Resting-state fMRI functional connectivity and mindfulness in clinical and non-clinical contexts: A review and synthesis

“The existence of VAN was initially identified through lesion studies notably in stroke patients with spatial hemi-neglect (Corbetta and Shulman, 2002; Vossel et al., 2014).”

“Additional networks are implicated in higher-order processes and include the default mode network (DMN; Raichle et al., 2001), bilateral frontoparietal network (FPN; often called the central executive network [CEN]; Seeley et al., 2007), and salience network (SN; Downar et al., 2002).”

“Seed-based functional connectivity computes correlations between the time-courses of an a priori defined region of interest (ROI) called a “ seed ” to other target regions, which may include all voxels of the entire brain (Biswal et al., 1995).”

“These anatomical and functional ambiguities are diminished when a given taxonomy refers to those networks instead as dorsal frontoparietal attention network (DAN) and lateral frontoparietal control network (FPN; Uddin et al., 2019), with an FPN B subsystem functionally connected to DAN (Dixon et al., 2018).”

“Conversely, the ICA method (Bell and Sejnowski, 1995; Cal-houn et al., 2001) is a whole-brain model-free method that provides a more data-driven and holistic approach to quantifying functional connectivity.”

“The SN is composed of primary nodes of bilateral anterior insular cortex and dACC, which are often used as seeds for seed-based approaches, in addition to other subcortical and limbic structures including the amygdala (Seeley et al., 2007).”

“Additionally, because of statistical complexity, ICA may complicate between-group comparisons and their translation to clinical contexts (Fox and Raichle, 2007; van den Heuvel and Hulshoff Pol, 2010).”

“The DMN (Greicius et al., 2003; Raichle et al., 2001) is comprised primarily of nodes in bilateral medial prefrontal cortex (mPFC), posterior cingulate cortex (PCC), precuneus and medial temporal regions (Greicius et al., 2009).”

“Extensive cytoarchitectural, lesion and neuroimaging evidence implicates a functional distinction in the ACC wherein the dorsal component, dACC, is related to regulation of cognitive processes, and the rostral component, rACC, in regulating emotional processes (Bush et al., 2000; Stevens, 2011).”

“The anterior insula receives interoceptive and external sensory information from other parts of the brain, and has been shown to function as a detector of behaviorally relevant information (Menon and Uddin, 2010).”

“Functional connectivity fMRI has led to the discovery of sets of highly functionally connected brain regions that are often referred to as resting-state brain networks or systems (Damoiseaux et al., 2006).”

“This 8-week program targets stress reduction and was originally designed to help chronic pain patients improve aspects of self-regulation related to pain management (Kabat-Zinn, 1982).”

“Indeed, psychiatric disorders have been associated with modulation of large-scale brain networks (Kaiser et al., 2015; Menon, 2011).”

“…exogenous stimulus to an internally focused endogenous stimulus, and is hypothesized to act as a major gateway or “ circuit-breaker ” linking the Ventral Attention Network (VAN) to the Dorsal frontoparietal Attention Network (DAN; Corbetta et al., 2008; Japee et al., 2015; Uddin et al., 2019).”

“Clinically, mindfulness has been shown to reduce the severity of symptoms of a variety of conditions, including anxiety (Hofmann et al., 2010), post-traumatic stress disorder (PTSD; Boyd et al., 2018), attention-deficit/hyperactivity disorder (ADHD; Poissant et al., 2019), eating disorder…”

“One neuroanatomical region that has been extensively linked to emotion regulation is the ACC (Etkin et al., 2011; Stevens, 2011).”

“…( “ I find myself doing things without paying attention ” ; reverse-scored), non-judgmental inner experience ( “ I disapprove of myself when I have illogical ideas ” ; reverse-scored), and non-reactivity ( “ I perceive my feelings and emotions without having to react to them ” ; Baer et al., 2008).”

“…been proposed that the insular and dACC SN nodes act as a “ switch ” between rsFC of the DMN, which is activated when individuals are not engaging in a cognitively demanding task, and the FPN, which is activated during cognitively challenging tasks that require attention (Sridharan et al., 2008).”

“…builds on Tang et al. ’ s model of pillar concepts of mindfulness improving self-regulation – attentional control, emotional regulation, and self-related awareness - (Tang et al., 2015) in order to define a resting-state functional connectivity (rsFC)-based neurobiological framework of mindfulness.”

“The heterogeneity of these studies is in part due to the ambiguous definition of the construct of mindfulness in the neuroscientific literature (Van Dam et al., 2018), indeed, modern science more broadly has had difficulty precisely defining the concept of mindfulness (Keng et al., 2011).”

“…Walach et al., 2006), Mindful Attention and Awareness Scale (MAAS; MacKillop and Anderson, 2007), Kentucky Inventory of Mindfulness Skills (KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006), Cognitive and Affective Mindfulness Scale Revised (CAMS-R; Feldman et al.,…”

“Indeed, psychiatric disorders have been associated with modulation of large-scale brain networks (Kaiser et al., 2015; Menon, 2011).”

“…structural, functional, and lesion studies leading to parcellation suggest that PCC has highly heterogenous subregions (Bzdok et al., 2015; Leech and Sharp, 2014; Leech and Smallwood, 2019; Scheperjans et al., 2008) and acts as a major hub involving distinct networks (Leech and…”

“However, reverse inferences can suggest new hypotheses and give direction to future experimental testing (Poldrack, 2006).”

“The DMN (Greicius et al., 2003; Raichle et al., 2001) is comprised primarily of nodes in bilateral medial prefrontal cortex (mPFC), posterior cingulate cortex (PCC), precuneus and medial temporal regions (Greicius et al., 2009).”

“That is, the insular cortex is often used to define the SN while its neuroanatomically neighboring region, the IFG/MFG node, (Vossel et al., 2014), is assigned to the VAN (Farrant and Uddin, 2015).”

“The FPN A subsystem is thought to be involved in internally-focused attention and interoceptive processes whereas the FPN B subsystem is thought to be involved in external attention (Dixon et al., 2018; Vincent et al., 2008).”

“Dorsal ACC additionally has been implicated in pain processing and the anticipation of pain (Vogt, 2005), which suggests a possible mechanism for mindfulness meditation training on altered functional connectivity in chronic pain (Su et al., 2016) and pain relief by mind-fulness (Fig.”

“For a long time, neuroscience research considered those “ task-negative low frequency fluctuations ” as noise or random (Deco et al., 2011).”

“While a growing number of studies have investigated therapeutic effects of mindfulness meditation for MDD (e.g., meta-analyzed in Goldberg et al., 2016), only one has investigated the effects of mindfulness meditation on rsFC in MDD. Specifically, Lifshitz and colleagues used a seed-based approach to compare MDD patie…”

“Taken those information together, it is hypothesized that the rlPFC is a flexible hub that facilitates adaptive functional connectivity and switching between networks according to ongoing task demands (Cole et al., 2013; Desrochers et al., 2015; Gilbert et al., 2005).”

“…practices have been linked to improvements in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al., 2014; Howell et al.,…”

“As chronic stress is linked to MDD (Dieleman et al., 2015; Mcewen, 2004; Tafet and Bernardini, 2003; Vyas et al., 2004), the modulation of disordered frontolimbic systems through mindful emotion regulation is a promising approach to non-pharmacologically treat mood and anxiety disorders, and…”

“…Skills (KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006), Cognitive and Affective Mindfulness Scale Revised (CAMS-R; Feldman et al., 2007), Philadelphia Mindfulness Scale (PHLMS; Cardaciotto et al., 2008) and Southampton Mindfulness Questionnaire (SMQ; Chadwick et…”

“…by using self-report questionnaires such as the Five Facet Mindfulness Questionnaire (FFMQ; Baer et al., 2008), Freiburg Mindfulness Inventory (FMI; Walach et al., 2006), Mindful Attention and Awareness Scale (MAAS; MacKillop and Anderson, 2007), Kentucky Inventory of Mindfulness Skills (KIMS;…”

“…Scale (MAAS; MacKillop and Anderson, 2007), Kentucky Inventory of Mindfulness Skills (KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006), Cognitive and Affective Mindfulness Scale Revised (CAMS-R; Feldman et al., 2007), Philadelphia Mindfulness Scale (PHLMS;…”

“To overcome issues related to the similarities of the SN and VAN and the FPN and DAN, increasingly specific and distinct regions could be used as seeds: for instance, TPJ for the VAN and the FEF for the DAN.”

“This may be explained by the fact that DMN-related processes, including mind-wandering and self-reflection, are conceptually opposed to the present-moment awareness component of mindfulness (Brewer et al., 2011).”

“Alternatively, randomized active-controlled studies are considered a gold standard in intervention research (Chalmers et al., 1981).”

“Additionally, the networks described using this framework are thought to have similar roles involving attentional processes, although the FPN is thought to be more so involved in higher-order cognitive processes — including cognitive control — than the DAN (Corbetta and Shulman, 2011).”

“…regions previously described in the literature, including: anterior and posterior cingulate cortex, insular region (Falcone and Jerram, 2018; Fox et al., 2016; H ¨ olzel et al., 2011 ), as well as resting-state large-scale brain networks (H ¨ olzel et al., 2011; Vago and Silbersweig, 2012).”

“In this sense, mindfulness practices have been linked to improvements in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al.,…”

“…together, these results suggest that both FA and OM meditation are associated with distinct patterns of increased activity in regions of the frontal lobe, notably the SMA and rACC; as well as, additionally, the insular region, located deep in the lateral sulcus of the brain (Uddin et al., 2017).”

“This network is linked to top-down control of attention activated by endogenous stimuli, a goal-directed type of attention (Spreng et al., 2010).”

“Because MDD patients exhibit hyperconnectivity of the DMN, especially in relation to the pgACC region (Horn et al., 2010; Sheline et al., 2010), Yang et al. hypothesized that reduced connectivity between pgACC of the rACC and the DMN could be a mechanism for the “ antidepressant ” effect of…”

“Those dysregulation of brain networks are often linked to neuropsychiatric disorders and have been postulated to underlie neuropsychiatric symptoms (Greicius, 2008; Woodward and Cascio, 2015).”

“Importantly, rlPFC has additionally been evidenced to integrate several cognitive processes for a behavioral goal (Ramnani and Owen, 2004).”

“Indeed, rsFC can additionally be used to visualize abnormalities in co-activation patterns (Fox and Greicius, 2010; Lee et al., 2013).”

“This modulation may be related to less self-referential processing and mind-wandering and more cognitive allocation to attentional processes (Andrews-Hanna, 2012; Poerio et al., 2017).”

“As aberrant sgACC-DMN hyperconnectivity has been observed in depression (Connolly et al., 2013; Drevets et al., 2008), it has been speculated that decreased connectivity in these regions mediated by mindfulness training may relate to an antidepressant effect (Yang et al., 2016).”

“Greater mPFC and nucleus accumbens connectivity has been evidence to predict transition from acute to chronic pain (Baliki et al., 2012).”

“Abnormalities in DMN activity and functional connectivity have been linked to psychiatric disorders, including major depressive disorder (Hamilton et al., 2015; Sambataro et al., 2014; Wise et al., 2017).”

“Moreover, future mindfulness and brain network research will benefit from recent developments in approaches in this field (e.g., connectomics; Smith et al., 2013).”

“…(KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006), Cognitive and Affective Mindfulness Scale Revised (CAMS-R; Feldman et al., 2007), Philadelphia Mindfulness Scale (PHLMS; Cardaciotto et al., 2008) and Southampton Mindfulness Questionnaire (SMQ; Chadwick et al., 2008).”

“Spontaneous neural oscillations resulting from evolutionary structural adaptations of high specificity (Buckner et al., 2013), differences in rsFC may indicate underlying differences in brain functioning.”

“Notably, the precuneus is extensively linked to self-referential processing and mind-wandering (Utevsky et al., 2014).”

“Indeed, task-free paradigms offer the advantage of reproducibility across different populations and study designs (Deco et al., 2011; Mulders et al., 2015).”

“In sum, to move beyond reverse inference , these links need to be subsequently studied directly (Chang et al., 2013; Sprooten et al., 2017).”

“However, while the VAN is mainly linked to attentional processes (Corbetta and Shulman, 2002; Vossel et al., 2014), the SN has been associated with processing internally and externally relevant (Seeley, 2019; Seeley et al., 2007).”

“This increased connectivity reported by Parkinson et al. could facilitate the theorized “ switch ” role of the insular node of the SN (Goulden et al., 2014; Sridharan et al., 2008), and thus provide a framework that links attentional networks and mindfulness.”

“…stress disorder (PTSD; Boyd et al., 2018), attention-deficit/hyperactivity disorder (ADHD; Poissant et al., 2019), eating disorder (Wanden-Berghe et al., 2011), substance use disorder (Priddy et al., 2018), and major depressive disorder (MDD; Hofmann et al., 2010; Piet and Hougaard, 2011).”

“One neuroanatomical region that has been extensively linked to emotion regulation is the ACC (Etkin et al., 2011; Stevens, 2011).”

“…” , and — among others — trait mindfulness, state mindfulness, mindfulness meditation training; the lack of adequate control groups in mindfulness training; and the difficulty in choosing adequate neural targets in analyses (Caspi and Burleson, 2005; Davidson, 2010; Davidson and Kaszniak, 2015).”

“Indeed, when activated by a task, dorsal PCC tends to exhibit higher functional connectivity with the FPN (Leech et al., 2011).”

“Notably, the rlPFC (BA 10) is hypothesized to play a role in switching between internally and externally-focused attention (Burgess et al., 2007).”

“…mental object or proprioceptive experience, to open monitoring (OM), which is defined as meta-awareness of present-moment processes (e.g., thoughts, emotions, bodily sensations) without systematically and repeatedly focusing one ’ s attention on any specific object of attention (Lutz et al., 2015).”

“Notably, Uddin and colleagues (Uddin et al., 2019) proposed six large-scale networks referred to with anatomical nomenclature: occipital, pericentral, dorsal fronto-parietal, lateral frontoparietal (FPN), midcingulo-insular, and medial frontoparietal networks.”

“SN has been observed to be involved in the awareness of mind-wandering (Hasenkamp et al., 2012).”

“There is a burgeoning interest in using functional connectivity methods in human neuroscience to advance neural models of myriad aspects of health and disease (Smitha et al., 2017).”

“PTSD is a debilitating condition that is characterized by intrusion and persistence of traumatic memories, as well as avoidance symptoms and negative alterations in cognition and mood, caused by the direct or indirect exposure to a major stressful event (Friedman et al., 2011).”

“Overall, FPN regions have been shown to be involved in sustaining attention through integration of bottom-up perception (Ptak, 2012).”

“The dlPFC is commonly used as an a priori-defined seed region in rsFC analyses to quantify effects related to the FPN (Dixon et al., 2018).”

“Chronic stress is a risk factor for MDD (Hammen, 2018; Yang et al., 2015) and causes increased inflammation that can lead to a reduction of neurogenesis (Schoenfeld and Gould, 2012), an increase in neurotox-icity (Lupien et al., 2018), and increased risk of cardiovascular disease and overall…”

“Emotional processing through awareness and acceptance of negative emotions is a core aspect of mindfulness and mindfulness-based trainings and therapies for psychiatric and physical conditions (Hill and Updegraff, 2012).”

“Emotional distancing cultivated by mindfulness meditation seems to be an effective mechanism of adaptive coping strategy for processing emotions with negative valence (Grecucci et al., 2015; Guendelman et al., 2017; Jones, 2018; Ortner et al., 2007).”

“Given that the PCC is hypothesized to be one of the core regions involved in self-referential processes, autobiographical memory, pro-spection, and planning (Davey et al., 2016; Maddock et al., 2001), these results related to mindfulness may be considered counterintuitive.”

“Given that the PCC is hypothesized to be one of the core regions involved in self-referential processes, autobiographical memory, pro-spection, and planning (Davey et al., 2016; Maddock et al., 2001), these results related to mindfulness may be considered counterintuitive.”

“In this sense, mindfulness practices have been linked to improvements in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al.,…”

“Additional networks are implicated in higher-order processes and include the default mode network (DMN; Raichle et al., 2001), bilateral frontoparietal network (FPN; often called the central executive network [CEN]; Seeley et al., 2007), and salience network (SN; Downar et al., 2002).”

“…(KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006), Cognitive and Affective Mindfulness Scale Revised (CAMS-R; Feldman et al., 2007), Philadelphia Mindfulness Scale (PHLMS; Cardaciotto et al., 2008) and Southampton Mindfulness Questionnaire (SMQ; Chadwick et al., 2008).”

“Several systematic reviews and meta-analyses have suggested a need to revise questionnaires (Baer et al., 2019; Bergomi et al., 2013; Park et al., 2013).”

“In this sense, mindfulness meditation can in this way be understood as a step toward equanimity , that is, a dispositional tendency of evenness of mind towards all experiences, regardless of their emotional valence (Desbordes et al., 2015).”

“Emotional distancing cultivated by mindfulness meditation seems to be an effective mechanism of adaptive coping strategy for processing emotions with negative valence (Grecucci et al., 2015; Guendelman et al., 2017; Jones, 2018; Ortner et al., 2007).”

“While a growing number of studies have investigated therapeutic effects of mindfulness meditation for MDD (e.g., meta-analyzed in Goldberg et al., 2016), only one has investigated the effects of mindfulness meditation on rsFC in MDD. Specifically, Lifshitz and colleagues used a seed-based approach to compare MDD patie…”

“…exogenous stimulus to an internally focused endogenous stimulus, and is hypothesized to act as a major gateway or “ circuit-breaker ” linking the Ventral Attention Network (VAN) to the Dorsal frontoparietal Attention Network (DAN; Corbetta et al., 2008; Japee et al., 2015; Uddin et al., 2019).”

“PTSD is associated with the remodulation of large-scale brain networks.”

“…Questionnaire (FFMQ; Baer et al., 2008), Freiburg Mindfulness Inventory (FMI; Walach et al., 2006), Mindful Attention and Awareness Scale (MAAS; MacKillop and Anderson, 2007), Kentucky Inventory of Mindfulness Skills (KIMS; Baer et al., 2004), Toronto Mindfulness Scale (TMS; Lau et al., 2006),…”

“…stress is a risk factor for MDD (Hammen, 2018; Yang et al., 2015) and causes increased inflammation that can lead to a reduction of neurogenesis (Schoenfeld and Gould, 2012), an increase in neurotox-icity (Lupien et al., 2018), and increased risk of cardiovascular disease and overall mortality…”

“This condition is characterized by long-lasting and continuous pain that is believed to be caused by dysregulation of corticolimbic circuitry involving regions including the PFC, ACC, amygdala and nucleus accumbens (Yang and Chang, 2019).”

“…shown to reduce the severity of symptoms of a variety of conditions, including anxiety (Hofmann et al., 2010), post-traumatic stress disorder (PTSD; Boyd et al., 2018), attention-deficit/hyperactivity disorder (ADHD; Poissant et al., 2019), eating disorder (Wanden-Berghe et al., 2011), substance…”

“…MDD (Hammen, 2018; Yang et al., 2015) and causes increased inflammation that can lead to a reduction of neurogenesis (Schoenfeld and Gould, 2012), an increase in neurotox-icity (Lupien et al., 2018), and increased risk of cardiovascular disease and overall mortality (Kopp and R ´ ethelyi, 2004).”

“Mindfulness training programs usually provide aspects of both FA and OM (Williams and Kabat-Zinn, 2011), and thus the unique effects of each practice are difficult to”

“As the dACC is implicated in control of attentional processes (Benedict et al., 2002; Tian et al., 2006; Weissman et al., 2005) and the PCC in internally directed cognition (Leech and Sharp, 2014), these findings may be related to control of attention and self-reflection.”

“…with the FPN and increased communication with the SN (Yang et al., 2016b), concomitant to insular functional connectivity dysfunctions (Manoliu et al., 2013) overall decreased PCC/caudate nucleus coupling (Bluhm et al., 2009), as well as decreased interhemispheric coupling (Guo et…”

“Several systematic reviews and meta-analyses have suggested a need to revise questionnaires (Baer et al., 2019; Bergomi et al., 2013; Park et al., 2013).”

“Indeed, methodological issues arise from a number of issues including the inconsistent and broad meaning applied for the term “ mindfulness ” , and — among others — trait mindfulness, state mindfulness, mindfulness meditation training; the lack of adequate control groups in mindfulness training; and the difficulty in…”

“…coupling to rACC are aberrant in MDD (Disner et al., 2011), and connectivity dysfunctions of rACC and the amygdala component of limbic circuitry are prevalent in mood disorders (Alexander et al., 2020; Connolly et al., 2013; Hakamata et al., 2020; Kamphausen et al., 2012; Scharnowski et al., 2020).”

“Similarly, increased DMN PCC to dACC SN connectivity was observed after MBSR-based training (Yang et al., 2016) and after an intensive retreat in undiagnosed participants (Kwak et al., 2019).”

“…fMRI activation studies are highly heterogenous in terms of the type of meditation practiced during fMRI acquisition by participants with different experience levels (Falcone and Jerram, 2018; Fox et al., 2016), and rarely relate to changes mediated by trait mindfulness (Lutz et al., 2014).”

“The dACC region is involved in attentional control ( Bush, 2011) and pain cognition and processing has been shown to be modified by attention monitoring (Kabat-Zinn, 1982).”

“For example, findings from Creswell and collaborators ’ study (Creswell et al., 2016) also suggest that increased functional coupling of the PCC node of the DMN and dlPFC node of the FPN is linked to reduced psychological symptoms in individuals with chronic stress.”

“Mindfulness meditation training has been shown to down-regulate activity of the DMN (Brewer et al., 2011; Garrison et al., 2015).”

“…with the FPN and increased communication with the SN (Yang et al., 2016b), concomitant to insular functional connectivity dysfunctions (Manoliu et al., 2013) overall decreased PCC/caudate nucleus coupling (Bluhm et al., 2009), as well as decreased interhemispheric coupling (Guo et al., 2013).”

“…structural, functional, and lesion studies leading to parcellation suggest that PCC has highly heterogenous subregions (Bzdok et al., 2015; Leech and Sharp, 2014; Leech and Smallwood, 2019; Scheperjans et al., 2008) and acts as a major hub involving distinct networks (Leech and Small-wood, 2019).”

“In sum, these two studies show similar effects of mindfulness meditation training on functional connectivity of dACC and rostral ACC regions (Kwak et al., 2019; Yang et al., 2016).”

“Yang and collaborators interpreted the reduced pgACC-DMN connectivity after mindfulness training to the ones found when comparing healthy controls after receiving antidepressant medication ( Scheidegger et al., 2012).”

“In this sense, mindfulness practices have been linked to improvements in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al.,…”

“As chronic stress is linked to MDD (Dieleman et al., 2015; Mcewen, 2004; Tafet and Bernardini, 2003; Vyas et al., 2004), the modulation of disordered frontolimbic systems through mindful emotion regulation is a promising approach to non-pharmacologically treat mood and anxiety disorders, and…”

“Participants who followed mindfulness training, compared to active control participants, had increased functional connectivity between the PCC and the left dlPFC node of the FPN and relatively decreased levels of IL-6 at 4-month follow-up.”

“Indeed, this burgeoning field is aimed toward characterizing inter-regional synchronized low frequency ( 0.1 Hz) spontaneous fluctuations during resting, or non-task, states (Snyder and Raichle, 2012).”

“Abnormalities in DMN activity and functional connectivity have been linked to psychiatric disorders, including major depressive disorder (Hamilton et al., 2015; Sambataro et al., 2014; Wise et al., 2017).”

“…in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al., 2014; Howell et al., 2008; Smith et al., 2015) in healthy populations.”

“Theoretically, this design is supported by the argument that long-term meditation practice leads to changes in brain connectivity and altered trait levels of mindfulness (Luders et al., 2011).”

“Those dysregulation of brain networks are often linked to neuropsychiatric disorders and have been postulated to underlie neuropsychiatric symptoms (Greicius, 2008; Woodward and Cascio, 2015).”

“Given this role, and taking all of this into account, the FPN is sometimes theorized to facilitate mindful present-moment interactions with the environment (Hasenkamp et al., 2012; Kajimura et al., 2020; Taren et al., 2017; Vago and Zeidan, 2016).”

“Reviews and meta-analytic syntheses have summarized relations between trait mindfulness and its relation to psychological and behavioral features (Keng et al., 2011; Mesmer-Magnus et al., 2017; Sala et al., 2019).”

“Taken those information together, it is hypothesized that the rlPFC is a flexible hub that facilitates adaptive functional connectivity and switching between networks according to ongoing task demands (Cole et al., 2013; Desrochers et al., 2015; Gilbert et al., 2005).”

“The dACC has been implicated in response selection and conflict monitoring (Ide et al., 2013; Menon, 2015).”

“This differential functional connectivity relating to trait mindfulness could be due to a lesser need for mindful individuals to suppress basal DMN activity, which has been related to mind-wandering (Poerio et al., 2017).”

“In sum, to move beyond reverse inference , these links need to be subsequently studied directly (Chang et al., 2013; Sprooten et al., 2017).”

“Lesion studies have determined a marked laterization of the VAN on the right hemisphere (Bartolomeo and Seidel Malkinson, 2019), The VAN has been shown to be involved in the orientation of attention towards unpredicted external exogenous stimuli (Vossel et al., 2014).”

“Emotional distancing cultivated by mindfulness meditation seems to be an effective mechanism of adaptive coping strategy for processing emotions with negative valence (Grecucci et al., 2015; Guendelman et al., 2017; Jones, 2018; Ortner et al., 2007).”

“That is, the insular cortex is often used to define the SN while its neuroanatomically neighboring region, the IFG/MFG node, (Vossel et al., 2014), is assigned to the VAN (Farrant and Uddin, 2015).”

“Notably, ventral PCC has been shown to communicate with vmPFC, a major node of the DMN, whereas the dorsal subregion of PCC has been related to increased connections to the dlPFC, a core node of the FPN (Bzdok et al., 2015; Fan et al., 2018).”

“While the function of the STG and its relations with the DMN remain unclear, some evidence suggests that this region is implicated in visuotemporal attention (Shapiro et al., 2002).”

“For example, with regard to the widely used FFMQ, only the non-judging subscale has been shown to exhibit strong validity, while the observing, describing, acting with awareness and non-reactivity subscales have not (Goldberg et al., 2016; Mattes, 2019).”

“It has been hypothesized that individuals who are more mindful will exhibit functional connectivity patterns that are similar to those who practice mindfulness meditation (Wheeler et al., 2017).”

“…al., 2018; Parkinson et al., 2019), cross-sectional (Bauer et al., 2019; Froeliger et al., 2012) and controlled longitudinal studies in undiagnosed (Doll et al., 2015; Kilpatrick et al., 2011; Kral et al., 2019; Kwak et al., 2019; Yang et al., 2016) and disordered populations (Creswell et al.,…”

“Indeed, higher rsFC changes of DMN and FPN have been associated with higher cognitive flexibility and cognitive performance, overall (Douw et al., 2016).”

“Given this role, and taking all of this into account, the FPN is sometimes theorized to facilitate mindful present-moment interactions with the environment (Hasenkamp et al., 2012; Kajimura et al., 2020; Taren et al., 2017; Vago and Zeidan, 2016).”

“There are interindividual differences in vulnerability and susceptibility to the development of PTSD after a trauma (Bomyea et al., 2012).”

“…with the FPN and increased communication with the SN (Yang et al., 2016b), concomitant to insular functional connectivity dysfunctions (Manoliu et al., 2013) overall decreased PCC/caudate nucleus coupling (Bluhm et al., 2009), as well as decreased interhemispheric coupling (Guo et al., 2013).”

“Abnormalities in DMN activity and functional connectivity have been linked to psychiatric disorders, including major depressive disorder (Hamilton et al., 2015; Sambataro et al., 2014; Wise et al., 2017).”

“While a growing number of studies have investigated therapeutic effects of mindfulness meditation for MDD (e.g., meta-analyzed in Goldberg et al., 2016), only one has investigated the effects of mindfulness meditation on rsFC in MDD. Specifically, Lifshitz and colleagues used a seed-based approach to compare MDD patie…”

“2) (Zotev et al., 2013), dysfunctions are frequently linked to mood disorders.”

“Dysregulated sgACC function is often observed in mood disorders, notably MDD (Ge et al., 2020; Ho et al., 2014).”

“While a growing number of studies have investigated therapeutic effects of mindfulness meditation for MDD (e.g., meta-analyzed in Goldberg et al., 2016), only one has investigated the effects of mindfulness meditation on rsFC in MDD. Specifically, Lifshitz and colleagues used a seed-based approach to compare MDD patie…”

“…MDD (Hammen, 2018; Yang et al., 2015) and causes increased inflammation that can lead to a reduction of neurogenesis (Schoenfeld and Gould, 2012), an increase in neurotox-icity (Lupien et al., 2018), and increased risk of cardiovascular disease and overall mortality (Kopp and R ´ ethelyi, 2004).”

“Stronger temporal relations are generally interpreted as indicating greater functional connectivity, or interaction, between regions (Mohanty et al., 2020).”

“They focused on the DAN, and found increased functional connectivity in experienced practitioners within the DAN related to IPS and the FEF nodes (Froeliger et al., 2012) as well as the visual area MT. Previous findings in rsFC suggest interactions between visual areas and the DAN (Yeo et al.,…”

“It is hypothesized that resting with eyes closed would increase functional connectivity in the retrosplenial cortex, implicated in scene viewing (McAvoy et al., 2008).”

“Given the role of these regions in emotion processing (LeDoux, 1994), the observed changes in functional coupling may provide a neural signature of improved emotion regulation.”

“Taken those information together, it is hypothesized that the rlPFC is a flexible hub that facilitates adaptive functional connectivity and switching between networks according to ongoing task demands (Cole et al., 2013; Desrochers et al., 2015; Gilbert et al., 2005).”

“…in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al., 2014; Howell et al., 2008; Smith et al., 2015) in healthy populations.”

“…stress disorder (PTSD; Boyd et al., 2018), attention-deficit/hyperactivity disorder (ADHD; Poissant et al., 2019), eating disorder (Wanden-Berghe et al., 2011), substance use disorder (Priddy et al., 2018), and major depressive disorder (MDD; Hofmann et al., 2010; Piet and Hougaard, 2011).”

“Future studies may benefit from more explicit descriptions of the type of mindfulness training studied or explicitly dismantling components of mindfulness training (Britton et al., 2018).”

“…coupling to rACC are aberrant in MDD (Disner et al., 2011), and connectivity dysfunctions of rACC and the amygdala component of limbic circuitry are prevalent in mood disorders (Alexander et al., 2020; Connolly et al., 2013; Hakamata et al., 2020; Kamphausen et al., 2012; Scharnowski et al., 2020).”

“Specific training programs have varied, and included MBSR (Kilpatrick et al., 2011; Kral et al., 2019; Su et al., 2016), variants of MBSR (Yang et al., 2016), study-specific audio recordings of mindfulness meditation training (Doll et al., 2015; Lifshitz et al., 2019), mindfulness meditation…”

“Notably, PTSD is evidenced to be related to within-DMN hypoconnectivity, decreased connectivity between the DMN and affective systems, as well as increased connectivity between DMN and the somatomotor network (Bao et al., 2021).”

“…structural, functional, and lesion studies leading to parcellation suggest that PCC has highly heterogenous subregions (Bzdok et al., 2015; Leech and Sharp, 2014; Leech and Smallwood, 2019; Scheperjans et al., 2008) and acts as a major hub involving distinct networks (Leech and Small-wood, 2019).”

“Dysregulation of the processing of painful and negative emotions, and pain avoidance, through disordered behavior is related to a variety of psychiatric conditions including MDD and PTSD (Asmundson et al., 1999; Xie et al., 2014).”

“In Pali, the term Sati was adopted and translated as mindfulness in western cultures (Sharf, 2014).”

“Studies additionally differed with respect to design, including: correlational (Bilevicius et al., 2018; Parkinson et al., 2019), cross-sectional (Bauer et al., 2019; Froeliger et al., 2012) and controlled longitudinal studies in undiagnosed (Doll et al., 2015; Kilpatrick et al., 2011; Kral et al.,…”

“…coupling to rACC are aberrant in MDD (Disner et al., 2011), and connectivity dysfunctions of rACC and the amygdala component of limbic circuitry are prevalent in mood disorders (Alexander et al., 2020; Connolly et al., 2013; Hakamata et al., 2020; Kamphausen et al., 2012; Scharnowski et al., 2020).”

“…including anxiety (Hofmann et al., 2010), post-traumatic stress disorder (PTSD; Boyd et al., 2018), attention-deficit/hyperactivity disorder (ADHD; Poissant et al., 2019), eating disorder (Wanden-Berghe et al., 2011), substance use disorder (Priddy et al., 2018), and major depressive disorder…”

“For example, with regard to the widely used FFMQ, only the non-judging subscale has been shown to exhibit strong validity, while the observing, describing, acting with awareness and non-reactivity subscales have not (Goldberg et al., 2016; Mattes, 2019).”

“…in cognitive processes (Chiesa et al., 2011; Gallant, 2016; Malinowski, 2013), stress-management (Chiesa and Serretti, 2009), social cognition (Campos et al., 2019; Tan et al., 2014), and general well-being (Campanella et al., 2014; Howell et al., 2008; Smith et al., 2015) in healthy populations.”

“…coupling to rACC are aberrant in MDD (Disner et al., 2011), and connectivity dysfunctions of rACC and the amygdala component of limbic circuitry are prevalent in mood disorders (Alexander et al., 2020; Connolly et al., 2013; Hakamata et al., 2020; Kamphausen et al., 2012; Scharnowski et al., 2020).”

“Disruption of amygdala and ventrolateral prefrontal cortex (vlPFC) circuitry has been extensively highlighted in populations with symptoms of anxiety and depression, especially in younger participants (Fowler et al., 2017; Greenberg et al., 2017; Guyer et al., 2008).”

“…of functional connectivity is related to years of mindfulness practice (Bauer et al., 2019; Froeliger et al., 2012), and a meta-analysis found unique functional connectivity patterns when comparing results of studies of novice and experienced meditation practitioners (Falcone and Jerram, 2018).”

“In sum, these two studies show similar effects of mindfulness meditation training on functional connectivity of dACC and rostral ACC regions (Kwak et al., 2019; Yang et al., 2016).”

“Several systematic reviews and meta-analyses have suggested a need to revise questionnaires (Baer et al., 2019; Bergomi et al., 2013; Park et al., 2013).”

“…assessed with questionnaires (Bilevicius et al., 2018; Parkinson et al., 2019), as well as in meditators compared to non-meditators (Bauer et al., 2019), decoupling between the DMN and the STG region of the VAN has been observed (Bauer et al., 2019; Bilevicius et al., 2018; Parkinson et al., 2019).”

“First, decreased functional connectivity between the cuneus and the SN has been related to trait mindfulness (Bilevicius et al., 2018).”

“…to altered and dysfunctional functional connectivity in the PCC, with weakened communication with the FPN and increased communication with the SN (Yang et al., 2016b), concomitant to insular functional connectivity dysfunctions (Manoliu et al., 2013) overall decreased PCC/caudate nucleus…”

“However, some studies suggest that the PCC has subregions that exhibit higher specificity, and, furthermore, functional connectivity may not always mirror activity (Lynch et al., 2018).”

“The cuneus, along with other midline cortical structure, was linked to reduced activation during mindful self-awareness (Lutz et al., 2016).”

“As the dACC is implicated in control of attentional processes (Benedict et al., 2002; Tian et al., 2006; Weissman et al., 2005) and the PCC in internally directed cognition (Leech and Sharp, 2014), these findings may be related to control of attention and self-reflection.”

“Multiple studies have reported changes in connectivity strength following a mindfulness training, both in structural [15] and functional connectivity [16].”

“In a previous review of functional connectivity in mindfulness meditation, Sezer et al. (2022) observed that meditation-related changes in self-awareness are linked to decreased connectivity between the SN and the cuneus.”

“Moreover, changes in functional connectivity in the frontoparietal, salience, and default mode networks during meditation are associated with increased attention and reduced occurrences of mind-wandering and self-referential processing [ 100].”

“The functional roles of DMN and ECN in mindful breathing and body-scan await further investigations.”

“, 2017), functional magnetic resonance imaging (fMRI) (Engström et al., 2022; Li et al., 2022; Sezer et al., 2022; Snyder et al., 2022), functional near-infrared spectroscopy (fNIRs) (Bergen-Cico et al.”

“…brain networks widely implicated in self-referential processing (default-mode network), awareness of bodily sensations (salience network), and attentional control (control network) during other meditation techniques involving focused attention (Ganesan, Beyer, et al., 2022; Sezer et al., 2022).”

“Several studies on the resting state have shown that prolonged and regular meditation practitioners have altered DMN FC compared to novices 67,69,77 .”

“This theory is supported by studies reporting that mindfulness- induced emotional regulation is associated with the corticolimbic system, including the prefrontal cortex and amygdala (Sezer et al., 2022).”

“Previous studies have frequently cited the effects of mindfulness and meditation on the subregions of the cingulate (Zsadanyi, Kurth, & Luders, 2021) in terms of local activity (Fox et al., 2014, 2016) and connectivity (Sezer, Pizzagalli, & Sacchet, 2022).”

“Second, these studies aimed to understand relationships among neural activation patterns and psychological processes involved in meditation, including the assessment of behavior, cognition, and emotion (40,41).”

“We concluded that OCD patients showed local brain functional changes, including higher activation in the left DLPFC (BA9), left IFG and left STG, as well as lower activation in the right IPL, right PCC (BA29) and bilateral caudate.”

“…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).”

“…resonance imaging (fMRI) during focused attention meditation ( Fox et al., 2016; Ganesan et al., 2022), changes observed in networks after mindfulness training ( Rahrig et al., 2022; Sezer et al., 2022), in addition to a robust cognitive neuroscience literature on these networks ( Menon, 2011).”

“Some regions overlap with what traditionally is referred to as the default mode (DMN) network (Raichle, 2015; Sezer et al., 2022), a network of brain regions that is activated during rest phases of a task, which has been linked to self-referential thought (Davey et al., 2016).”

“In addition, to compare our fi ndings with more conventional connectivity approaches (44), and suggested by recent studies (22,26), we derived 2 other metrics, the between-network dispersion and the within-network dispersion.”

“…involvement of the default mode network (DMN), salience network (SN), and central executive network (CEN) in studies evaluating the neurobiological impact of mindfulness (Melis et al., 2022; Rahrig et al., 2022; Sezer et al., 2022), though precise underlying neural mechanisms remain unclear.”

“…studies have revealed the involvement of specific brain regions and functional connections in FA, typically associated with cognitive processes including conflict monitoring, interoception, and cognitive control ( H ö lzel et al., 2011; Lutz et al., 2008; Sezer et al., 2022; Treves et al., 2024).”

“Mindfulness decreases default mode network and frontoparietal network connectivity, and changes in medial prefrontal-amygdala connectivity are critically implicated in the regulation of emotion, which may be related to improved emotional regulation [52, 53].”

“41-42 These findings were interpreted as a dissolution of the traditional sense of self (DMN) with advanced meditators and cognitive effort to maintain the meditative state (ECN integrity) in novices.”

“While there is a growing literature on brain connectivity in mindfulness and meditation (Sezer et al., 2022; Treves, Pichappan, et al., 2024), only a few studies have applied DFC to meditation, and none to advanced meditation.”

“Resting-state fMRI (rs-fMRI) metrics such as Regional Homogeneity (ReHo) and Amplitude of Low-Frequency Fluctuation (ALFF) offer unique insights into early inflammation-driven neural reorganization preceding functional decline [19].”

“…in particular that mindfulness practices induce connectivity changes between SN, DMN and CEN in experienced meditators (Brewer et al., 2011; Sezer et al., 2022), as well as in novice meditators after training (Bremer et al., 2022; Creswell et al., 2016; Farb et al., 2007, 2013; Kral et…”

“Additionally, studies on the neurophysiological mechanisms have shown that mindfulness can influence default mode (DMN), frontoparietal (FPN), and salience (SN) networks, explaining its favorable effects (70).”

“…positive effects of meditation, including the roles of self-awareness, attentional control, and emotion regulation (Abellaneda-Pérez et al., 2024; Sezer et al., 2022), and have informed theoretical accounts of how traditional descriptions of meditative phenomena can be reconciled with…”

“…have also been observed in the functional connectivity between: 1) the default mode network, which is involved in self-awareness and introspection, and 2) brain networks that support cognitive control and interoceptive awareness (the central executive control and salience networks) [142, 143].”

“Lines of treatment include standard pharmacological treatments with the following treatment options – rehabilitation, therapy, restoration, maintenance, and surgery (Sezer et al., 2022).”

“Emerging evidence also highlights inter-individual variations in self-report measures of dispositional mindfulness (33,39,42 – 44) and differences in meditation experience between adept practitioners and novices (33,34,39,45 – 48) that in fl uence functional connectivity (FC) and activation changes…”

“…in turn impact myriad functions including enhanced cognitive and emotional functioning; reduction of clinical symptoms like anxiety, depression, and PTSD; and increases in compassion (Schuman-Olivier et al., 2020; Seppälä et al. 2014; Sezer et al., 2022; Whitfield et al., 2022; Young et al., 2018).”

“These brain regions are core nodes of large-scale neurocognitive networks, suggesting that mindfulness training affects activity in a set of distributed brain regions operating within larger networks (Sezer et al., 2022).”

“Since the high heritability of structural and functional brain connectivity in families with a history of ADHD (Sudre et al., 2017) and for consistent results with weaker FC in DMN-FPN and DMN-VAN among subjects with ADHD symptoms (Castellanos & Aoki, 2016), we might infer that an increase in FC in DMN-FPN and DMN-VAN…”

“The interaction of these 3 networks (and their constituent brain regions) contributes to mindfulness meditative states that have been explored more in recent reviews (21,22).”

“Similar analysis with LEC did not find significant differences on score changes of SCS and SCS-SJ between those above and below median LEC scores.”

“Extensive mindfulness practice also leads to changes in the functional connectivity of the default mode and salience networks associated with enhanced self-awareness and attention/ emotion regulation 49,51 .”

“…on resting-state FC. Resting-state FC effects of meditation typically include PCC-DLPFC positive coupling (Creswell et al., 2016; King et al., 2016; Kral et al., 2019; Sezer et al., 2022) mediated by a positive integration with the awareness-related SN (Bremer et al., 2022; Rahrig et al., 2022).”

“Reduced functional connectivity within the salience network has been associated with experiential acceptance and present-moment awareness (Sezer et al., 2022).”

“Third, the default mode network (DMN) is associated with self-referential and social-cognitive processing [27].”

“71 However, this study found that mindfulness was negatively associated with depression, rather than anxiety.”

“26,27 Meditation also is associated with change in brain functional connectivity or plasticity on functional resonance imaging, 28,29 perhaps suggesting that meditation reverses maladaptive plasticity through cortical reorganization.”

“MBSR has been reported to improve the activation and connectivity of the areas including the frontal cortex, which are involved in emotion regulation and self-referential processing 30–33 .”

“Functional MRI studies have shown that mindfulness does not simply increase or decrease brain activity, but rather modulates brain connectivity of the corticolimbic circuits, resulting in well-organized functioning in clinical samples.(66) Therefore, mindfulness, which helps patients reestablish the ability to navigat…”