiological changes associated with mind-
fulness and meditation practices. The results highlight the broad benefits of MBSR across
neuroanatomical, psychological, and social domains. MBSR programs, including abbrevi-
ated versions, have been shown to increase cortical thickness in areas such as the right insula
and improve psychological outcomes like reduced anxiety, depression, and alexithymia.
Moreover, mindfulness meditation has a specific effect on reducing pain and improving
emotional awareness; research indicates that it activates different brain mechanisms than
placebo treatments. It also shows promise for smoking cessation, particularly in women,
by lowering brain reactivity to smoking cues. While mindfulness does not consistently
affect impulsivity, it benefits social interactions by increasing inter-brain synchrony during
conversations [69–77]. Therapies, such as MBSR, have demonstrated the ability to induce
neurobiological changes that could directly benefit neurorehabilitation. First, the increases
observed in the cortex, particularly in areas including the right insula and somatosensory
cortex, would indicate improved sensory and interoceptive processing. This could be
particularly useful during neurorehabilitation, as patients often face sensory deficits or
altered body awareness, as in the case of stroke and traumatic brain injury [78,79]. This
is especially true given that the finding that mindfulness practice also modulates pain
through mechanisms separate from placebo effects speaks to its utility in managing chronic
pain conditions that are often comorbid with neurological impairments, including both
spinal cord injury and multiple sclerosis [80,81]. Simpson et al. (2023) targeted participants
with multiple sclerosis, but in their study, they found that mindfulness-based interventions
result in neurobiological changes of relevance for improved quality of life. Functional neu-
roimaging has revealed increased connectivity in the structures responsible for emotional
regulation and brain function, such as the DMN and executive control areas. These changes
were associated with reductions in the symptoms of depression and anxiety, thus proving
that mindfulness alters brain activity in a manner conducive to improved well-being in
multiple sclerosis [82].

4.1. Pain, Stress, & Anxiety

Regarding pain, Hunt et al. (2023) showed that mindfulness-based treatments for
episodic migraines are accompanied by considerable neurobiological changes. As medita-
tion practice continued, decreases in both the frequency and intensity of migraines were
linked to alterations in brain areas known to be involved in pain perception and regula-
tion, such as the ACC and insula. This may mean that mindfulness practices alter neural
pathways involved in the processing of pain, thereby contributing to relief from episodic
Biomedicines 2024, 12, 2613 14 of 21

migraines [83]. Since mindfulness practices reduce pain perception through the prefrontal
cortex and anterior cingulate cortex, interventions could benefit patients through pain man-
agement, as well as through the simultaneous improvement of emotional regulation, which
is an important predictor of rehabilitation outcomes, reducing symptoms of anxiety, stress,
and depression [84–87]. Pagni et al. (2023) were found to report that MBSR had a significant
effect in reducing symptoms of anxiety in adults with autism. They detected neurophysio-
logical changes at the level of brain wave patterns and increased connectivity in the areas of
emotional regulation, by which MBSR might help manage anxiety in autism [88]. Similarly,
a broader perspective on anxiety disorders has provided additional neurobiological insights
related to mindfulness-based interventions. A systematic review conducted by Gerber and
Matuschek (2023) on mindfulness-based interventions concerning anxiety disorders found
that such interventions can bring about significant changes in these neuroanatomical stress
vulnerabilities, including amygdala and prefrontal cortex activation [89]. Based on these
findings, short-term mindfulness interventions have also shown significant neurobiological
effects. Diez et al. (2023) identified the benefit of a short-term mindfulness and compassion
retreat in improving stress reduction and mental well-being. Their study emphasized
the interplay between gene expression pathways activated in stress responses, with an
observed increase in neural activity in regions associated with emotional regulation [90].
Bakshi and Srivastava (2024) explained the neurobiological understanding of yoga and
mindfulness, and their effects on neuronal function, stress, and well-being. Their review
showed that the impact of mindfulness and yoga on the brain areas responsible for reg-
ulating stress, emotional control, and cognitive processes is positive. More specifically,
function is enhanced in areas like the prefrontal cortex and amygdala, which substantiate
emotional resilience and improve cognitive control. They further noted positive effects on
brain health in settings related to addictive disorders and palliative care, which means that
mindfulness might have a bearing on neural circuits underpinning stress response and
reward processing [91]. Siew and Yu (2023) conducted a meta-analysis of structural brain
changes discovered in mindfulness-based randomized controlled trials. They reported that
mindfulness practice is associated with increased cortical thickness and changes in brain
areas responsible for attention and self-regulation [92].

4.2. Healthy Lifestyle and Well-Being

In the context of brain health, Mace et al. (2024) conducted a systematic review of
the effects of mindfulness-based interventions on lifestyle behaviors. Mindfulness-based
interventions, according to their findings, have resulted in neurobiological changes that
underlie a healthy lifestyle. More specifically, mindfulness practices are associated with
better functioning in self-regulation- and executive function-related areas of the brain, such
as the prefrontal cortex. These changes encourage better involvement in health-enhancing
behaviors, and thus foster improved brain health [93]. In addition to their effects on lifestyle,
mindfulness interventions have been studied for their roles in functional connectivity and
neural processes. Hammersjö Fälth and Eklind (2024) conducted a systematic review of
the effects of cognitive behavioral therapy and MBSR on functional connectivity. They
discovered that although both therapies are known to alter brain connectivity, MBSR is
associated with increased connectivity in regions related to self-awareness and parts of
the emotional regulation network, while cognitive behavioral therapy alters regions in-
volved in cognitive control and emotional processing [94]. Overall, the results provide
evidence for the induction by mindfulness and meditation practice of significant neu-
robiological changes at several levels of experiential processing, from cortical structure
and connectivity to emotional and cognitive processing. Since neurological damage is
usually accompanied by psychological trauma, one of the positive roles of mindfulness
interventions in neurorehabilitation may be related to improved mental health, which
can help increase patient motivation and engagement [95]. Such mindfulness practice
can improve emotional resilience during recovery by reducing anxiety, depression, and
alexithymia, thus playing an important role in the long-term rehabilitation process [96,97].
Biomedicines 2024, 12, 2613 15 of 21

Finally, mindfulness was found to enhance functional connectivity in a network of inter-

connected regions that covered cognitive control and self-regulation [98]. In particular,
this may be an important application in cognitive rehabilitation, where executive function
and attentional control must be regained [99,100]. In this respect, improved connectivity
within networks, such as the default mode network and regions of emotional regulation,
may allow for the restoration of cognitive and behavioral functioning in patients [101]. In
summary, neurorehabilitation, in combination with mindfulness, will result in improved
sensory processing, manageable pain, emotional regulation, and cognitive control, all
factors that aim to ensure better recovery and improvement in the quality of life of persons
with neurological impairment [102,103].

4.3. Neurobiological Benefits of Mindfulness Alternatives in Neurorehabilitation

Other options besides MBSR involve practices like yoga, diaphragmatic breathing,
biofeedback, guided imagery, Tai Chi, and Qigong, which influence brain function and struc-
ture. These approaches, while varied, encompass common elements related to mindfulness
concepts, such as self-awareness, self-regulation, and mind-body integration, which have
shown the ability to promote neuroplastic alterations. Understanding these neurobiological
effects is essential to acknowledge the benefits of neurorehabilitation. Yoga, for instance,
includes a blend of physical poses (asanas), breathing exercises (pranayama), and medita-
tion, which activate the brain’s sensory and motor pathways alongside regions associated
with emotional control, such as the prefrontal cortex and limbic system. Neuroimaging
research indicates that yoga activities enhance connectivity between the prefrontal cortex
and limbic areas, which is crucial for managing stress and emotions [104–106]. Diaphrag-
matic breathing influences the autonomic nervous system, especially the parasympathetic
part, by activating the vagus nerve. This activity boosts heart rate variability, which is
associated with better emotional control and decreased physiological stress reactions. The
soothing influence on the autonomic nervous system assists in reducing the sympathetic
“fight or flight” reaction, encouraging a feeling of relaxation and mental clarity. In neurore-
habilitation, diaphragmatic breathing may help to reduce anxiety and boost concentration,
which in turn enhances patient participation in therapy [107,108]. Biofeedback, involving
the immediate tracking of physiological responses, such as heart rate, muscle tension, or
skin conductivity, instructs patients on how to consciously control these reactions using
different methods, including breathing exercises or muscle relaxation techniques. This type
of self-regulation may result in enduring alterations in brain function by enhancing the link
between the brain and body, especially in regions associated with emotional regulation
and executive control. Studies have indicated that biofeedback can increase activity in the
prefrontal cortex, an area involved in decision-making, emotional control, and advanced
cognitive function [109,110]. Guided visualization is an additional method that, while
not explicitly mindfulness, aligns with its fundamental principles. By mentally practicing
tranquil images or imagining preferred results, individuals can stimulate brain regions
linked to emotion control and sensory processing, including the amygdala and the occipital
cortex. Visualization has been demonstrated to encourage neural alterations similar to those
seen in mindfulness practice, creating a calm state that can enhance pain control and reduce
anxiety [111,112]. Tai Chi and Qigong, traditional practices from China that integrate move-
ment, breath management, and meditation, have been discovered to affect brain regions
linked to motor control, balance, and emotional regulation. Tai Chi has demonstrated
the ability to enhance motor coordination and balance, which are crucial for individuals
healing from neurological issues such as stroke or Parkinson’s disease. Neuroimaging
research has shown that Tai Chi enhances brain activation in regions like the basal ganglia,
which govern motor control, and the prefrontal cortex, which is associated with executive
function and focus [113–115]. To summarize, yoga, diaphragmatic breathing, biofeedback,
guided visualization, Tai Chi, and Qigong all encompass neurobiological processes that
promote self-regulation, emotional strength, and neuroplasticity.
Biomedicines 2024, 12, 2613 16 of 21

4.4. Strengths & Limitations

This systematic review has several strengths. It focuses on neurobiological changes
due to the practice of mindfulness and meditation through a thorough database search,
which includes PubMed, Web of Science, Cochrane Library, and Embase. Moreover, it
encompasses a broad range of studies and techniques, such as MBSR, and applies rigorous
criteria for the studies to be included in the review that cater to specific neurobiological
changes through neuroimaging and electrophysiological techniques. This systematic review
is only an overview of research on how mindfulness is linked to brain structures and
functions, thus leading to better pain management, smoking cessation, and outcomes
of chronic diseases. However, reviews of these types have limitations: differences in
study designs, populations, and methodologies limit the drawing of integrated results and
generalization of conclusions. This could compromise the reliability of the findings, as
they rely heavily on small sample sizes in some of the studies and on differences between
short-term practices and the long-term effects of the practices of mindfulness. A focus on
specific populations and settings has implications for the generalizability of the results.
Moreover, disparities related to impulsivity indicate that further research is needed. Finally,
limiting the search to publications in English might have excluded relevant research articles
on the current topic published in other languages and, as a consequence, limited the
review’s comprehensiveness.

5. Conclusions
This review discusses the neurobiological changes attributed to mindfulness and
meditation practices regarding their impact on mental health and changes in brain structure.
It emphasizes increased cortical thickness within emotion regulation and sensory processing
areas, reduced level of anxiety, and improved regulation of emotions. Mindfulness also
enhances the functioning of the brain through improvement in connectivity and increasing
neurotransmitter systems, hence improving mood and reducing anxiety. Future studies
should analyze diverse populations, study designs, and the impact of mindfulness on
a variety of mental health disorders. Advanced neuroimaging and electrophysiological
techniques can be combined to probe the neural mechanisms underlying mindfulness.
Moreover, further research on mindfulness effects within naturalistic settings and cultural
contexts will greatly improve the applicability and understanding of such practices. While
the review recommends that the major benefits of mindfulness on mental and physical
health are involved, more studies are needed to completely understand its neurobiological
effects and maximize the efficacy of its clinical applications.

Author Contributions: Conceptualization, A.C. and D.L.; methodology, F.I.; validation, R.S.C.;
resources, F.F.; data curation, P.d.P.; writing—original draft preparation, A.C.; writing—review and
editing, R.S.C. and A.Q.; visualization, R.S.C.; supervision, A.Q.; project administration, A.Q. All
authors have read and agreed to the published version of the manuscript.
Funding: This study was supported by Current Research Funds 2024, Ministry of Health, Italy.
RCR204-23684848.
Conflicts of Interest: The authors declare no conflicts of interest to disclose.

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