Brain Research Bulletin 82 (2010) 4656

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Brain Research Bulletin

journal homepage: www.elsevier.com/locate/brainresbull

Research report

Neural correlates of focused attention and cognitive monitoring in meditation

Antonietta Manna a,b,,1 , Antonino Raffone c,e,1 , Mauro Gianni Perrucci a,b , Davide Nardo c,d ,
Antonio Ferretti a,b , Armando Tartaro a,b , Alessandro Londei c,d , Cosimo Del Gratta a,b ,
Marta Olivetti Belardinelli c,d , Gian Luca Romani a,b
a

ITAB, Institute for Advanced Biomedical Technologies, G. DAnnunzio University Foundation, Chieti, Italy
Department of Clinical Sciences and Bioimaging, University of Chieti, Chieti, Italy
Department of Psychology, La Sapienza University, Rome, Italy
d
ECONA (Interuniversity Center for Cognitive Processing in Natural and Articial Systems), Rome, Italy
e
Perceptual Dynamics Laboratory, RIKEN Brain Science Institute, Wako-shi, Saitama, Japan
b
c

a r t i c l e

i n f o

Article history:
Received 4 September 2009
Received in revised form 1 March 2010
Accepted 2 March 2010
Available online 16 March 2010
Keywords:
Meditation
Attention
Consciousness
Prefrontal cortex
Cognitive control

a b s t r a c t
Meditation refers to a family of complex emotional and attentional regulatory practices, which can be
classied into two main styles focused attention (FA) and open monitoring (OM) involving different
attentional, cognitive monitoring and awareness processes. In a functional magnetic resonance study we
originally characterized and contrasted FA and OM meditation forms within the same experiment, by an
integrated FAOM design. Theravada Buddhist monks, expert in both FA and OM meditation forms, and
lay novices with 10 days of meditation practice, participated in the experiment. Our evidence suggests
that expert meditators control cognitive engagement in conscious processing of sensory-related, thought
and emotion contents, by massive self-regulation of fronto-parietal and insular areas in the left hemisphere, in a meditation state-dependent fashion. We also found that anterior cingulate and dorsolateral
prefrontal cortices play antagonist roles in the executive control of the attention setting in meditation
tasks. Our ndings resolve the controversy between the hypothesis that meditative states are associated
to transient hypofrontality or deactivation of executive brain areas, and evidence about the activation of
executive brain areas in meditation. Finally, our study suggests that a functional reorganization of brain
activity patterns for focused attention and cognitive monitoring takes place with mental practice, and
that meditation-related neuroplasticity is crucially associated to a functional reorganization of activity
patterns in prefrontal cortex and in the insula.
2010 Elsevier Inc. All rights reserved.

1. Introduction
Meditation can be conceptualized as a family of complex emotional and attentional regulatory practices, involving different
attentional, cognitive monitoring and awareness processes. Many
recent behavioral, electroencephalographic and neuroimaging
studies have revealed the importance of investigating meditation
states and traits to achieve an increased understanding of cognitive
and affective neuroplasticity, attention and self-awareness, as well
as for relevant clinical implications [7,28].
Given that regulation of attention is the central commonality
across the many different meditation methods [14], medita-

Corresponding author at: Department of Clinical Sciences and Bioimaging, ITAB,

Institute of Advanced Biomedical Technologies, G. DAnnunzio University, via Dei
Vestini, Campus Universitario, 66100 Chieti, Italy, Tel.: +39 0871 3556952;
fax: +39 0871 3556930.
E-mail address: amanna@unich.it (A. Manna).
1
These authors have contributed equally to this work.
0361-9230/$ see front matter 2010 Elsevier Inc. All rights reserved.
doi:10.1016/j.brainresbull.2010.03.001

tion practices can be usefully classied into two main styles

focused attention (FA) and open monitoring (OM) depending on how the attentional processes are directed [7,28]. In the
FA (concentrative) style, attention is focused on an intended
object in a sustained fashion. The second style, OM (mindfulnessbased) meditation, involves the non-reactive monitoring of the
content of experience from moment to moment, primarily as
a means to recognize the nature of emotional and cognitive
patterns.
FA meditation entails the capacities of monitoring the focus
of attention and detecting distraction, disengaging attention from
the source of distraction, and (re)directing and engaging attention to the intended object [28]. These attentional and monitoring
functions have been related to dissociable systems in the brain
involved in conict monitoring, selective and sustained attention
[12,28,32,40]. A study with a binocular rivalry paradigm showed
that Tibetan Buddhist monks were able to perceive a stable, superimposed percept of two dissimilar, competing images presented to
separate eyes for a longer duration both during and after FA meditation, but not during and after a form of compassion (emotional

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

OM) meditation [10]. These extreme increases in perceptual dominance durations suggest that extensive training in FA meditation
might improve the abilities to sustain attention focus on a particular
object and to control the ow of items being attended for conscious
access. A recent fMRI study investigated the neural correlates of
FA meditation in experts (following Tibetan Buddhist traditions)
and novices, with meditation focus on an external visual point
[4]. FA meditation compared with a rest condition, was associated
with activation in multiple brain regions involved in monitoring,
such as dorsolateral prefrontal cortex (DLPFC), attentional orienting (e.g., the superior frontal sulcus and intraparietal sulcus) and
engaging attention (visual cortex). The meditation-related activation patterns depended on the level of expertise of the meditation
practitioners.
OM meditation involves no explicit attentional focus, and therefore does not seem associated to brain areas implicated in sustained
or focused attention, but to brain regions involved in vigilance,
monitoring and disengagement of attention from sources of distraction from the ongoing stream of experience [28]. OM practices
are based on an attentive set that is characterized by an open
presence and a nonjudgemental awareness of sensory, cognitive
and affective elds of experience in the present moment, and
involves a higher-order awareness or observation of the ongoing
mental processes [7]. The cultivation of this reexive awareness
in OM meditation is associated to a more vivid conscious access
to the rich features of each experience and enhanced metacognitive and self-regulation skills [28]. Behavioral studies have shown
a more distributed attentional focus [39], enhanced conict monitoring [37] and reduced attentional blink or more efcient resource
allocation to serially presented targets [34] in OM meditation practitioners.
Despite the increasing number of studies on neural correlates of
meditation states and traits, the differential brain activity patterns
in focused attention and open monitoring meditation forms have
not been contrasted yet in a neuroimaging experiment. Therefore,
in an fMRI experiment we studied the FA and OM meditationrelated brain activity patterns of Buddhist monks who are expert in
Samatha (FA) and Vipassana (OM) meditation forms, and follow the
oldest (Theravada) currently active Buddhist tradition. Vipassana
(insight) meditation is central in mindfulness-based clinical interventions and studies [8,38]. Although lay practitioners of Vipassana
have participated in recent research (e.g., [17,34]), to our knowledge this is the rst study in which Theravada Buddhist monks are
involved.
Our integrated FAOM experimental design allows testing of
whether FA and OM meditation styles enhance or, by contrast,
reduce brain activations in frontal and other executive areas, given
controversial evidence and theoretical stances. Indeed, it has been
recently argued that meditative states are associated to transient
hypofrontality or deactivation in executive networks [15,26]. In
contrast, other authors have emphasized the activation of executive areas in meditation [7,28]. We hypothesize that the brain
regions associated with conict monitoring, such as the dorsal anterior cingulate cortex (ACC) and DLPFC [9,40], selective attention,
such as the temporalparietal junction, ventrolateral prefrontal
cortex, intraparietal sulcus and frontal eye elds [12] and sustained attention, such as right frontal and parietal areas, and the
thalamus [13,32], were more involved in inducing and maintaining the state of FA meditation as compared to the conditions of
OM meditation and non-meditative rest [28]. Given neuropsychological [22] and psychophysical [18] evidence of dominance of
the left cerebral hemisphere in conscious access, and theoretical
bases to hypothesize a leading role of this hemisphere in conscious
experiences [2,22,27], we predict a leftward bias of activation in
fronto-parietal areas in OM meditation as compared to the other
conditions.

47

2. Materials and methods

2.1. Participants
Participants included 8 Theravada Buddhist monks (males, mean age 37.9 years,
range 2553 years, SD 9.4 years), with 15,750 h on average of balanced Samatha (FA)
and Vipassana (OM) meditation practice in Theravada monasteries (SD 9900 h). The
monks were from the Santacittarama monastery, in central Italy, following a Thai
Forest Tradition (the order was funded by Ajahn Chah, one of the most inuential
Buddhist teachers in the 20th century). In this tradition, monks experience regular intensive meditation retreats, with a balanced practice of FA (Samatha) and
OM (Vipassana) meditation forms, including an about 3-month long winter retreat.
Outside the retreat period, the monks typically practice SamathaVipassana meditation, with a balance of FA and OM meditation, 2 h per day with the monastery
community. Individual meditation practice, with a balance of FA and OM meditation forms is also emphasized. Thus, on average the expertise of the monks in the
studied group can be estimated in 15,750 h of balanced FAOM meditation practice. Participants also included a group of 8 novice meditators (males, mean age
32 years, ages 2636 years, SD 3 years), recruited from the local community. All
novice subjects were interested in meditation but had no prior meditation experience. The novice participants were given oral and written instructions on how to
perform Samatha and Vipassana meditation styles, and during the 10 days before
the fMRI scan session practiced each of the two meditation styles 30 min per day.
The meditation instructions were written by Ajahn Chandapalo, the abbot of the
Santacittarama monastery, expert SamathaVipassana meditation teacher. All participants were right-handed. Subjects gave their written informed consent according
to the Declaration of Helsinki [41].
2.2. Task and protocol
The FAOM experimental paradigm consisted of 6 min FA (Samatha) and 6 min
OM (Vipassana) meditation blocks, each preceded and followed by a 3 min nonmeditative resting state block (Rest), for three times (see Fig. 1). The total duration
of the experiment was 57 min. The condition switch was instructed by an auditory
word-signal during the experiment.
To perform FA meditation, participants were given the following instruction:
gently engage in sustaining the focus of your attention on breath sensations, such
as at the nostrils, noticing with acceptance and tolerance any arising distraction,
as toward stimuli or thoughts, and return gently to focus attention on the breath
sensations after having noticed the distraction source. In OM meditation, participants were given the following instruction: observe and recognize any experiential
or mental content as it arises from moment to moment, without restrictions and
judgement, including breath and body sensations, percepts of external stimuli, arising thoughts and feelings. The instruction for Rest was the following: rest in a
relaxed awake state. In FA and OM meditation as well as in the Rest task condition,
the subjects did not employ any discursive strategy, recitation, breath manipulation or visualization technique. During all the conditions, the participants kept eyes
closed. At the end of the experiment, all participants reported they could perform
the FA, OM and Rest task conditions according to the given instructions, with no
differences in the experienced difculty to perform FA and OM meditation conditions.
2.3. Functional MRI recording
Functional MRI scans were acquired on a Siemens Magnetom Vision scanner
at 1.5 T, equipped with a standard receiver head coil. BOLD contrast functional
imaging was performed using a T2 -weighted echo planar (EPI) free induction decay
(FID) sequence with: TR = 4 s, 28 slices, voxel size 4 mm 4 mm 4 mm, 860 functional volumes for each run. A high-resolution T1 -weighted whole-brain image
was also acquired at the end of each session via a 3D-MPRAGE sequence (sagit-

Fig. 1. Sequence of the experimental conditions during the measurements.

48

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

tal matrix = 256 256, FOV = 256 mm, slice thickness =1 mm, no gap, in-plane voxel
size = 1 mm 1 mm, ip angle = 12 , TR/TE = 9.7/4.0 ms).
2.4. Physiological measures
Respiration rate and ECG were recorded throughout each scanning session in all
subjects. EEG was also recorded, with data to be analyzed for a subsequent report.
2.5. Data analysis
Raw data were analyzed using Brain Voyager QX 1.7 software (Brain Innovation,
The Netherlands). The rst ve scans of each run were discarded to avoid the T1 saturation effect. Preprocessing consisted in a 3D motion correction and in a temporal
ltering of voxel time series. The data set of one of the monks was discarded from
further analysis due to excessive motion. Preprocessed functional volumes were
co-registered with the corresponding structural data set. Co-registration transformation was determined using the slice position parameters of the functional images
and the position parameters of the structural volume. Temporal ltering included
linear and non-linear (high-pass lter of two cycles per time course) trends removal.
Structural and functional volumes were than transformed into the Talairach space
[36]. No spatial or temporal smoothing was applied.
Statistical analysis was carried out for individual subjects and condition using
the General Linear Model [21]. To account for the hemodynamic delay, the boxcar waveform of each task condition was convolved with the Boynton empirically
founded hemodynamic response function [3]. In order to search for activated areas
common to the entire group of subjects, a voxel-wise random effect group analysis
was performed, distinguishing between monks and novice meditators (withingroup analysis). To this purpose, all the subjects time series were z-normalized
and individual betas computed by specifying subject-specic regressors in the GLM.
Moreover, to quantitatively evaluate differences between the brain activity patterns of monks and novices, and to include group as a factor in our statistical model
of the fMRI data, between-group inferences were also computed (between-group
analysis).
Group statistical maps were thresholded at an overall signicance level (the
probability of a false detection for the entire functional volume) of p < 0.01, corrected
for multiple comparisons. The correction for multiple comparisons was performed
using a cluster-size thresholding algorithm [19] based on Monte Carlo simulations
and implemented in the BrainVoyager QX software. A threshold of p < 0.01 at the
voxel level, a FWHM = 1 voxel as Gaussian kernel of the spatial correlation among
voxels and 5000 iterations were used as input in the simulations, yielding a minimum cluster size of 8 voxels ( < 0.01).
2.6. Activations
Clusters of activation were obtained from the group activation maps considering voxels showing a signicant response (p < 0.01, corrected) to any experimental
condition. The Talairach coordinates of Table 1 are referred to the most signicant
voxel in each activated cluster. A correlation analysis between these activations was
performed. Specically, the correlation between the time course of the mean BOLD
response (in a certain experimental condition) recorded from voxels belonging to
two given clusters was evaluated. In order to assess the signicance level for the
r-Pearson coefcient, the Bonferroni correction for multiple comparisons among
clusters was applied to the p-threshold.
Moreover, an additional correlation between BOLD estimated parameters (betaweights resulting from both contrasts, FA > Rest and OM > Rest) and both expertise
level and age was evaluated, with reference to the ROIs emerging from the withingroup analysis for the monks.
2.7. Respiration rate correlation
Since all subjects underwent physiological monitoring during scanning, a
correlation analysis between the average respiration rates recorded in different
meditation/rest conditions was performed; in addition, t-test across conditions was
evaluated for both monks and novices, to probe condition-related breathing behavior differences between expert and novice meditators.

3. Results
3.1. Within-group analysis
In order to test the involvement of differential brain activations
in FA and OM meditation styles, with reference to Rest, the FA
meditation vs. Rest condition (FA > Rest) and OM meditation vs.
Rest condition (OM > Rest) contrasts were considered. Complementarily, we analyzed brain activations in the OM meditation vs. FA
meditation (OM > FA) contrast. The results revealed by these contrasts for monks and novices are summarized in Table 1.

3.1.1. Contrasts in the monk group

The contrast FA > Rest (Fig. 2) showed a wide pattern of deactivations in the left hemisphere, comprising multiple clusters in the
middle frontal gyrus (MFG), dorsolateral prefrontal cortex (DLPFC)
(BAs9/46), lateral anterior prefrontal cortex (aPFC) (BA10), precuneus (BA7), transverse temporal gyrus (TTG) (BA41), anterior and
posterior insula (BA13). The deactivations of the inferior frontal
gyrus (IFG) (BA44 and BA46) and the superior temporal gyrus (STG)
(BA22) were found in the right hemisphere. Moreover, three medial
frontal areas exhibited an increased activity as compared to Rest,
located in the left and right dorsal anterior cingulate cortex (ACC)
(BA24), and in the right medial aPFC (BA10).
The contrast OM > Rest (Fig. 3) revealed three activations in
the left hemisphere: medial aPFC (BA10), superior temporal gyrus
(STG) (BA22) and superior parietal lobule (SPL)/precuneus (BA7;
the activation cluster was centered in the SPL but extended medially in the precuneus). The contrast OM > FA (Fig. 4) showed
a large pattern of activations in the left hemisphere, including
DLPFC (BAs9/46, superior frontal gyrusSFG and middle frontal
gyrusMFG), lateral aPFC (BA10, MFG), medial frontal gyrus
(MeFG) (BA9), precuneus (BA7), superior parietal lobule (SPL)
(BA7), and anterior insula (BA13). In the right hemisphere, the SFG
in lateral aPFC (BA10), the IFG (BA46) and the TTG (BA41) were
activated. It is due to make the reader notice that a large part of the
activations are actually due to the fact that those areas are deactivated during FA, rather than activated during OM. A deactivation
of the dorsal ACC (BA24) and the medial aPFC (BA10) was found in
the right hemisphere.
3.1.2. Contrasts in the novice group
As regards the novices, the contrast FA > Rest (Fig. 5) showed a
single activation in the left posterior cingulate (BA31). The contrast
OM > Rest (Fig. 5) showed activations in the left dorsal ACC (BA32),
the right rostral ACC (BA32), the right lateral orbitofrontal cortex
(IFG, BA47) and the right medial aPFC (BA10). The contrast OM > FA
yielded no signicant activation.
3.1.3. Cluster correlations
In order to explore the differential state-dependent signal correlations between areas activated in the FA > Rest and OM > Rest
contrasts, as well as between these areas and other hypothesisrelevant regions, we conducted a correlation analysis with clusters
of activation derived from the contrasts, in the monks and control subject groups. We evaluated the correlation between time
courses of the mean BOLD response recorded from voxels belonging to two given clusters, in FA, OM or Rest conditions. In the monks,
the correlations between clusters which were (positively) activated
in the FA > Rest contrast were considered, with the correlations
between such clusters and a subset of hypothesis-relevant clusters deactivated in the same contrast, including left lateral DLPFC
and right IFG. In the same way, the correlations between the clusters which were (positively) activated in the OM > Rest contrast
were considered, with the correlations between such clusters and a
subset of hypothesis-relevant clusters activated in the OM > FA contrast, including anterior insula, lateral aPFC and SPL. Correlations
were also computed between clusters activated in the FA > Rest and
OM > Rest contrasts in the novices.
In particular, we hypothesized positive correlations within the
set of the three clusters activated in FA > Rest, and within the set
of the three clusters activated in OM > Rest, to reect two interarea brain circuitries for FA and OM meditation: a medial frontal
(prevalently right) circuitry for attentional focusing and monitoring in FA meditation; a left fronto-parieto-temporal circuitry for
open awareness in OM meditation, We also hypothesized negative correlations between the three activated clusters and a set of
deactivated executive clusters in the FA > Rest contrast, including

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

49

Table 1
Contrast results for monks and novices.
Group/contrast/area

10
9
48
47
12
12
54
48
0
41
57
42
42
41

OM meditation > FA meditation

Left MFG, BA46
Left MeFG, BA9
Left SFG, BA9
Left MFG, BA10
Left MFG, BA46
Right SFG, BA10
Right dorsal ACC, BA24
Right MeFG, BA10
Right IFG, BA46
Left STG, BA22
Left TTG, BA41
Right TTG, BA41
Left precuneus, BA7
Left SPL, BA7
Left anterior insula, BA13
OM meditation > Rest
Left MeFG, BA10
Left SPL/precuneus, BA7
Left STG, BA22

Monks
FA meditation > Rest
Left SFG, BA10
Left dorsal ACC, BA24
Left MFG, BA46
Left MFG, BA9
Right MeFG, BA10
Right dorsal ACC, BA24
Right IFG, BA44
Right IFG, BA46
Left precuneus, BA7
Left TTG, BA41
Right STG, BA22
Left anterior insula, BA13
Left anterior insula, BA13
Left posterior insula, BA13

Novices
FA meditation > Rest
Left posterior cingulate, BA31
OM meditation > Rest
Left dorsal ACC, BA32
Right rostral ACC, BA32
Right MeFG, BA10
Right IFG, BA47

66
26
39
32
50
32
17
41
70
25
51
8
19
10

19
16
13
28
13
14
13
14
49
10
13
3
1
7

81
270
618
243
351
432
135
270
270
314
54
270
162
1458

5.579
6.832
5.706
6.338
8.821
6.639
5.174
8.225
3.806
8.552
6.439
5.627
3.581
8.714

***
***
***
***
****
***
**
****
*
****
***
***
*
****

41
3
27
26
48
24
12
15
51
48
39
54
6
30
32

44
38
48
59
32
41
30
48
29
31
31
20
73
55
23

25
29
31
13
13
22
14
10
14
5
13
13
46
61
5

108
162
54
57
81
54
270
351
108
297
513
189
297
108
513

7.319
8.993
4.333
5.201
4.311
5.386
9.525
4.726
4.683
4.157
5.183
8.367
6.957
3.69
7.76

***
****
**
**
**
**
****
**
**
**
**
****
***
*
***

3
18
54

53
64
37

10
43
7

216
108
297

7.867
4.503
5.63

23

25

37

216

8.889

****

12
12
15
21

20
39
56
23

22
4
14
5

117
378
243
2125

4.809
7.892
6.812
8.035

**
****
****
****

***
**
***

Contrast results for monks and novices. Single-voxel uncorrected p-value are denoted by *(p < 0.01), **(p < 0.005), ***(p < 0.001), ****(p < 0.0001).

lateral frontal areas in the left and right hemispheres. Finally, in this
correlation analysis we considered activated clusters in the insula,
given the observed massive deactivation of it in the FA > Rest condition, and its implication in meditation from previous studies (see
[7,28]).
Average positive and negative signicant correlations in each
of the groups, were computed. We considered clusters as correlated if the r-value was larger than 0.4 in at least four subjects
in the group, for positive correlations, and lower than -0.4 in at
least four subjects, for negative correlations. The positively or negatively correlated clusters, according to such criteria, are shown in
Table 2.
3.1.4. Correlations in the monk group
As regards clusters resulting from FA > Rest contrast, we found a
high positive correlation between the activated medial frontal areas
during FA, OM and Rest conditions (with the left dorsal ACC and the
right medial aPFC correlating only in the Rest condition). A negative
correlation, during the Rest condition, was found between these
activated right and left dorsal ACC clusters, and a deactivated left
DLPFC cluster, as well as between the left dorsal ACC and a right IFG
activation in the same contrast. A negative correlation between the

(activated) right medial aPFC cluster and a (deactivate) left DLPFC

cluster, was also found in Rest.
With reference to clusters activated in the OM > Rest and
OM > FA contrasts, we found that left medial aPFC positively correlated with lateral aPFC, STG and anterior insula in the left
hemisphere. The left STG also correlated with the left superior parietal lobule and the left anterior insula. These correlations were
found in all experimental conditions. Finally, the left SPL/precuneus
correlated with the left STG, in the Rest condition.
3.1.5. Correlation with expertise level
The probed correlation between beta-weights recorded in
both contrasts (FA > Rest and OM > Rest) and meditation expertise
revealed no signicant outcomes for the OM condition, whereas
during FA meditation both deactivations of right IFG (BA46) and
of the posterior insula (BA13) positively correlate with meditation
expertise (Fig. 7). Finally, no signicant correlation with age was
observed.
3.1.6. Correlations in the novice group
Correlations between the four activations in the FA > Rest
or OM > Rest contrasts, in the novices group, were computed.

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A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

Fig. 2. Activations and deactivations revealed by the FA > Rest contrast, in the monks group. Note the deactivation of insula (BA13), MFG (BA46), TTG (BA41) and precuneus
(BA7) in the left hemisphere, and of IFG (BAs44/46) in the right hemisphere. Note also the activation of left/right dorsal ACC (BA24) and right medial aPFC (BA10). In this and
in all the gures the right hemisphere is on the left side (radiological convention).

Fig. 3. Activations revealed by the OM > Rest contrast, in the monks group, including medial aPFC (BA10), SPL/precuneus (BA7) and STG (BA22), in the left hemisphere.
Table 2
Correlations between activations resulting from the FA > Rest, OM > Rest and OM > FA contrasts, for both monks and novices. The percentage of subjects for which signicance
was displayed, is indicated.
Group

Contrast

Condition

Areas

Average r-value

Subjects percentage

Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Monks
Controls
Controls

FA > Rest
FA > Rest
FA > Rest
FA > Rest
FA > Rest
FA > Rest
FA > Rest
OM > Rest
OM > Rest
OM > Rest
OM > Rest
OM > Rest
OM > Rest
OM > Rest
OM > Rest

Rest
All
All
Rest
Rest
Rest
Rest
All
All
All
Rest
All
All
All
All

Dorsal ACC, L-BA24/MeFG, R-BA10

Dorsal ACC, L-BA24/Dorsal ACC, R-BA24
MeFG, R-BA10/Dorsal ACC, R-BA24
Dorsal ACC, L-BA24/MFG, L-BA9
Dorsal ACC, L-BA24/IFG, R-BA44
MeFG, R-BA10/MFG, L-BA9
Dorsal ACC, R-BA24/MFG, L-BA9
MeFG, L-BA10/MFG, L-BA10
MeFG, L-BA10/STG, L-BA22
MeFG, L-BA10/Anterior insula, L-BA13
Precuneus, L-BA7/STG, L-BA22
STG, L-BA22/SPL, L-BA7
STG, L-BA22/Anterior insula, L-BA13
Dorsal ACC, L-BA32/MeFG, R-BA10
Rostral ACC, R-BA32/MFG, R-BA11

0.46
0.59
0.58
0.44
0.44
0.43
0.44
0.47
0.40
0.43
0.44
0.43
0.53
0.41
0.53

88%
100%
100%
62.5%
62.5%
62.5%
88%
62.5%
62.5%
62.5%
62.5%
62.5%
75%
63%
75%

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

51

Fig. 4. Activations and deactivations revealed by the OM > FA contrast, in the monks group. Note the activation of anterior insula (BA13), STG (BA22), TTG (BA41), MFG (BA46),
MeFG (BA9) and SPL (BA7) in the left hemisphere. Note the activation of IFG (BA46) in the right hemisphere as well. The deactivation of right dorsal ACC (BA24) and medial
aPFC (BA10), are also shown.

We found a positive correlation between left dorsal ACC and

right medial aPFC, and between right rostral ACC and the right
orbitofrontal cluster, in all conditions.
3.2. Between-group analysis
Quantitative differences emerged from the between-group
analysis (see Fig. 6). In order to evaluate if these differences were
signicant and determine their direction, t-tests (between the
BOLD of monks and controls) were computed on the resulting clusters. This analysis conrms that the monks, as compared to the
novices, increased dorsal ACC (BA24) and right MeFG (BA6 and
BA10) activity bilaterally to focus their attention (FA meditation).

In OM meditation, the monks engaged more than novices the left

precuneus/SPL (BA7), the right dorsal ACC (BA32) and the right
parahippocampal gyrus. By contrast, novice participants showed
a higher engagement of the right rostral ACC (BA32), bilateral IFG
(BA47), right orbitofrontal (BA11) and right medial aPFC in open
monitoring.
3.2.1. Respiration rate correlation
The resulting correlation matrix (Fig. 8) showed no signicant
differences between OM, FA and Rest conditions for the monks,
i.e. individual differences in basal rates of respiration (e.g., during
rest) are preserved in the other conditions (FA, OM). In contrast,
the novices differentiated their respiration rates, above all between

Fig. 5. Activations and deactivations revealed by the FA > Rest and OM > Rest contrasts, in the novice group. On the left, the deactivation of the left posterior cingulate (BA31)
in FA > Rest is shown. The other sections show the activation of the orbitofrontal IFG (BA47), medial aPFC (BA10) and rostral ACC (BA32), in the right hemisphere, and dorsal
ACC (BA32) in the left hemisphere. The orientation of the sections follows the radiological convention.

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A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

Fig. 6. Quantitative differences between monks and novices emerging from the between-group analysis during FA (left) and OM (right) meditations. The letter k indicates
the number of voxels, and p the signicance of the peak voxel.

OM and rest condition (Fig. 8(a) and (b), t-test across conditions).
This result attests a peculiar condition-related breathing behavior
of each group, and indicates that novice participants might reduce
their focus on respiration when they are asked to emphasize the
monitoring faculty.
4. Discussion
For the rst time brain activity patterns in FA and OM meditation were contrasted in a neuroimaging (fMRI) experiment,
in expert (Buddhist monks) and lay novices, with an integrated
FAOM paradigm. Overall, we found striking differences between
the patterns of brain activity of monks and novices, in OM and
FA meditation styles. The brain activity patterns of the monks
in OM meditation resembled their ordinary brain resting state,
whereas their brain activity in focused attention meditation
sharply contrasted with both these states. In the monks, the
larger differentiation between brain activity patterns in FA and
OM meditation conditions, as compared to the OM > Rest contrast, indeed suggests that open monitoring (mindfulness) is also
reected and thus practiced in ordinary non-meditative conditions.
It has been recently argued that meditative states are associated
to transient hypofrontality or deactivation in executive networks
[15,26]. In contrast, other authors have emphasized the activation of executive areas in meditation [7,28]. As expected, the
results with our experimental design resolve this controversy: we
conclude that FA meditation is associated to an enhanced (predom-

inantly right) medial frontal and a reduced (predominantly left)

lateral prefrontal activation, and OM meditation to an increased
(predominantly left) medial frontal activation, as compared to
rest. We also conclude that OM meditation, as compared with FA
meditation, is characterized by a lateral prefrontal activation in
both hemispheres, with a more subtle differentiation in medial
frontal brain activations associated to these fundamental meditation styles.
Unlike brain activations, signal correlations between areas
activated in the contrasts were mostly not sensitive to the
meditation/rest conditions, against our expectations. The resulting correlation patterns suggest that networks or large-scale
multiregional assemblies of neurons, plausibly emerging by neuroplasticity, are recruited throughout different rest and meditation
conditions. However, the activation level of their components
located in different brain areas would depend on the meditation or
rest conditions, as shown by our contrasts. As discussed below, the
resulting positive and negative correlation patterns contribute to
shed light on functional connections between subsets of activated
brain areas, and suggest that a functional reorganization of the brain
resting state takes place in mental practice experts, with spatially
distributed neural activations modulated by different meditation
states.
We will rst consider meditation state effects in the monks and
then in the novices, and subsequently discuss these two groups
comparatively. The implications of our ndings for the cognitive
neuroscience of attention and awareness, will be thoroughly considered through the discussion.

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

Fig. 7. Signicant correlation results between clusters emerging from the withingroup analysis in the monk group. The correlation between beta-weights recorded
in each condition (FA and OM) and both meditation expertise and age reveals that
during the FA meditation the right IFG (BA46) and the left posterior insula (BA13)
positively correlate with meditation expertise. No signicant correlation with age
was observed.

4.1. Self-regulation of cognitive engagement in the monks

At a macroscopic level, a relative left-lateralization of brain
activity patterns resulted in our experiment. Most of the deactiva-

53

tions in FA > Rest and activations in OM > FA, and all the activations
in OM > Rest, were in the left hemisphere. These patterns include
a massive deactivation of left anterior and posterior insula in
FA > Rest, and a consistent activation of the left anterior insula
in OM > FA. We also found that the precuneus, which has been
associated to self-referential activations [11,26], was involved in
the left hemisphere in all contrasts. Specically, it was deactivated in FA > Rest, and activated in OM > Rest and OM > FA, as
hypothesized. The left precuneus was the only neural site to show
a similar pattern in our contrasts. This evidence suggests that
the left precuneus might plausibly act as a key brain region in
the self-induced transitions between brain resting states associated to meditative and non-meditative attentional sets, in the
monks.
The OM > Rest and OM > FA contrasts, with related correlation
analyses, also point out the relevance of the left SPL in BA7, which
might act in an inter-area circuitry with the left STG activated in
the OM > Rest contrast.
In asymmetry with the left hemisphere, in the right hemisphere
we did not nd activation or deactivation of (medial and lateral)
BA9, insula and posterior parietal areas, in any of the contrasts.
This evidence could not be predicted in light of the ndings of
a set of thicker cortical areas (including anterior insula) in the
right hemisphere of insight meditators [25]. This set of right hemisphere areas found in previous MRI structural studies might be
involved in an ongoing phenomenal awareness of the elds of
experience, independent on meditation-related attention focus and
open monitoring. Indeed, during FA and OM meditations (as well
as in rest), Buddhist monks experience an ongoing phenomenal
awareness of sensory elds, even though items in these elds may
not be intentionally accessed and investigated [28]. Conscious
access to selected contents of experience might instead take place
in fronto-parietal areas of the left hemisphere, consistent with psychophysical evidence about dominance of the left hemisphere in
perceptual awareness [18,29,31].
Finally, the activations found in the OM > Rest contrast in the
monks concern three main regions typically associated to selfreferential processing [32], with a peculiar left-lateralization. This

Fig. 8. Correlations between respiration frequencies recorded in different meditation/rest conditions: correlation matrix (left) and t-test evaluated for monks (a) and novices
(b). No signicant differences emerged in monks, i.e. individual differences in basal rates of respiration (e.g., during rest) are preserved in the other conditions (FA, OM); in
contrast, novices differentiated their respiration rates, above all between OM and Rest condition (a and b: t-test across conditions). This further result reveals condition-related
breathing behavior differences between expert and novice meditators.

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A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

left-lateralized network of cortical midline and superior temporal

areas might reect a neuroplasticity-based cognitive reorganization of the monk brains, with neuronal populations in brain regions
ordinarily linked to self-referential processing reallocated to a
metacognitive observation of phenomenal experience and of the
experiencing subject [33,35]. This refers to a crucial aspect of
Buddhist practice and meditation, i.e. the cultivation of wholesome mental states going beyond the cognition of a separated self
[1].
As we did not nd activation (or deactivation) of the left IFG,
hypothesized to play an important role in self-awareness as an
inner speech-related region [30], it can be inferred that the leftlateralized activity patterns we have observed are likely not to be
associated with a narrative self function theorized to account for
the dominance of the left hemisphere in conscious access [2,22].
Moreover, given that it has been recently shown that the left IFG
is characterized by a higher regional grey matter concentration in
Vipassana meditators [23], the activation of this area could have
been revealed in OM > Rest and OM > FA. Our negative evidence,
however, indicate that the monks did not perform a language-based
access to present moment experiences.
Our evidence suggests that the monks might control cognitive
engagement and broadcasting in brain networks for conscious
access to sensory-related, thought and emotion contents, by massive self-regulation of fronto-parietal and insular areas in the left
hemisphere, in a meditation state-dependent fashion. Specically,
this self-regulation appears mostly controlled by alternated activations of (right) dorsal ACC and a set of neural sites in (left)
MFG(right) IFG, and reected in massive changes in activity levels
of (left) insula and parietal posterior cortex, as well as in a shift of
medial aPFC activation.
4.2. Control of focused attention in the monks
The observed dorsal ACC (left and right BA24) activations
showed by the FA > Rest contrast are consistent with their predicted involvement in conict monitoring during FA meditation.
The right medial aPFC (BA10) area activated in the same contrast
is plausibly involved in focused awareness during FA meditation. The observed high positive correlations between these three
frontal medial areas suggest that these midfrontal areas interact
in a unitary circuitry, with a higher activation during FA meditation. The hypothesized activations in FA meditation associated
to selective attention (temporalparietal junction, ventrolateral
prefrontal cortex, intraparietal sulcus and frontal eye elds) and
sustaining attention (right frontal and parietal areas, and the thalamus) were not observed. As an unpredicted result, in FA > Rest we
observed a remarkable deactivation of DLPFC, more pronounced in
the left hemisphere. The deactivation of the left DLPFC was even
more widespread in OM > FA, with the additional involvement of
the dorsolateral SFG. This evidence suggests that sustaining the
attentional focus in FA meditation implies a deactivation of DLPFC
areas.
Interestingly, a negative correlation (in Rest) between both the
right and left dorsal ACC clusters activated in FA > Rest, and a left
DLPFC cluster deactivated in FA > Rest, was found. This negative correlation suggests that, especially when distraction is more likely
(Rest), dorsal ACC and left DLPFC play a contrasting role in maintaining cognitive focus and opening the eld of cognitive monitoring.
We also found evidence about a role of the right IFG (BA44 and
BA46) and the left DLPFC, as shown in contrasts and correlations
(see Section 3).
It can therefore be concluded that dorsal ACC and lateral (in
particular dorsolateral) prefrontal cortex play antagonist roles in
executive control of the attention setting, as observed in meditation
tasks without goal-related actions.

4.3. Meditation state effects in the novices and comparison with

the monks
As regards the novices, we only found the deactivation of the left
posterior cingulate cortex in FA > Rest. Considering this result and
the evidence about the precuneus in the monks, consistent with
the recent proposal that the precuneus/posterior cingulate cortex
plays a pivotal role in the default mode network [20], it can be
hypothesized that the left precuneus/posterior cingulate region is
the component of the default mode network which can be more
sensitively affected by a goal-independent task, such as FA meditation. We also conclude that unlike FA meditation on a visual
point [4], breath-centered FA meditation may demand a longer
(more intensive) practice to observe more differentiated fMRI activations, at least with a paradigm characterized by short-duration
meditation blocks as in our experiment. It can also be concluded
that the monks performed a demanding FA meditation task in our
experiment.
Four activated clusters in the OM > Rest contrast were found in
the novice group. Overall, it seems that in novices open monitoring
mostly involved right prefrontal areas. The activation of the left
dorsal ACC might be explained by the executive demand to novices
in OM meditation performance. The positive correlation between
left dorsal ACC and right medial aPFC in the novice group, suggest
that these two areas co-operate in enabling cognitive focus.
The activations in novices of (right) rostral ACC and (right) lateral orbitofrontal cortex (IFG), which were not found in the monk
group, suggest that in novices open monitoring may reect an
evaluation-based stance. Indeed, there is evidence that rostral ACC
and lateral orbitofrontal cortex are involved in affective and cognitive evaluation processes [6,16,24]. These two areas were found to
be positively correlated in the novice group.
The key involvement of dorsal anterior cingulate cortex (BA24)
and right medial frontal gyrus for focused attention in monks, and of
the right rostral anterior cingulate cortex (BA32) and orbitofrontal
cortex/inferior frontal gyrus (BA11/BA47) for open monitoring in
novices, is conrmed by the between-group analysis.
4.4. Correlates of monk expertise
The probed correlation between beta-weights recorded in both
conditions and meditation expertise of the monks revealed that
during FA meditation both the deactivations of the right IFG (BA46)
and of the left (posterior) insula (BA13) positively correlate with
meditation expertise, i.e. larger deactivations are observed in more
expert practitioners. This evidence suggests that the expertisedependent sustained focused attention implies the deactivation
of such regions, likely to be involved in a more transient awareness of experience contents [28]. The disengagement of these
areas might also be plausibly related to a more effortless maintenance of attentional (cognitive) focus in expert meditators [28],
in terms of a reduced background of neural activations which can
potentially reorient the allocation of limited attention-related brain
resources.
4.5. Potential caveats and further investigations
It remains to be seen how the differential brain activity patterns
we have found correlate with hours of practice in a larger group
of FA/OM meditation practitioners. Although all the participants in
our study reported that they could perform FA and OM meditation
forms according to the given instructions and with no differences
in experienced difculty, in a further study with a larger group
of participants, a quantitative measure of effort (e.g., self-ratings
on a Likert-scale), performed block by block, can provide a more
accurate information about effort for FA and OM meditation forms.

A. Manna et al. / Brain Research Bulletin 82 (2010) 4656

Indeed, effort is an important aspect of meditation performance

[4,28]. Consideration of other aspects, such as balancing for subjective efcacy (how well FA and OM were performed), intensity (how
intense the experience was) and stability (for how long the meditative state persisted) within and between groups, also appears
important. As suggested by previous research [4], controlling for
subjective motivation while meditating and structural differences
in brain anatomy due to ethnicity, are further relevant aspects to
keep into account.
In further studies it may be insightful to compare brain activity
patterns in OM meditation conditions with differential awareness
of elds of experience, such as body sensory eld, external sensory
elds and internal thoughts and feelings. There is indeed evidence
with a related theoretical account [5], to differentiate prefrontal
regions involved in stimulus-oriented and stimulus-independent
processing. Moreover, it appears useful to design an experiment
comparing brain activity patterns in FA meditation with focus on
breath-related sensations (as in our study) and on an external visual
point (as in [4]). FA meditation might involve a different level of
engagement with external world processing and modulation of
self-referential (default mode) brain networks, depending on the
focused object.
To conclude, the present study appears to shed light on fundamental aspects of meditation practice and the related set of
cognitive functions, beyond and complementarily to the ndings
in previous studies, as related to the unique participants in it and
its design. The present ndings might open the way to deeper
investigations of meditation-based awareness, with important
implications for the domain of neural correlates of consciousness
[33].
Conict of interest
The authors declare that they have no competing nancial interests.
Acknowledgements
First of all, we thank the monks of the Santacittarama monastery
for their kind participation in the study, as well as for useful feedback at different stages of the work. We thank Antoine Lutz for
important suggestions to improve the design of the experiment.
We also thank Alessandro DAusilio and Valerio Santangelo for useful comments, and Luca Simione for kind assistance. Finally, we
have beneted from helpful and critical discussions about statistical data analysis from two of our close colleagues, Gianna Sepede
and Nicoletta Cera and we wish to sincerely thank their continuous
support in this study. Finally, we would like to thank two anonymous reviewers for important remarks and comments which have
conducted to a signicantly improved manuscript.
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