NEUROSCIENCE

REVIEW
Leila Chaieb et al. / Neuroscience 409 (2019) 69–80

New perspectives for the modulation of mind-wandering using transcranial

electric brain stimulation
Leila Chaieb, a* Andrea Antal, b Marlene Derner, a Marcin Leszczyński cd and Juergen Fell a
a
Department of Epileptology, University of Bonn, Sigmund-Freud Str. 25, 53105 Bonn, Germany
b
Department of Clinical Neurophysiology, University Medical Center, Robert-Koch-Str. 40, 37073 Göttingen, Germany
c
Department of Neurological Surgery, Columbia University Medical Center, New York, USA
d
Translational Cognitive Neuroscience Program, Nathan Kline Institute, Orangeburg, New York, USA

Abstract—When our attention is decoupled from an ongoing task and becomes coupled to thoughts and feelings not
being subject to task engagement, we are mind-wandering. This transient and pervasive mental process can occupy a
considerable amount of our waking hours. Mind-wandering is understood to exert both positive and negative effects
on well-being, and has been shown to play a role in mood disorders and depression. Here we summarize recent
research aiming to investigate whether states of mind-wandering can be modulated using transcranial direct current
stimulation (tDCS), a non-invasive, reversible means of altering neuronal excitability and in turn, cortical activity. We
examine and compare the methodologies underlying the existing studies on this topic, and evaluate the commonal-
ities and contrasts of their outcomes. So far, existing studies tentatively suggest an influence of tDCS on the contents
and propensity to mind-wander. However, these studies exhibit considerable methodological differences and changes
in the propensity to mind-wander are inconsistent with task performance. Moreover, replication of the results of two
studies from the same group by another group has recently failed. We discuss the implications of these findings, in
particular, regarding therapeutic targets in mood disorders, and propose perspectives for future investigations. For
instance, tDCS effects on deliberate versus undeliberate mind-wandering should be disentangled. The hippocampus
as an important hub for mind-wandering-related processes may be targeted. Most importantly, research efforts related
to mind-wandering and rumination should be integrated. © 2019 IBRO. Published by Elsevier Ltd. All rights reserved.

Key words: transcranial electrical stimulation, mind-wandering, task-unrelated thought, meta-awareness, dorsolateral prefrontal
cortex, hippocampus.

INTRODUCTION when a task is monotonous or not cognitively engaging,

Mind-wandering is a pervasive and ubiquitous mental phe-
nomenon. It is often understood as a process during which
our attention shifts from an ongoing task (often referred to
as the ‘task at hand’) to thoughts and feelings not being sub-
ject to task performance (for a review see (Smallwood and
Schooler, 2015). In general, when task demands are low, or
*Corresponding author. Tel.: + 49 228 287 19344; fax: + 49 228 287
16294.
E-mail address: leila.chaieb@ukbonn.de (Leila Chaieb).
Abbreviations: tES, transcranial electric stimulation; MW, mind-wander-
ing; TUT(s), task-unrelated thought(s); MA, meta-awareness; DLPFC,
dorsolateral prefrontal cortex; tDCS, transcranial direct current stimula-
tion; tACS, transcranial alternating current stimulation; tRNS, transcra-
nial random noise stimulation; DMN, default mode network; NIBS, non-
invasive brain stimulation; SART, sustained attention to response task;
LPFC, left prefrontal cortex; rIPL, right inferior parietal lobule; PCC,
posterior cingulate cortex; mPFC, medial prefrontal cortex; MDD, major
depressive disorder.
mind-wandering episodes become more prevalent
(Smallwood et al., 2004). In addition, individuals who show
deficits in working memory have a higher propensity to
mind-wander during demanding tasks and display behaviors
associated with goal-neglect (McVay and Kane, 2009). It
has been reported that this shifting of attention to focus on
spontaneously-generated thoughts is commonly associated
with more negative mood states (Killingsworth and Gilbert,
2010), in particular, when mind-wandering is related to the
past (Ruby et al., 2013a). This in turn, can lead to rumination
and feelings of persistent worry (Killingsworth and Gilbert,
2010; Stawarczyk et al., 2013). Studies have shown that per-
sistent rumination typically increases depressive symptoms
and contributes to dysphoric mood states, leading to a reduc-
tion in wellbeing (Hertel, 1998; Smallwood and O'Connor,
2011). However, mind-wandering also has positive functions
such as promoting creative thinking (Baird et al., 2012;
Leszczynski et al., 2017), enabling better social problem

https://doi.org/10.1016/j.neuroscience.2019.04.032
0306-4522/© 2019 IBRO. Published by Elsevier Ltd. All rights reserved.

69
70 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
solving (Ruby et al., 2013b), fostering future planning to over-
come potential obstacles (Oettingen and Schwörer, 2013)
and providing mental breaks by allowing us to dissociate from
monotonous activities (Mooneyham and Schooler, 2013;
Ruby et al., 2013b).
Nevertheless, despite evidence of the beneficial effects of
mind-wandering, many studies indicate that an increased
propensity to mind-wander generally impacts negatively on
mood states and even on cognitive processes. This effect
appears to be larger when the thoughts are repetitive and
pervasive (Ottaviani et al., 2014). A recent study reported
that patients with major depressive disorder (MDD) are
more likely to show a higher propensity to mind-wander, in
addition to exhibiting a specific pattern of self-related, rumi-
native and anxious thoughts (Hoffmann et al., 2016). Such
thoughts were persistently reported during the performance
of a continuous response task, when compared to healthy,
age-matched controls. These findings suggest that both
an increased propensity to mind-wander, combined with
highly specific thought patterns, may serve as markers of
depressive states. Such markers may act as targets for clin-
ical interventions and may allow for the development of new
therapeutic strategies.
WHAT EXACTLY IS MIND-WANDERING?
The term “mind-wandering” is used to describe conceptually
different thought processes, however, with overlapping char-
acteristics. According to the family-resemblances view, as
discussed by Seli et al. (2018), it is sufficient to provide sev-
eral related definitions or descriptions of mind-wandering
instead of precisely demarcating the term “mind-wandering”.
Such descriptions encompass, for instance, task-unrelated
thought (TUTs), unintentional thought, stimulus-independent
thought and even daydreaming (Schooler et al., 2011;
Shrimpton et al., 2017; Seli et al., 2018). However, what com-
plicates the case is that mind-wandering can be also entered
into with intent (Seli et al., 2016). For instance, subjects may
intentionally mind-wander during experimental settings used
to assess mind-wandering, since the implemented tasks are
typically boring by design and subjects try to cope with their
boredom. In view of these overlapping, but also distinct mean-
ings the need to understand which processes are targeted
when investigating the neural correlates of mind-wandering
is of utmost importance. Hence, it is currently being discussed
how to approach designing experimental paradigms targeting
particular components of mind-wandering and their effects
(Christoff et al., 2018; Seli et al., 2018). Up to now, most
experimental studies addressing mind-wandering have
investigated this phenomenon in terms of task-unrelated
thought. To examine this aspect, researchers typically
presented experience sampling probes inquiring about
mind-wandering during perceptual/attentional tasks requiring
continuous monitoring. This approach has also been applied
in the tDCS studies described in this review. As a conse-
quence, the findings of these studies should be primarily
interpreted with regard to the modulation of task-unrelated
thought.
WHICH BRAIN REGIONS ARE INVOLVED IN
MIND-WANDERING?
Studies examining the effects of mind-wandering have often
focused on cortical regions comprising the default mode net-
work (DMN; for a review see (Andrews-Hanna et al., 2014).
Since activity within DMN regions is registered when fixation
is contrasted with task performance or when self-related
tasks are contrasted with non-self-related tasks, and since
mind-wandering is often self-related and more prominent in
the absence of a task (Chou et al., 2017), this suggests a
contribution of DMN regions to mind-wandering. Indeed,
the DMN is – through activation of different subsystems –
understood to play a vital role in the generation of self-
generated thoughts (Smallwood and Schooler, 2015; Axelrod
et al., 2017), as well as in mental time travel, which is an
important component of retrospective and future-oriented
mind-wandering (Karapanagiotidis et al., 2017). In line with
these ideas, a decreased frequency of mind-wandering among
patients with ventromedial prefrontal cortex lesions, a DMN
region, compared to both healthy and brain-damaged controls
has been reported (Bertossi and Ciaramelli, 2016). However, a
meta-analysis of 24 neuroimaging studies investigating spon-
taneous thoughts and mind-wandering (Fox et al., 2015) iden-
tified neural correlates of these activities not only within DMN
areas, such as medial prefrontal cortex, posterior cingulate
cortex, medial temporal lobe, including the hippocampus,
and inferior parietal lobule, but also within several non-DMN
regions, such as dorso-, rostro- and ventrolateral prefrontal
cortex, dorsal anterior cingulate cortex, insula, temporopolar
cortex, secondary somatosensory cortex, and lingual gyrus.
The authors concluded that regions beyond the DMN and in
particular, regions comprising the frontoparietal executive con-
trol network are in addition involved in mind-wandering.
Furthermore, fMRI studies using connectivity analyses have
shown that fluctuations in functional connectivity between a
ventral DMN subsystem and posterior cingulate cortex reflect
spontaneous mind-wandering (Kucyi and Davis, 2014), and
that mind-wandering is characterized by lapses in executive
control which can be predicted by co-activation of the DMN
and subregions of the task-positive network (Mittner et al.,
2014).
As research into mind-wandering continues to develop,
there is emerging evidence from a few studies to suggest
a role of the hippocampus in the initiation of spontaneous
thoughts (Andrews-Hanna et al., 2010; Ellamil et al.,
2016). A recent fMRI study examining which cortical regions
support the early generation of spontaneous thoughts in
highly trained meditators, revealed a sequential recruitment
of brain regions prior to, during and subsequent to the onset
of spontaneous thoughts (Ellamil et al., 2016). The primary
region that showed activation prior to the arising of sponta-
neous thoughts was the hippocampus. Subsequently, other
regions comprising the DMN and executive control regions
were activated (for instance, (prior to) the posterior cingulate
cortex, right inferior parietal lobule, and right posterior insula
and (during) medial prefrontal cortex, left and right temporo-
polar cortex and left and right mid-insula). In a recent review
by the same group, the hippocampus has been suggested
 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
to act as a kind of relay, reactivating hippocampal–neocorti-
cal and, in turn, neocortical–neocortical ensembles, both
prior to and during the generation of spontaneous thoughts
(Christoff et al., 2016). New findings from a study examining
mind-wandering in six patients with selective bilateral hippo-
campal damage also emphasize the role of the hippocam-
pus in mind-wandering, but question its relevance for the
propensity to mind-wander (McCormick et al., 2018).
McCormick and colleagues compared the incidence and
contents of mind-wandering episodes of the patients versus
an age-matched, healthy control group. Patients and partici-
pants reported whether they engaged in mind-wandering at
probed time-points, in addition to recording the contents of
their mind-wandering episodes using an adapted version
of the descriptive experience sampling method (Hurlburt
and Heavey, 2001; Hurlburt and Akhter, 2006). The authors
observed that while patients engaged in mind-wandering
just as frequently as the control group (two-tailed Mann–
Whitney U test, p = 0.12), the contents of their mind-
wandering were almost exclusively related to the present
and comprised of verbally mediated semantic thoughts. In
contrast, the instances of mind-wandering reported by the
control group were dispersed across the three temporal
categories (past, present and future) and were described
using vivid imagery (McCormick et al., 2018).
In summary, there is evidence to support that both DMN
regions, most prominently the hippocampus, as well as non-
DMN regions, such as parts of lateral prefrontal cortex, are
involved in mind-wandering. Up to now, this evidence is
mainly based on correlational neuroimaging findings and very
few lesions studies. Non-invasive brain stimulation may offer
an alternative approach to investigate in greater depth, the
causal role of these candidate regions in mind-wandering.
USING NON-INVASIVE BRAIN STIMULATION
(NIBS) TO MODULATE COGNITIVE PROCESSES
Transcranial direct current stimulation (tDCS) is one mem-
ber of a group of non-invasive transcranial electrical stimu-
lation techniques, used to safely and reversibly alter levels
of cortical excitability. Other techniques include transcranial
alternating current stimulation (tACS) and random noise
stimulation (tRNS; in which stimulation frequencies are ran-
domly shuffled during application (Terney et al., 2008).
These techniques use currents that are applied through at
least two electrodes attached to the scalp, at low intensities
(most frequently using 1–2 mA) in order to target the under-
lying cortical area and related neuronal networks. Their
efficacy is dependent upon the duration of stimulation,
intensity, and in the case of tDCS, polarity of stimulation
(for an overview of NIBS guidelines see Bikson et al.,
2016; Woods et al., 2016; Antal et al., 2017).
Of the three most frequently used TES methods, tDCS is
the only stimulation technique that induces cortical excitabil-
ity increases or decreases depending upon the direction of
current flow (for a review see Filmer et al., 2014). TRNS
and tACS are in relation to tDCS relatively novel methods,
and are more suited to generating plasticity-like after-
effects and alterations in cortical excitability via entrainment
71
of cortical oscillations, due to the nature of their applied
sinusoidal currents (for a review see Antal and Herrmann,
2016). Early animal studies have demonstrated that tDCS
induces cortical excitability changes via the modulation of
neuronal resting membrane potentials (Bindman et al.,
1962). Generally it is stated that current flow in an anodal
direction causes depolarization, whereas cathodal stimula-
tion induces a hyperpolarization of the resting membrane.
However, there is evidence that at the cellular level, anodal
vs. cathodal tDCS affects different cellular components due
to the different current polarities (Bikson et al., 2012, 2013;
Rahman et al., 2013, 2014). In studies examining the appli-
cation of direct currents using cell-cultures, it was observed
that anodal tDCS hyperpolarizes the membrane potential in
the apical dendritic regions and depolarizes it in the somatic
region, whereas the cathodal electrode had the reverse
effect (Rahman et al., 2013, 2014). Besides cell type and
morphology, the extent to which neurons are affected by
tDCS also depends on the orientation of the cells with
regard to the induced electric field (Rahman et al., 2014).
While tDCS is in itself insufficient to elicit action potentials,
its application impacts the spontaneous firing rate of the
neurons underneath the stimulating electrode (Bindman et
al., 1962; Purpura and Mcmurtry, 1965) and associated
neuronal networks (Vöröslakos et al., 2018). Stimulation of
increased duration has been shown to induce
neuroplastic-like effects, for instance, in the motor system
and visual cortex (Nitsche and Paulus, 2001; Lang et al.,
2005; Antal et al., 2014).
An increasing number of studies have examined the role
of tDCS in cognitive processes such as working memory
(Zaehle et al., 2011); (for a review see (Albouy et al.,
2018), attention (Jacoby and Lavidor, 2018; Miler et al.,
2018), associative verbal memory (Flöel et al., 2008) and
episodic memory (Penolazzi et al., 2010). However, tDCS
in a low-definition montage lacks some spatial focality, and
so it is difficult to determine exactly which superficial cortical
structures are affected by the stimulation (Bortoletto et al.,
2016).
STUDIES INVESTIGATING THE IMPACT OF TDCS
ON MIND-WANDERING
Up to now, seven studies have investigated the modulation
of mind-wandering in healthy subjects by tDCS (see Table 1
and Fig. 1). In a first study, 1-mA anodal tDCS over the left
dorsolateral prefrontal cortex (DLPFC; vs. right supraorbital
ridge cathode) was applied for 20 min (Axelrod et al., 2015).
Control conditions comprised of anodal tDCS over the occi-
pital lobe and sham stimulation (using the DLPFC montage,
but administered only for 2 min). During stimulation partici-
pants performed a sustained attention to response task
(SART), which has been shown to facilitate mind-
wandering (Robertson et al., 1997). Digits were presented
successively on a screen and participants were asked to
respond to each digit displayed with a button press except
when a target digit was presented. Mind-wandering was
accessed via experience sampling probes, i.e. participants
were asked approximately every 1.5 min to report to what
 72
Table 1. Summary of methodological approaches and outcomes of studies examining the effects of tDCS on mind-wandering.

Study Site of Site of Stimulation Control Timing of task Task Additional methods Effects of Effects of tDCS Additional findings
(year) stimulation: stimulation: conditions condition(s) (relative to tDCS) tDCS on on task
anode cathode mind-wandering performance
Axelrod left DLPFC (F3) right supra- 1 mA, 20 min, 1) sham during stimulation SART – increased vs. none –
et al., orbital 7 × 5 cm 2 cathode, (2 min stimulation) (online) both control
2015 4 × 4 cm2 anode, 2) occipital (Oz, conditions
saline soaked sponges anode vs. Cz)
Kajimura left LPFC (AF7)/ right IPL/ 1.5 mA, 20 min, sham, 30 s post-stimulation target – increased/inverse load- –
and right IPL (P4) left LPFC 7 × 5 cm 2, saline stimulation detection/ decreased dependent flanker
Nomura, soaked sponges flanker task vs. sham effect for tDCS vs.
2015 sham
Kajimura left LPFC/ right right IPL/ 1.5 mA, 20 min, sham, 30-s post-stimulation target resting state increased/ not analyzed rIPL-PCC/
et al., IPL left LPFC 7 × 5 cm 2, saline stimulation detection/ fMRI before decreased mPFC-PCC
2016 soaked sponges flanker task and after tDCS; vs. sham connection
DCM inhibited/

Leila Chaieb et al. / Neuroscience 409 (2019) 69–80

facilitated
mind-wandering
Bertossi right deltoid mPFC 2 mA, 15 min, 1) sham, 30-s before and after CRT temporal direction opposite none opposite changes
et al., (left of Fpz) 5 × 5 cm 2 cathode, stimulation stimulation and self-relatedness changes of self-relatedness
2017 5 × 7 cm 2 anode, 2) occipital probes (after–before) of MW for tDCS
saline soaked sponges (Oz, cathode) vs. both vs. sham, only
control conditions in males
only in men
Kajimura right IPL/ left IPL contra- 1.5 mA, 20 min, sham, 30-s post- stimulation SART meta-awareness decreased for reaction time No effect of
et al., (P3) lateral cheek 7 × 5 cm 2, saline stimulation probe; resting state right IPL vs. decreased for left stimulation
2018 soaked sponges fMRI; DCM; sham IPL vs. sham condition on
machine learning meta-awareness;
within-DMN
connectivity
and amplitude
changes
for right IPL
vs. sham
Axelrod left DLPFC (F3) right 1 mA, 30 min, 1) sham during stimulation SART meta-awareness increased vs. none No effect of
et al., supra-orbital 7 × 5 cm 2 cathode, (2-min stimulation) (online) probe both control stimulation
2018 7 × 5cm2 anode, 2) occipital (Oz, conditions condition on
saline soaked sponges anode vs. Cz) meta-awareness;
decreased MW
during high levels
of meta-awareness
Boayue left DLPFC (F3) right 1 mA, 20 min, sham during stimulation SART Bayesian statistics absence of any none No online effect of
et al., supra-orbital 7 × 5 cm 2 cathode, (15-s fade-in (online) and effect of tDCS stimulation on
2019 4 × 4 cm 2 anode, fade-out) post-stimulation on MW task performance;
saline soaked sponges (offline) no offline effects on
MW or task
performance
Abbreviations: tDCS, transcranial direct current stimulation; DLPFC, dorsolateral prefrontal cortex; LPFC, lateral prefrontal cortex (ventrolateral/orbitofrontal); IPL, inferior parietal lobule; mPFC, medial prefrontal cortex; PCC, posterior
cingulate cortex; SART, sustained attention to response task; CRT, choice reaction task; fMRI, functional magnetic resonance imaging; DCM, dynamic causal modeling; DMN, default mode network.
 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
Fig. 1. Example of a tDC stimulation montage and a summary of elec-
trode positions used by the studies examining tDCS effects on mind-
wandering. Stimulation was applied to the left DLPFC (F3) vs. right
supraorbital region (Axelrod et al., 2015). Active stimulation was com-
pared to a sham condition where stimulation was applied over the occi-
pital cortex (Oz) and over the vertex (Cz). Color-coded electrodes
positions depict sites of stimulation: Red- Axelrod et al., 2015 and Axel-
rod et al., 2018; green- Kajimura and Nomura, 2015 and Kajimura et al.,
2016; blue- Bertossi et al., 2017; orange- Kajimura et al., 2018; violet-
Boayue et al., 2019.
extent they had just experienced task-unrelated thoughts
(on a scale of 1–4). Axelrod and colleagues reported an
increased propensity to mind-wander during anodal
73
DLPFC stimulation compared to both control conditions.
No influence of tDCS on task performance was observed.
In a commentary on the findings published by Axelrod and
colleagues, Fox and Christoff (2015) raised an important
question concerning the participants' self- reporting of their
mind-wandering episodes, since a prerequisite of reporting
mind-wandering is that subjects are so-called meta-aware
of this process. Stimulation of the DLPFC may have influ-
enced not only the propensity to mind-wander, but also the
meta-awareness of mind-wandering, i.e. subjects may have
been more often aware that their mind was not focused on
the task due to DLPFC stimulation. Indeed, there is evi-
dence to suggest that not only mind-wandering, but also
meta-awareness is mediated by the DLPFC (Christoff et
al., 2009; Fleming and Dolan, 2012). Thus, DLPFC
stimulation-related enhancement of meta-awareness may
putatively have contributed to the increase of self-reported
mind-wandering episodes. An experimental option to con-
trol for this influence is to implement a second experience
sampling probe related to meta-awareness (e.g. (Christoff
et al., 2009).
Axelrod and colleagues recently addressed this critique
by conducting a replication study employing a large cohort
of 91 subjects (in the earlier study the participant population
consisted of 47 subjects (Axelrod et al., 2015)). The design
was similar to their previous study, however, with the addi-
tion of a meta-awareness probe (“To what extent have you
been aware of where your attention was focused?; Possible
responses: 1 (minimal)–4 (maximal)”) and a longer stimula-
tion duration of 30 min instead of 20 min. Confirming their
previous findings, Axelrod and colleagues reported an
increased propensity to mind-wander during anodal
DLPFC stimulation versus both control conditions and no
effect of tDCS on task performance. Importantly, they found
that meta-awareness was not modulated by stimulation
condition. They further observed that high levels of meta-
awareness were associated with decreased levels of
mind-wandering. This finding is in line with results by
Christoff and colleagues (2009) showing that meta-
awareness during mind-wandering is related to decreased
activity in DMN and executive control regions, and suggests
a mitigating effect of meta-awareness on mind-wandering.
Using a different approach, Kajimura and Nomura (2015)
implemented a 2 × 2 design, where they applied anodal vs.
cathodal tDCS in a left prefrontal cortex (LPFC) vs. right
inferior parietal lobule (rIPL) montage, as well as in the
reverse montage (anode: rIPL vs. cathode: LPFC), at an
intensity of 1.5 mA for 20 min (Kajimura and Nomura,
2015). The authors chose to stimulate the rIPL (a DMN
region) using the return electrode, as this region is under-
stood to be involved in the regulation of task-unrelated
thoughts (Mason et al., 2007; Christoff et al., 2009). The
control condition consisted of sham stimulation, during
which current was passed through the stimulating electro-
des only for 30 s before being switched off. The LPFC sti-
mulation electrode (position AF7; extended 10/20 system)
chosen by Kajimura and Nomura (2015) was located slightly
more lateral/anterior than the DLPFC electrode (position F3)
selected by Axelrod et al. (2015). In the case of Kajimura
74 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
and Nomura (2015), the stimulation site was rather ventro-
lateral/ orbitofrontal. Importantly, the cognitive paradigm
was conducted after the stimulation period and not during
stimulation (i.e. online), as in the study by Axelrod et al.
(2015). The paradigm consisted of a visual target detection
task with letter stimuli and additional congruent or incongru-
ent flanker letters, which the subjects were instructed to
ignore. The task was performed in a low vs. high perceptual
load condition (one vs. five competing letters). Thought
probes examined the contents of the participants' thoughts
immediately prior to the appearance of the probe (partici-
pants were asked to choose from the following categories
to describe their experiences: task contents, task perfor-
mance, a memory from the past, something in the future,
current state of being, fantasy, other; only the latter five
choices were classed as mind-wandering; please note that
thinking about task contents and task performance may
not necessarily equate to “being on-task”). Kajimura and
Nomura (2015) reported that compared to sham stimulation,
the propensity to mind-wander was significantly increased/
decreased by LPFC (anode) vs. rIPL (cathode)/rIPL (anode)
vs. LPFC (cathode) stimulation respectively. Furthermore,
they described that the dependence of the flanker effect
(i.e. the accuracy of target detection for (congruent-incon-
gruent)/congruent flankers) on perceptual load was inverted
for both stimulation conditions (larger flanker effect for the
high load) compared to sham stimulation (larger flanker
effect for the low load).
In a subsequent fMRI study, Kajimura et al. (2016),
sought to examine the neural underpinnings of the tDCS-
induced increase/decline in the propensity to mind-
wander, which they reported in their earlier study
(Kajimura and Nomura, 2015). This study used the same
stimulation protocols and cognitive paradigm, but different
experience sampling probes (probe 1: were you thinking of
something?: yes/no; probe 2: where was your attention
focused just before the probe?: on-task/off-task; instances
of mind-wandering were categorized based on participants'
responses to probe 2). Prior to and after the stimulation per-
iod, the authors conducted two resting-state fMRI scans,
which were subjected to functional and effective connectiv-
ity analyses. Participants performed the cognitive paradigm
after the second fMRI scan. Replicating their earlier beha-
vioral findings, the authors again observed that LPFC vs.
IPL/IPL vs. LPFC stimulation increased/decreased the like-
lihood of mind-wandering compared to sham stimulation.
Based on dynamical causal modeling of the fMRI data, the
authors reported that anodal stimulation of the rIPL
decreased the afferent connections of the posterior cingu-
late cortex (PCC) from the rIPL and the medial prefrontal
cortex (mPFC). Furthermore, based on mediation analysis,
they found that the connection from the rIPL to the PCC
inhibited, while the connection from the mPFC to the PCC
facilitated, mind-wandering (Kajimura et al., 2016).
In a recent fMRI study, Kajimura et al. (2018) moreover
aimed to investigate the idea of a functional asymmetry
between the IPLs with regard to mind-wandering. For this
purpose, they alternatively applied anodal tDCS to the rIPL
and lIPL (cathode: contralateral cheek). In their article, the
anode positions were specified as right/left angular gyrus
(i.e. subregions of r/lIPL), however, the right stimulating
electrode was placed at the same position (P4), as reported
in their previous studies (left electrode at P3) (Kajimura et
al., 2018). The study design was similar to that in Kajimura
et al. (2016) with two resting state fMRI scans, one prior to
and one post stimulation, however, using a different cogni-
tive paradigm, this time a standard SART task (Robertson
et al., 1997), being conducted after the second scan.
Experience sampling probes were in accordance with those
used by Christoff et al. (2009), with an initial probe addressing
instances of mind-wandering (“where was your attention
focused just before the probe?: on-task/off-task), and a sec-
ond probe addressing meta-awareness (“how aware were
you of where your attention was focused?: aware/unaware).
For analysis of the fMRI data the authors applied dynamic
causal modeling and a machine learning approach. With
regard to the behavioral results, the authors reported a
decrease in the propensity to mind-wander for rIPL stimula-
tion vs. sham, but not for lIPL stimulation vs. sham. However,
lIPL stimulation vs. sham stimulation decreased reaction
times in the SART task. No stimulation effects were
observed regarding meta-awareness. Based on the ana-
lyses of blood-oxygen-level dependent signals during rest-
ing state, the authors reported that only rIPL (and not lIPL)
stimulation vs. sham modulated within-DMN connectivity
and amplitudes. They concluded that there is indeed beha-
vioral and functional evidence for an IPL asymmetry with
regard to mind-wandering.
Finally, Bertossi et al. (2017) investigated the role of the
medial prefrontal cortex (mPFC) in mind-wandering. To this
aim, they applied cathodal tDCS (2 mA for 15 min) over the
mPFC (left of position Fpz), while the anodal return elec-
trode was attached to the right deltoid (i.e. to the right
shoulder). Control conditions consisted of stimulation with
a cathodal electrode over the occipital cortex (position Oz)
and sham stimulation (30-s stimulation). Before and after sti-
mulation a choice reaction time task (CRT) was performed.
The CRT task consisted of monitoring a stream of digits pre-
sented in two colors (one frequently, one rarely), where parti-
cipants were asked to report whether the digits of the
infrequently presented color were even or odd. Experience
sampling was performed based on three subsequent probes.
The first probe addressed the degree of mind-wandering (on/
task vs. off task: visual analog scale from 0 to 100), the sec-
ond probe addressed the content of mind-wandering in terms
of temporal direction (past/present/future/external distraction/
time not clear/unaware), and the third probe addressed the
content of mind-wandering in terms of self- vs. other-
relatedness (self-related/other-related/unrelated to people).
The authors evaluated the changes in incidences of mind-
wandering during post-tDCS vs. pre-tDCS task performance.
They observed that stimulation-related changes in the pro-
pensity to mind-wander occurred in different directions for
cathodal mPFC stimulation compared to both control condi-
tions. Mind-wandering decreased after cathodal mPFC stimu-
lation, but increased after occipital and sham stimulation.
Importantly, this effect was only observed in the male partici-
pants, but not in the females. The authors furthermore
 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
reported opposite changes in self-relatedness of mind-
wandering after both stimulation conditions compared to
sham (decreases vs. increase), again, only for the male sub-
jects, but not for the female participants (Bertossi et al., 2017).
While it remains unclear what these gender differences may
be attributed to, other studies using tDCS have also demon-
strated gender-specific effects of direct current stimulation,
for instance, with regard excitability changes within motor cor-
tex (Kuo et al., 2006) and visual cortex (Chaieb et al., 2008).
Very recently, Boayue et al. (2019) published the results
of a registered multicenter study, re-examining the findings
of Axelrod et al. (2015, 2018). The analysis and data collec-
tion plan had been preregistered at the European Journal of
Neuroscience. In total, Boayue et al. (2019) recruited 192
subjects in three study centers (Amsterdam, Göttingen
and Tromsø). They implemented the exact same stimula-
tion parameters as those used by Axelrod et al. (2015) in
their initial study. Mind-wandering during anodal left
DLPFC (F3) stimulation (cathode: right supraorbital ridge)
was compared to sham stimulation. Additionally, mind-
wandering for anodal stimulation vs. sham condition was
compared offline during a 20-min phase immediately follow-
ing stimulation. Boayue et al. (2019) found neither an online,
nor an offline effect of anodal DLPFC stimulation on mind-
wandering or on task performance. Using Bayesian statis-
tics they showed that based on their data, a null effect is
about 10 times more probable (Bayes factor = 10.65) than
an increase of mind-wandering due to anodal DLPFC stimu-
lation, as reported by Axelrod et al. (2015). They further
reported that there is extreme evidence that the findings of
the initial study (Axelrod et al., 2015) were not replicated
(Bayes factor = 1:500). Boayue et al. (2019) concluded that
they were unable to detect an effect of anodal stimulation
over the DLPFC on mind-wandering, but on the contrary
found evidence for the absence of such an effect (Boayue
et al., 2019).
DISCUSSION AND FUTURE PERSPECTIVES
The studies investigating the modulation of mind-wandering
via tDCS, described in this review, exhibit considerable
methodological differences, in particular regarding stimula-
tion montages, the choice of cognitive task used, timing of
the tasks and which experience sampling probes were
implemented (see Table 1 and Fig. 1). For instance, with
regard to task timing, it has been shown that tDCS effects
on motor sequence-learning qualitatively differ for stimula-
tion prior to, compared to during execution of the task
(Stagg et al., 2011). So far, at least the studies by
Kajimura and Nomura (2015); Kajimura et al. (2016, 2018)
and Bertossi et al. (2017) tentatively suggest that it is indeed
possible to modulate the propensity to mind-wander via
tDCS, and point to a causal role of the DMN and executive
control regions in mind-wandering. Nevertheless, we com-
pletely agree with the assessment of Boayue et al. (2019),
in view of their recent failure to replicate the findings of
Axelrod et al. (2015, 2018): “Given the negative results of
the current study, however, it is important to replicate any
75
of these positive effects before accepting them as facts.”
Keeping this limitation in mind, the observed tDCS-related
effects support the hypotheses that activity within left
dorso/ventrolateral PFC (i.e. executive control regions)
and within medial PFC (i.e. a DMN region) is positively cor-
related with mind-wandering (in the latter case only in men),
and that activity in the right IPL (i.e. a DMN region) is nega-
tively correlated with mind-wandering. To be more precise,
these findings indicate that mind-wandering in the sense
of task-unrelated thought, can be modulated using these
tDCS interventions. It remains an open question, whether
these findings are specifically related to undeliberate mind-
wandering, or whether they also apply to intentional mind-
wandering or indeed, other definitions of mind-wandering
(Seli et al., 2016). For instance, it has been suggested that
activity in executive control regions may be associated with
deliberate, rather than with undeliberate mind-wandering
(Seli et al., 2017). Hence, future studies may aim at disen-
tangling the influence of tDCS applied to dorso/ventrolateral
PFC on deliberate vs. undeliberate mind-wandering. Such
studies may also investigate whether any findings can be
transferred to other forms of mind-wandering, such as, for
instance, daydreaming or deliberate meandering. Further-
more, cognitive modeling of subprocesses contributing to
mind-wandering may help to elucidate differences and
commonalities between various forms of mind-wandering
(Mittner et al., 2014; Axelrod and Teodorescu, 2015). More-
over, it has to be noted, that in five of the seven studies no
meta-awareness probe has been implemented and it thus
remains unclear, whether and to which extent changes in
the meta-awareness of mind-wandering may have contribu-
ted to the observed changes in the propensity to mind-
wander (Fox and Christoff, 2015). However, the recent stu-
dies by Kajimura et al. (2018) and Axelrod et al. (2018) mon-
itored this variable, and could largely exclude a possible
bias.
A much more puzzling aspect of the reported findings is
that the effects on task performance did not mirror those
changes in the likelihood to mind-wander. An increase/
decrease in mind-wandering is expected to have a
detrimental/positive effect on task performance, for
instance, in terms of increased/reduced error rates or
increased/reduced reaction times. Task performance was
analyzed in six of the seven described studies. In four stu-
dies no effects of tDCS on task performance were found
(Axelrod et al., 2015; Bertossi et al., 2017; Axelrod et al.,
2018). In the other two studies, task effects were reported,
which are incoherent with the reported mind-wandering
effects. In the study by Kajimura and Nomura (2015) both
stimulation conditions (applying reverse montages) exhib-
ited inverse load-dependent flanker effects when compared
to sham. In the following study by Kajimura et al. (2018), a
reaction time decrease, compared to sham, was found for
the tDCS condition (left IPL anode), that showed no effect
on mind-wandering. Thus, the effects on task performance
do not fit into the overall picture of the reported modulation
of mind-wandering by tDCS. Speculatively, direct effects of
tDCS on task performance may have neutralized the indir-
ect effects due to mind-wandering. For instance, anodal
76 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
stimulation of executive control regions like dorso-/ventro-
lateral PFC may not only increase mind-wandering, but
may also facilitate task performance. However, evidence
for such counterbalancing effects is still lacking. Possibly,
more refined measures examining task performance, such
as the variation of reaction times across trials (e.g. Bastian
and Sackur, 2013; Leszczynski et al., 2017), may reveal
tDCS-induced effects, which can be related to the observed
modulations of mind-wandering. Moreover, machine learn-
ing techniques using functional measures, such as EEG,
fMRI or pupillometry, have been shown to be capable of
estimating mind-wandering on short time scales (Mittner et
al., 2014; Jin et al., 2019).
So far only tDCS has been used to modulate mind-
wandering. An alternative approach would be to apply
tACS. This technique uses sinusoidal waveforms, impacts
upon the amplitude and frequency of ongoing brain oscilla-
tions, and provides the unique opportunity to interfere with
cortical oscillations within a specific frequency range (for a
review see Antal and Herrmann, 2016). Although its effects
have been first characterized in the motor cortex (Antal et
al., 2008; Chaieb et al., 2011), recent studies have demon-
strated that tACS can modulate a range of perceptual and
cognitive processes (Herrmann et al., 2013; Fröhlich et al.,
2015) like visual, somatosensory and auditory perception
(Riecke et al., 2015; Alekseichuk et al., 2016; Gundlach et
al., 2016), working memory (Polanía et al., 2012; Vosskuhl
et al., 2015; Alekseichuk et al., 2017), speech (Rufener et
al., 2016; Zoefel et al., 2018), attention (Hopfinger et al.,
2017; Wöstmann et al., 2018) and cognitive control
(Santarnecchi et al., 2013; Hsu et al., 2017). Unlike tDCS,
tACS is understood to modulate brain activity via entrain-
ment of endogenous oscillations (Fröhlich and McCormick,
2010) and/ or by influencing the phase coherence of
ongoing oscillations (Neuling et al., 2012). For instance,
Braboszcz and Delorme (2011) recorded EEG from healthy
participants while they were asked to perform a breath-
counting task. The authors sought to investigate oscillatory
activity during episodes of mind-wandering, where partici-
pants' attention had drifted away from the breath-counting
task (as indicated by button presses). They reported an
increase in occipital and parieto-central theta EEG activity
during episodes of mind-wandering compared to the
breath-focus state (Braboszcz and Delorme, 2011). Specu-
latively, this occipital/parieto-central activity could be related
to processes within posterior cingulate cortex and/or precu-
neus, DMN regions likely to be involved in mind-wandering
(Fox et al., 2015). Accordingly, tACS could, for instance,
be applied to test the link between occipital/parieto-central
theta activity and mind-wandering. In this sense, tACS
may be more suitable to specifically target oscillatory pro-
cesses underlying mind-wandering.
The studies described in this review applied tDCS to
different neocortical regions in order to modulate mind-
wandering. However, deeper structures such as the hippo-
campus have also been reported to be involved in the
generation of spontaneous thoughts. Is it possible to target
deep brain structures such as the hippocampus, using
non-invasive brain stimulation? Indeed several potentially
suitable techniques have recently been proposed. Pulsed
transcranial ultrasound stimulation has been shown to be
able to modulate hippocampal activity in mice (Tufail et al.,
2010), and has been successfully applied to humans for
the modulation of activity in primary somatosensory
(Legon et al., 2014) and visual cortex (Lee et al., 2016).
Transcranial electric stimulation with temporally interfering
electric fields (Grossman et al., 2017) uses the superposi-
tion of high-frequency electric fields comprised of slightly
different frequencies. The interfering electric field patterns
exhibit a low-frequency amplitude modulation and produce
a spatially focused form of deep brain stimulation. This tech-
nique has been demonstrated to selectively stimulate the
hippocampus in mice (Grossman et al., 2017), and will likely
be transferred to humans, in the near future. A similar audi-
tory technique is so-called beat stimulation, consisting
either of the application of the physical superposition of
two nearby frequencies (resulting in an amplitude modu-
lated signal) to one or both ears simultaneously (monaural
beats), or the application of each individual frequency sepa-
rately to each ear (binaural beats) (for a review see Chaieb
et al., 2015). Research in epilepsy patients undergoing inva-
sive presurgical evaluation with intracranial electrodes has
shown that hippocampal activity can be modulated by audi-
tory beat stimulation (Becher et al., 2015). Of course in
these cases, deep brain stimulation could also be used to
unravel the brain regions contributing to mind-wandering
and to substantiate or disprove non-invasive stimulation
findings (e.g. Foster and Parvizi, 2017).
Another important issue is the actual localization and
direction of the stimulation currents induced by tDCS. For
instance, Boayue et al. (2019) have argued that the effects
observed by Axelrod et al. (2015, 2018) could be in part
due to indirect stimulation of the medial prefrontal cortex in
spite of targeting the DLPFC. Recent research has focused
on the use of realistic head models to address the issue of
focality of stimulation. One such study demonstrated that
tDCS-induced electric fields tend to spread beyond the
stimulating electrodes and are largely dependent upon indi-
vidual anatomical differences, in addition to being subject to
the positioning, size and type of electrodes used (Opitz et
al., 2015). Also, combined fMRI-tDCS studies showed that
surrounding cortical structures are inadvertently exposed
to stimulation (Keeser et al., 2011). Further research inves-
tigating the effects of tES in epilepsy patients with implanted
depth electrodes has demonstrated that electric fields tend
to be higher around the edges of stimulating electrodes,
resulting in inhomogeneous field distributions in the stimu-
lated tissue (Opitz et al., 2018). To improve upon these
issues electrodes of different shapes and sizes or multiple
electrodes may be applied in combination with computa-
tional modeling of current densities based on realistic head
models (Seo et al., 2017; Wang et al., 2018). On the other
hand, the precision of electrode placement can be
increased, for instance, by training technical personnel in
electrode placement using 3D head models in conjunction
with landmarks based on the 10/20 system (Indahlastari et
al., 2019), or by using MRI-guided neuronavigation systems
(De Witte et al., 2018).
 Leila Chaieb et al. / Neuroscience 409 (2019) 69–80
Apart from shedding light upon the causal role of different
brain regions, investigating the modulation of mind-
wandering using non-invasive brain stimulation may have
eminent clinical relevance: methods lessening incidences
of mind-wandering may also be effective in reducing
depressive rumination, i.e. a subtype of mind-wandering
being characteristic for patients suffering from major
depressive disorder or obsessive compulsive disorder.
These repetitive thoughts with similar content, often con-
cerning negative issues originating from past experiences
(Nolen-Hoeksema, 2000), have been linked to the onset
(Robinson and Alloy, 2003) and maintenance of depression
(Nolen-Hoeksema et al., 1997), as well as to its severity
(e.g. (Morrison and O'Connor, 2008). Patients suffering
from major depression were described to have more
mind-wandering episodes with less positive, more past-
oriented and self-related thoughts than healthy controls
(Hoffmann et al., 2016). Thus, non-invasive brain stimula-
tion methods aimed at reducing the propensity to mind-
wander, or modulating its content characteristics may open
new avenues for clinical interventions in major depression.
Nevertheless, it is still an open question as to how these
results from studies applying tDCS to modulate mind-
wandering, can be merged with findings from investigations
examining tDCS-effects on rumination. For example, Axelrod
et al. (2015) observed that anodal tDCS applied to the left
DLPFC increased the propensity to mind-wander, a finding
which could not be replicated by Boayue et al. (2019). Actually,
the rationale behind this choice of stimulation site is not quite
clear, since rather the right (but not the left) DLPFC has been
consistently associated with mind-wandering (see the meta-
analysis by Fox et al., 2015). However, it has also been
reported that anodal vs. cathodal tDCS applied bilaterally to
right vs. left DLPFC, increases angry rumination in healthy sub-
jects (Kelley et al., 2013), contrary to what would be expected
based on the mind-wandering findings by Axelrod et al.
(2015). Nonetheless, no modulatory effects of anodal vs. cath-
odal tDCS applied to left vs. right DLPFC on rumination were
observed in depression patients (Vanderhasselt et al., 2015).
There is converging evidence that the DLPFC, aside from
its role in executive control and mind-wandering, also plays
an integral role in the top-down control of emotions (for review
see (Okon-Singer et al., 2015; Dixon et al., 2017), and that
there is an activity imbalance between left DLPFC (hypoactiv-
ity) and right DLPFC (hyperactivity) in major depression
(Grimm et al., 2008). Accordingly, anodal tDCS of the left
DLPFC has been applied in several studies for the modulation
of depressive symptoms, and this approach has gained
recognition due to a probable antidepressant effect in patients
suffering from MDD in the absence of pharmacoresistance
(Lefaucheur et al., 2017). However, recent clinical trials show
that implementation of anodal tDCS to the left DLPFC as a
therapeutic option in the acute phase of MDD has not consis-
tently demonstrated robust efficacy (Borrione et al., 2018). In
view of these partially contradictory findings related to the
effects of tDCS on mind-wandering vs. rumination, conver-
ging these lines of research may provide new insights into
the mechanisms underlying the generation of non-
ruminative vs. ruminative thoughts. In addition, such an
77
integrative approach may provide more robust data with
regard to tDCS parameters suitable for clinical interventions.
DECLARATION OF INTEREST
LC, MD and JF were supported by the German Research
Foundation via the SFB1089.
ROLE OF THE FUNDING SOURCE
The funding sources provided financial support only and
had no involvement in the writing of this publication. The
authors declare no conflicts of interest.
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