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The left prefrontal cortex supports inhibitory processing during semantic

memory retrieval

Article in Cortex · January 2021

DOI: 10.1016/j.cortex.2020.11.001

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Research Report

The left prefrontal cortex supports inhibitory

processing during semantic memory retrieval

Martin Marko a,b and Igor Rie
canský a,c,*

a
Department of Behavioural Neuroscience, Institute of Normal and Pathological Physiology, Centre of Experimental
Medicine, Slovak Academy of Sciences, Bratislava, Slovakia
b
Department of Applied Informatics, Faculty of Mathematics, Physics and Informatics, Comenius University in
Bratislava, Bratislava, Slovakia
c
Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology,
Faculty of Psychology, University of Vienna, Vienna, Austria

article info abstract

Article history: Semantic control refers to a set of neural and cognitive mechanisms that govern semantic
Received 24 April 2020 processing and retrieval. Neuroimaging studies have indicated that controlled semantic
Reviewed 2 September 2020 processing engages the left prefrontal cortex (PFC), yet the functional role of the prefrontal
Revised 5 October 2020 activity in semantic control is poorly understood and was therefore addressed in the
Accepted 2 November 2020 present study. We used a double-blind randomized controlled experiment, in which par-
Action editor Angela Sirigu ticipants from three distinct groups received anodal transcranial direct current stimulation
Published online 20 November 2020 (tDCS) over left lateral PFC (n ¼ 40), a control tDCS over temporoparietal cortex (n ¼ 40), or
sham stimulation (n ¼ 41), while executing automatic and controlled semantic retrieval
Keywords: tasks as well as additional control tasks assessing working memory and semantic judge-
Semantic cognition ment. We demonstrate that anodal tDCS of the left lateral PFC improved inhibition of
tDCS prepotent semantic associations but had no significant effect on retrieval of habitual as-
Prefrontal cortex sociates or switching between retrieval rules. The prefrontal tDCS also enhanced working
Executive functions memory capacity, but this effect did not account for the improved semantic inhibition. The
Working memory control temporoparietal tDCS did not affect semantic retrieval. Our findings show that
semantic inhibition and switching represent distinct components of the semantic control
system and indicate that the left lateral PFC is involved in a filtering process that constrains
the accessible semantic representations (i.e., a proactive pre-retrieval inhibition) or sup-
presses already retrieved responses (i.e., a retroactive post-retrieval inhibition). The
recognition of such an inhibitory process could inspire novel treatments targeting altered
semantic processing.
© 2020 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

* Corresponding author. Department of Behavioural Neuroscience, Institute of Normal and Pathological Physiology, Centre of Experi-
mental Medicine, Slovak Academy of Sciences, Sienkiewiczova 1, 813 71 Bratislava, Slovakia.

anský).
E-mail addresses: martin.marko@savba.sk (M. Marko), igor.riecansky@savba.sk (I. Riec
https://doi.org/10.1016/j.cortex.2020.11.001
0010-9452/© 2020 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6 297

semantic processing gradually changes during the progres-

1. Introduction sion of verbal fluency tasks since retrieving information from
semantic memory becomes increasingly demanding as
Controlled semantic cognition supports multiple forms of
typical (i.e., easily-accessible) retrieval candidates are being
adaptive behavior and thought. Growing evidence suggests
depleted (Golestanirad et al., 2015; Marko et al., 2019a). Thus, it
that semantic control engages the left lateral prefrontal cortex
is currently not clear whether the previously observed im-
(PFC), which interacts with the temporal-parietal brain re-
provements in verbal fluency due to left prefrontal tDCS
gions involved in semantic representation, i.e., a system
reflect enhanced automatic retrieval (i.e., a bottom-up, stim-
forming generalizable, multi-modal concepts and their
ulus-driven retrieval at early stages of the task), controlled
mutual relations (Ralph et al., 2017; Chiou & Lambon Ralph,
semantic retrieval (i.e., a top-down, goal-directed retrieval
2019; Chiou et al., 2018). Multiple lines of research converge
employed when the automatic retrieval becomes insufficient),
to indicate that PFC is not involved in the encoding of se-
or domain-general executive abilities not uniquely related to
mantic representations per se, but it executively manipulates
semantic processing, such as working memory and response
the activation of semantic knowledge (Kapur et al., 1994;
monitoring.
Fedorenko & Thompson-Schill, 2014; Stampacchia et al., 2019;
In the present study, we addressed this problem using
Jefferies, 2013). For instance, neuroimaging studies have
anodal tDCS guided by computational models of electric field
shown that the left lateral PFC is engaged in semantic tasks
distribution to target the left lateral PFC. Semantic processing
requiring inhibition of habitual associates (Allen et al., 2008;
was assessed using the associative chain test (ACT; Marko
Burgess and Shallice, 1996; Cattaneo et al., 2011), retrieval of
et al., 2019a; Marko et al., 2019b) that implements a word-
less accessible knowledge from semantic memory (e.g., weak
production approach by which automatic (free association)
or infrequent associates; Badre & Wagner, 2002; Badre et al.,
and controlled (inhibition and switching) semantic processing
2005; Badre & Wagner, 2007), or a flexible switching between
can be effectively disentangled. Besides the prefrontal and
semantic features and/or categories (Troyer et al., 1998; Marko
sham stimulation, a montage targeting temporal-parietal
& Riec
anský, 2018). Nevertheless, despite a large number of
cortex bilaterally (i.e., not affecting the PFC) was included as
studies coupling general executive control with the activity of
a control montage to assess the anatomical specificity of the
PFC (Kiesel et al., 2010; Yuan & Raz, 2014; Diamond, 2013), little
stimulation effects. Our main prediction was that, compared
research has been dedicated to the executive processes in the
to sham and the temporal-parietal tDCS, the anodal prefrontal
semantic domain (such as inhibition or switching), and causal
tDCS would enhance controlled but not automatic semantic
evidence that PFC is involved in the semantic control func-
processing.
tions is limited.
Furthermore, two additional control measures were
Non-invasive brain stimulation provides a safe and effec-
introduced in the interest of establishing behavioral speci-
tive approach to investigate functional relations between
ficity of the expected stimulation effects. Working memory
brain and behavior (Polanı́a et al., 2018). In particular, trans-
capacity (WMC) was assessed as a domain-general executive
cranial direct current stimulation (tDCS) has been successfully
function that had been shown to be sensitive to prefrontal
used to modulate the excitability and functional connectivity
tDCS (Ruf et al., 2017; Ohn et al., 2008; Zaehle et al., 2011;
of cortical regions (Nitsche & Paulus, 2000; Rahman et al., 2013;
Brunoni & Vanderhasselt, 2014) and involved in semantic
Stagg & Nitsche, 2011; Filmer et al., 2019), resulting in altered
memory retrieval (Unsworth et al., 2011, 2013). Moreover, a
cognitive processing and behavior in healthy participants as
semantic judgement task (SJT) was used to discern whether
well as neuropsychiatric patients (Filmer et al., 2020). Accu-
left prefrontal tDCS also affects post-retrieval evaluation
mulating evidence indicates that prefrontal tDCS can modu-
and selection processes (Badre & Wagner, 2002; Badre et al.,
late semantic cognition (Joyal & Fecteau, 2016; Monti et al.,
2005; Badre & Wagner, 2007; Whitney et al., 2011), which are
2013). For instance, it has been reported that excitatory
considered domain-general and engage right rather than left
anodal tDCS over left lateral PFC improved picture naming
PFC (Hayama & Rugg, 2009; Sharp, 2004). Thus, both WMC
(Fertonani et al., 2010) and enhanced verbal fluency (Cattaneo
and SJT were assessed to refine the precise functional role of
et al., 2011; Pisoni et al., 2018). Interestingly, Cattaneo et al.
the left lateral PFC in controlled semantic processing and to
(2011) reported that prefrontal tDCS modulated semantic
challenge the main hypothesis with the alternative that
fluency more strongly than phonemic fluency. This indicates
tDCS affects semantic retrieval indirectly, via an improve-
that the prefrontal stimulation might exert stronger effect on
ment in domain-general cognitive abilities (working mem-
semantic processing compared with domain-general execu-
ory or post-retrieval response evaluation). Finally, all
tive processing, which is engaged more intensely in phonemic
cognitive tasks were administered before, during, and
than semantic tasks (Henry & Crawford, 2004; Shao et al.,
shortly after the tDCS in order to investigate both the im-
2014). For verbal fluency, on the other hand, the role of se-
mediate effects (“online effects”) and after-effects (“offline
mantic versus executive processing remains a matter of
effects”), as they may result from partially distinct neurobi-
debate (Shao et al., 2014; Whiteside et al., 2016) and several
ological processes (Stagg & Nitsche, 2011; Medeiros et al.,
theoretical models have posited distinct accounts for the
2012). The differences between online and offline tDCS ef-
domain-general executive involvement (e.g., suppression,
fects were assessed in an exploratory manner to inform
switching, monitoring, and/or maintenance; Mayr, 2002;
optimal application of tDCS for the modulation of semantic
Unsworth et al., 2013). Moreover, growing evidence suggests
processing in future studies.
that the relative engagement of automatic versus controlled
298 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6

2. Methods
2.1. Participants and study design
The study had a randomized, double-blind, placebo-
controlled design. The required sample size was estimated
using an a priori power analysis (5% Type I error rate, 20%
Type II error rate, and expected effect size hp2 > .10). In total,
121 participants (49 males; age 23.1 ± 3.8 years) completed a
single experimental session involving either PFC (n ¼ 40),
TPC (n ¼ 40) or sham (n ¼ 41) stimulation; the participants
were randomly assigned to balanced experimental groups.
Two more individuals participated (in the PFC and the TPC
groups) but did not finish the session due to technical is-
sues. All participants were right-handed (Edinburgh Hand-
edness Inventory e short form, Veale, 2014; mean score
92.3 ± 11.6), with no history of psychiatric disorder, neuro-
logical condition, or recent medication. Research was con-
ducted in accordance with the Declaration of Helsinki
(World Medical Association, 2013) and approved by the
institutional review board. All procedures and methods
were carried out in accordance with the relevant guidelines
and regulations. All participants gave written informed
consent and received a financial reward for their
participation.
2.2. Procedure
The outline of the main experimental procedure is depicted in
Fig. 1A. Each session started with a brief anamnestic inter-
view, which was followed by assessment of verbal fluency,
schizotypal personality traits using Schizotypal Personality
Questionnaire (SPQ), The Community Assessment of Psychic
Experience (CAPE), and current emotional state using affect
scales. Thereafter, a baseline measurement block (assessment
of WMC, ACT and SJT; see section 2.4) was carried out. After
the baseline block, the stimulation was set up and initiated. A
scale assessing adverse stimulation effects and a second block
of affect scales were administered immediately after the
current had ramped-up. The online measurement block
started after 5 min of full-intensity stimulation. At the tDCS
offset, participants provided perceived intensity rating for the
stimulation and completed a third measurement of the
emotional state using affect scales. The offline measurement
block was initiated 5 min after the stimulation offset. A short
debriefing was carried out at the end the session.
2.3. Transcranial direct current stimulation
Stimulation was delivered using a certified battery-driven
direct-current stimulator (DC-STIMULATOR PLUS, Neuro-
Conn, Illmenau, Germany). Two conductive rubber electrodes

Fig. 1 e Experimental procedure and tDCS. Panel A: Timeline of the experimental session. The session was initiated with a
block including short anamnestic interview, assessment of verbal fluency and schizotypal traits (INI). Cognitive tests were
administered in three blocks: baseline (BSL), online (ON), and offline (OFF) with respect to stimulation (tDCS). After the BSL
block, the stimulation was set up (Setup). Immediately after stimulation onset and offset, participants filled scales and
rested shortly (S). Panel B: Schematic depiction of electrode placement. Anode (red) and cathode (blue) are plotted with
respect to positions of the 10-10 system. Panel C: Simulated electric field intensity for tDCS. The models show that PFC, but
not TPC, stimulation montage induced peak electric field within left lateral PFC (highlighted region 1), whereas both
montages induced comparable electric fields in the left temporal cortex (highlighted region 2).
 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6
(5  7 cm2) were attached on the head using an EEG cap, elastic
bands, and conductive electrode paste (Ten20, Weaver and
Co., Aurora, CO, USA). For the PFC stimulation, the anode was
located in-between F3 and AF3 and cathode was centered
between T7 and P5 of the international 10-10 system of EEG
electrode placement. For the TPC stimulation, the anode and
cathode were attached between T8 and P6 and between T7
and P5, respectively (see Fig. 1B). Both active conditions were
set to 2 mA (0.057 mA/cm2), applied for 25 min, including an
additional 60 sec ramp-up and ramp-down period. The
placement of electrodes was guided by electric field intensity
models (estimated using SimNIBS version 3) to achieve left
PFC polarization in the PFC stimulation but not in the TPC
stimulation, while keeping the polarization of temporal and
parietal regions of the left hemisphere equal with both mon-
tages (see Fig. 1C). In the sham stimulation, the montage was
either that of PFC or TPC (balanced across participants), but
the full intensity current was delivered only for 50 sec. The
data-collector was not aware whether real or sham simulation
was applied (double-blinding). After the ramp-up period,
participants reported adverse effects using 6 Likert scales
(“itching”, “pinching”, “tingling”, “burning”, “tickling”, and
“pain”, rated from “not at all” e 0 to “very much” e 4). Overall,
tDCS was well-tolerated and there were no statistically sig-
nificant differences in the adverse scores among the groups
[mean ± SE ¼ .90 ± .11, .77 ± .09, and .77 ± .09, for sham, PFC
and TPC, respectively; one-way ANOVA: F(2,116) ¼ .56,
p ¼ .571, hp2 ¼ .01]. At the end of the stimulation, participants
were asked to rate the perceived intensity of the stimulation
using a Likert scale (“none” e 0 to “very strong” e 10). The
groups did not significantly differ in the perceived stimulation
intensity [one-way ANOVA: F(2,118) ¼ 1.48, p ¼ .233, hp2 ¼ .02],
indicating that the blinding of the participants was successful.
2.4. Cognitive assessment
Each measurement block (i.e., baseline, online-stimulation
and offline-stimulation) included three cognitive tests
administered in a fixed order: interference digit-span task, the
associative chain test, and semantic judgement task.
The interference digit span task was used to assess work-
ing memory capacity (WMC). Participants listened to series of
digits presented either in high or low pitch. They were
instructed to attend and remember high-pitch targets and
ignore low-pitch distractors. The stimuli were presented in
500 msec intervals in alternating pitch (always starting with a
low-pitch distractor). At the end of each trial, acoustic beep
signalized that participants should report the remembered
items in the reverse order. The starting span was set to 3,
which was increased by 1 after each correct recall or
decreased by 2 after each incorrect recall. The task continued
until the total number of 5 incorrect trials were made. The
overall capacity was then calculated as the average span of
the five incorrect trials minus 1. Cronbach's a indicated an
acceptable internal consistency of the test in each assessment
block (a ¼ .81 to .84).
Lexicalesemantic retrieval was assessed using the asso-
ciative chain test (ACT; Marko. et al., 2019b). In this test, par-
ticipants continuously generated word chains according to
299
specific conditions, starting with a word stimulus. In the
associate condition, participants produced words so that each
new response was semantically associated to the previous
one (e.g., “Hospital e Doctor e Health e Sport…”) for 50 sec.
Unrelated responses were considered as errors. In the disso-
ciate condition, participants were instructed to generate a
chain of semantically unrelated words, i.e., each new
response should not relate to the previous one (e.g., “Teacher e
Kitchen e Hockey e Apple…”) for 70 sec. Participants were
instructed that delivering a related word would count as an
error. In the associateedissociate condition, participants were
asked to deliver associations and dissociations in alternation
(i.e., switching between the two previously described condi-
tions after each response; e.g., “Phone e Call e Banana e Mon-
key…”) for 120 sec. All three conditions were repeated three
times within each assessment block in the same order. Par-
ticipants entered the words via keyboard, they were instruc-
ted to keep fluent word production, ignore grammatical or
typing errors, and not repeat words within the same chain.
Each response was assessed for response time (RT), i.e., the
latency of entering first letter of the word. The RTs were
structured by two independent factors with two levels each:
the response type (associative or dissociative) and the sequence
type (fixed or alternating) and averaged for each combination
of the two factors separately for each individual and assess-
ment block. Three main measures were then derived from the
averaged RT values: (i) response initiation, calculated as the
average RT in associative fixed and associative alternating re-
sponses [i.e., response initiation RT ¼ (associative fixed
RT þ associative alternating RT)/2]; (ii) inhibition cost, indicating
the RT difference between delivering dissociative versus asso-
ciative responses in the fixed sequence type (i.e., inhibition
cost ¼ dissociative fixed RT e associative fixed RT); and (iii)
switching cost, calculated as the RT difference between alter-
nating versus fixed sequences in the dissociative response type
(i.e., switching cost ¼ dissociative alternating RT e dissociative
fixed RT). See Fig. 2 for further details.
Fig. 2 e Panel A: Response times in ACT by response type
and sequence type. Panel B: Inhibition cost in the fixed
(FIX) and alternating (ALT) sequence type. Panel C:
Switching cost in the associative (A) and dissociative (D)
response type. Error bars represent ± SE; ***p < .001, ns e
non-significant difference.
300 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6
Semantic judgement task (SJT) was completed to assess
post-retrieval response monitoring, response selection and
decision making. Each assessment block included 7 trials
presented in random order. In each trial, participants were
introduced with a trial word (displayed in the upper half of the
screen throughout the trial) and a set of 21 probe words pre-
sented sequentially in the lower half on the screen in random
order. Following the onset of each probe, participants were
required to decide whether the displayed word pair (i.e., the
trial word and the current probe) was semantically related
(yes/no response using computer keyboard). After each
judgement, next probe was presented. The set of 21 probes
words included three types of stimuli: dominant associates
(n ¼ 3), remote associates (n ¼ 6), and unrelated words (n ¼ 12);
the lexical-semantic strengths of the word pairs were derived
from a large corpus and matched across the assessment
blocks (see supplementary Table S1 in the Supplementary
information online for more details). For each stimulus type,
average RT and detection rate (proportion of “yes” responses)
was calculated. These measures showed an acceptable inter-
nal consistency across the assessment blocks, as indicated by
Cronbach's a (aRT ¼ .90 to .96 and a% ¼ .79 to .94).
Verbal fluency (which was assessed in the initial block
only) was estimated using three distinct conditions, each
including two trials. In all trials, participants were asked to
deliver as many instances of a category (animals and super-
market items), letters (F, K), or designs (things for drawing/
painting, things that can make a hole) as possible within 1 min
each. The performance index was calculated as the sum of
responses across the six trials (Cronbach's a ¼ .82). The
fluency performance was not statistically different across the
tDCS groups [one-way ANOVA: F(2,118) ¼ 1.05, p ¼ .352,
hp2 ¼ .02], indicating no pre-existing group differences.
2.5. Self-reported measures
Schizotypal personality questionnaire (SPQ; Raine, 1991;
Slovak version: Chylova  et al., 2017) and Community
Assessment of Psychic Experiences Questionnaire (CAPE;
c
Stefanis et al., 2002; Slovak version: Forga
ova
 et al., 2019)
was completed at the beginning of the experimental session
in order to assess schizotypal personality traits as control
measures since they have been previously associated with
altered lexicalesemantic processing (Merten, 1993; Kiang
et al., 2010). One-way ANOVAs showed no significant dif-
ferences in the schizotypal dimensions across the tDCS
groups, F(2,116) < .25, p > .781, hp2 < .01 for the SPQ and
F(2,116) < 1.52, p > .223, hp < .03 for CAPE scales. Further-
2 response type, F(1,120) ¼ 260.25, p < .001, hp2 ¼ .684, sequence
more, short scales were administered at the beginning of
each cognitive assessment block in order to evaluate
possible differences in participants' affect state. The scale
included nine Likert items (0 e “not at all”, 4 e “very much”),
which were averaged to form three scores: distress (“stress”,
“tension”, and “insecurity”; Cronbach's a ¼ .81 to .84), arousal
(“energy”, “arousal”, and “vigilance”; Cronbach's a ¼ .82 to
.92), and engagement (“motivation”, “interest”, and “enthu-
siasm”; Cronbach's a ¼ .82 to .90). There were no statistically
significant differences in the baseline affect scores across
the tDCS groups [one-way ANOVA: F(2,116)  1.57, p  .213,
hp2  .026; see supplementary Table S2 in the Supplementary
information online for more details].
2.6. Data processing and analysis
The data were processed in SPSS (IBM Corp, 2017) and JASP
(JASP Team, 2019). As in the previous studies (Marko et al.,
2019a; Marko et al., 2019b), the error rate in the ACT was
very low (<5%), and thus only RT was analyzed. First, a 2  2
repeated measures (RM) ANOVA was computed to assess the
effect of response type (associative vs dissociative) and sequence
type (fixed vs alternating) for baseline ACT data. Next, full
factorial 3  3 RM ANOVAs, including a between-subjects
factor stimulation (sham/PFC/TPC) and a within-subjects fac-
tor block (baseline/online/offline) were computed for ACT and
WMC measures. The ANOVA for SJT included an additional
within-subjects factor trial type (dominant/remote/unrelated).
Greenhouse-Geisser sphericity correction was applied where
appropriate. The effects of tDCS on the cognitive measures
were assessed as the respective block  stimulation interaction
in the ANOVA models. To evaluate significant interactions,
difference values of the cognitive performance between the
baseline and online assessment block (e.g., for response time,
DRTON-BSL) and between the baseline and offline block (e.g.,
DRTOFF-BSL) were calculated. These difference values were
then compared across the tDCS conditions (i.e., groups) using
planned pairwise one-sided t-tests to test the hypotheses. The
changes in performance between online and offline block (e.g.,
DRTOFF-ON) were assessed in an exploratory manner with two-
sided t-tests. In order to account for familywise error due to
multiple testing, p-values of the pairwise comparisons across
all measures were adjusted by sequential Bonferroni correc-
tion (Cramer et al., 2016; adjusted p-values are reported). Effect
sizes for ANOVA and t-tests were estimated using hp2 and
Cohen's d, respectively. Finally, an exploratory analysis was
performed to assess relations between the dependent cogni-
tive measures (ACT, WMC, and SJT) using Pearson
correlations.
3. Results
3.1. Baseline semantic retrieval measures
A RM ANOVA of RTs in ACT revealed significant effects of
type, F(1,120) ¼ 44.01, p < .001, hp2 ¼ .268, and their interaction,
F(1,120) ¼ 31.62, p < .001, hp2 ¼ .209, showing that the
switching cost was evident only in dissociative but not in
associative responses (see Fig. 2). Pearson correlation analysis
of the baseline measures showed that response initiation was
not significantly correlated with inhibition cost, r(120) ¼ .010,
p ¼ .916, or switching cost, r(120) ¼ .052, p ¼ .571. Inhibition cost
and switching cost were negatively correlated, r(120) ¼ .378,
p < .001. Verbal fluency was significantly correlated only with
 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6 301

response initiation, r(120) ¼ .552, p < .001. None of the ACT switching cost and inhibition cost was negative (average
measures was significantly correlated with WMC (see r ¼ .437, SD ¼ .13). Complete reports for all the RM ANOVA
supplementary Table S3 in the Supplementary information models and tests are shown in Supplementary Tables S4 and
online for the complete report of the correlation analysis). S5 and Figure S1 in Supplementary information online.

3.2. Effects of tDCS on semantic retrieval measures 3.3. Effects of tDCS on control measures

The effects of tDCS on the derived ACT measures were eval-
uated as the block  stimulation interaction effects using RM
ANOVAs and planed pairwise t-tests on RT change values
(DRTON-BSL and DRTOFF-BSL) between the tDCS groups. For
response initiation, the block  stimulation interaction was not
significant, F(3.4,202.6) ¼ .15, p ¼ .945, hp2 ¼ .003 (Fig. 3A). For
inhibition cost, the block  stimulation interaction was signifi-
cant, F(4,236) ¼ 4.62, p ¼ .001, hp2 ¼ .073 (Fig. 3B). The com-
parisons of the difference scores showed that PFC tDCS
significantly decreased inhibition cost during (online block)
and after the stimulation (offline block) compared to baseline
(one-sided hypothesis, t-test: p ¼ .003, d ¼ .658 and p ¼ .003,
d ¼ .693, respectively). The change in inhibition cost during
and after TPC stimulation with respect to baseline were not
significant (two-sided t-test: p ¼ .394, d ¼ .215 and p ¼ .395,
d ¼ .223, respectively). Finally, for switching cost, block 
stimulation interaction was not significant, F(4,236) ¼ .45,
p ¼ .771, hp2 ¼ .008 (Fig. 3C). Overall, across the tDCS groups
and pairwise contrasts, the correlation between the change in
A RM ANOVA for working memory capacity showed a significant
block by stimulation interaction, F(3.6,214.8) ¼ 4.94, p < .001,
hp2 ¼ .077 (Fig. 3D). The comparisons of the difference scores
showed that PFC tDCS significantly increased WMC during
(one-sided hypothesis, t-test: p ¼ .006, d ¼ .660) and after the
stimulation (one-sided hypothesis, t-test: p ¼ .007, d ¼ .525)
compared to baseline assessment. The changes during and
after TPC tDCS were not significant (two-sided t-test: p ¼ .684,
d ¼ .089 and p ¼ .374, d ¼ .200, respectively). A RM ANOVAs for
semantic judgement detection rate (%) and semantic judgement
response time (RT) involved an additional within-subjects
factor trial type (dominant/remote/unrelated). Overall, as ex-
pected, the analysis revealed that the detection rate was
significantly higher for the trials including dominant associ-
ation (~73 ± 2.1%) than in the trials including remote associ-
ation (~48 ± 2.6%) or unrelated word pairs (~10 ± 1.6%).
Furthermore, the RTs were longer for remote as compared to
both dominant and unrelated trials. However, tDCS did not
significantly affect or interact with any of the two SJT

Fig. 3 e The effects of tDCS on ACT measures (panels A e C) and working memory capacity (panel D). Each panel shows
mean difference scores with respect to baseline (ON-BSL and OFF-BSL) as a function of tDCS (between subject factor).
Descriptive plots indicating distributions and individual values for each participant are presented in Supplementary
information (Supplementary Figure S1). PFC e prefrontal tDCS (red), TPC e temporoparietal tDCS (blue), BSL e baseline block,
ON e online block, OFF e offline block. Error bars represent ± SE. ***p < .001, **p < .01, ns e non-significant difference (p-
values of the pairwise contrasts were corrected for multiple comparisons using sequential Bonferroni correction).
302 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6
measures (see supplementary Table S6 and supplementary
Figures S2 and S3 in the Supplementary information online
for full report of the SJT data).
Finally, we evaluated whether the improvement in WMC
could account for the improvement in inhibition cost by tDCS
of the PFC. A negative correlation would be expected if WMC
mediated the effects of PFC tDCS on inhibition cost, as smaller
inhibition cost represents better performance. Pearson cor-
relations showed that the association between the difference
scores in WMC and inhibition cost was positive and not sig-
nificant for both online (r ¼ .258, p ¼ .107) and offline block
(r ¼ .280, p ¼ .081).
4. Discussion
The goal of the present study was to investigate the involve-
ment of the left lateral PFC in controlled semantic cognition.
Based on the recent discoveries in the field of semantic
cognition (Ralph et al., 2017; Chiou & Lambon Ralph, 2019;
Chiou et al., 2018), we hypothesized that excitatory anodal
tDCS targeting the left lateral PFC would enhance controlled
but not automatic semantic processing. In line with this pre-
diction, we observed that the processing time for semantic
inhibition (i.e., inhibition cost) decreased during and shortly
after the stimulation of PFC while this stimulation had no
significant effect on free-associative retrieval that does not
considerably engage semantic control. On the other hand, a
control stimulation of temporoparietal cortex did not show
such an effect, suggesting specificity of the prefrontal tDCS.
Contrary to our expectations, however, we found no signifi-
cant effect of the prefrontal tDCS on flexible semantic
switching (i.e., switching cost), which has been also consid-
ered a semantic control function (Troyer et al., 1998; Unsworth
et al., 2011).
Our study replicated previous findings (Marko et al., 2019a;
Marko et al., 2019b) showing that the production of semanti-
cally unrelated responses requires substantially more pro-
cessing time than the continuous production of free word
associations, which has been attributed to increased demands
on semantic control. In particular, we but also others (Allen
et al., 2008; Collette et al., 2001) have argued that the addi-
tional processing time in dissociative tasks reflects the sup-
pression of automatically evoked prepotent associations.
Thus, shorter inhibition cost in response to the excitatory
tDCS suggests that left lateral PFC may be a part of the neural
mechanism that regulates stimulus-driven semantic activa-
tion. We have also confirmed that alternating the retrieval
rule further increased response latencies (i.e., switching cost)
for dissociative but not associative retrieval. This interaction
was expected as the dissociative retrieval requires constant
inhibition to control habitual associates, and thus less re-
sources remain available to resolve the demands on flexible
switching between semantic sets (Kiesel et al., 2010; Marko M.
et al., 2019a; Marko et al., 2019b), resulting in larger difficulty of
the dissociative performance. However, the anodal tDCS did
not affect both semantic inhibition and switching, indicating
that they involve at least partially specific cognitive processes
(as in the case of domain-general inhibition and switching, see
Friedman & Miyake, 2017; Karr et al., 2018). Prefrontal tDCS
thus seem to interact with the prefrontal computations sup-
porting inhibition, i.e., the ability to suppress strong semantic
activation, but not with the computations supporting a flex-
ible shifting between remote semantic sets, which is likely
underpinned by partially distinct brain circuits (Lemire-
Rodger et al., 2019; Niendam et al., 2012). Yet, the negative
correlation between semantic inhibition and switching sug-
gests that these processes may either compete for a limited
domain-general resources or interfere with each other
(Fedorenko & Thompson-Schill, 2014; Chai et al., 2016;
Hagoort, 2005). This account is consistent with the proposal
that the brain networks supporting semantic control are
dynamically coupled with or de-coupled from the domain-
general executive brain network depending on current task
demands (Fedorenko & Thompson-Schill, 2014; Sabb et al.,
2007). Despite the fact that inhibition and switching may tap
on shared resources, our data indicate that they represent two
distinct components of the semantic control architecture, and
that the computations in the left lateral PFC are specifically
linked to inhibitory processing (see Allen et al., 2008; Collette
et al., 2001), but not switching.
An alternative account for the present data could be that
semantic inhibition and switching are implemented by the
same (or substantially overlapping) neural network involving
left lateral PFC, but the tDCS altered the operation of this
functional network from one state/function to another. This
account draws upon the evidence that tDCS can bias compu-
tations within functional networks and so enhance one pro-
cess or function at the cost of other processes (Bikson and
Rahman, 2013; Rahman et al., 2013). Following this rationale,
PFC tDCS could have redistributed the available neural and
cognitive resources of the semantic control network1 and so
biased its operation towards a more constricted (inhibitory)
rather than exploratory (switching) mode of processing. Such
case would not exclude that inhibition and switching share a
common neural and cognitive ground. Hence, it could also be
speculated that a distinct electrode montage or stimulation
type could favor flexible switching or a more exploratory
mode of semantic processing. Further research is thus needed
to shed light on the precise neural and cognitive computations
involved in the inhibition of semantic activation and switch-
ing between semantic sets as well as their interaction.
Consistently with previous studies (Ruf et al., 2017; Ohn
et al., 2008; Zaehle et al., 2011; Brunoni & Vanderhasselt,
2014), the prefrontal tDCS improved working memory capac-
ity. Interestingly, however, this effect was not related to
enhanced inhibition cost due to tDCS, indicating that an
extended working memory span does not necessarily trans-
late into a better ability to suppress habitual associates
(Baddeley, 2012). In fact, our study showed that improved
ability to maintain and operate on the information in working
memory was associated with marginally increased inhibition
costs, suggesting that improved maintenance of information
in phonological loop may hinder the ability to disentangle
from the intrusive semantic associates. Thus, given our
1
This principle suggests that, instead of referring to tDCS as
either depolarizing or hyperpolarizing, in some cases it may be
more accurate to describe tDCS as redistributing polarization
within the targeted neuronal assembly.
 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6
findings, we can reject the alternative hypothesis that se-
mantic inhibition was enhanced as a consequence of
improved working memory capacity.
On the other hand, prefrontal tDCS did not significantly
modulate semantic relatedness judgement. This finding sug-
gests that the improvement in inhibition was not related to
post-retrieval response monitoring and evaluation of already
retrieved word candidates. This indicates that the improve-
ment to inhibition by prefrontal tDCS was not due to a mod-
ulation of participants' response criterion, i.e., an increased
threshold to evaluate the candidate words as “associated”
versus “not-associated”. Based on the results from these
control measures we conclude that the observed changes of
semantic inhibition elicited by the left-lateral prefrontal tDCS
reflect a modulation of inhibitory processing that shapes the
activation or access to semantic representations (i.e., a pro-
active inhibitory process) or implements a post-retrieval
rejection of inappropriate candidates (i.e., a retroactive
inhibitory process). A more fine-grained experimental
manipulation will be required in the future studies to further
specify the precise nature of the inhibitory processing that is
sensitive to left-lateral prefrontal tDCS.
Our study also has limitations that should be considered.
Previous neuroimaging studies showed that controlled se-
mantic cognition involves partially distinct computations that
are putatively implemented by specific parts of the left lateral
PFC. For instance, Badre et al. (Badre. et al., 2005; Badre &
Wagner, 2007) suggested that left anterior ventrolateral PFC
(pars orbitalis) interacts with activated semantic representa-
tions and so supports controlled access to stored knowledge,
whereas left middle ventrolateral PFC (pars triangularis) im-
plements a more general mechanism that operates after
retrieval to resolve response competition. Due to low
anatomical specificity of tDCS, it is not clear to what extent
these two putative mechanisms could be modulated. Relat-
edly, as tDCS may affect larger functional networks (Bikson,
Name, & Rahman, 2013; Pisoni et al., 2018), the effects of PFC
tDCS could affect functional coupling of left lateral PFC with
other areas involved in semantic cognition, such as anterior/
middle temporal cortex (that support semantic representa-
tion) or parietal cortex (that may also support semantic con-
trol; Ralph et al., 2017; Chiou & Lambon Ralph, 2019; Badre
et al., 2005; Whitney et al., 2011). It is important to note that
the effect of anodal PFC stimulation could be conditioned on
or interact with the effects of cathode, which was placed over
left temporoparietal regions. Thus, future studies involving
neuroimaging should investigate the possible modulation of
the left lateral prefronto-temporal and/or prefrontoeparietal
interactions and their role in controlled semantic retrieval
(high-density tDCS may be suitable for a more detailed opti-
mization of the stimulation protocol in further studies).
Notably, although we did not find significant effect of the TPC
condition on the selected semantic measures, previous
research has suggested that right temporoparietal regions
may support coarse and diffuse semantic encoding, which
underpins creativity and insightful thinking (Jung-Beeman,
2005; Kounios & Beeman, 2014; Luft et al., 2018). We might
not be able to find such effect as our measures were focused
303
on the processing speed (RT) rather than originality of the
responses. Research utilizing measures sensitive to creative
semantics and response quality may provide more insight
into the role of the right hemisphere in semantic information
processing. Finally, from another methodological perspective,
between-subject experimental design could be considered a
limitation due to possible pre-treatment group differences.
However, we addressed this concern by assessing baseline
performance of each participant (showing no statistically
significant group differences) and performing statistical tests
of tDCS effects using the baseline-corrected (change) scores.
5. Conclusions
Using tDCS in a double-blind randomized controlled experi-
ment, we have demonstrated that the left lateral PFC supports
controlled semantic retrieval by exerting inhibition of habit-
ually activated semantic representations. The effects of tDCS
were specific for response inhibition while switching or
automaticeassociative retrieval was unaffected, and were
elicited by prefrontal but not temporoparietal stimulation.
Furthermore, we have shown that the enhanced semantic
inhibition was not due to a general improvement in the
cognitive ability (working memory capacity) or post-retrieval
response monitoring and evaluation (semantic judgement
speed or criterion). Taken together, our findings indicate that
the left lateral PFC may be specifically involved in a filtering
process that sculptures the space of available responses
within semantic representation system (i.e., a proactive pre-
retrieval inhibitory mechanism) or rejects/selects already
retrieved response candidates (i.e., a retroactive post-retrieval
inhibitory mechanism). The recognition of this inhibitory
process and the possibility of its modulation using tDCS could
eventually inspire novel biological treatments targeting
altered semantic processing.
Author contributions
M.M.: Conceptualization, Methodology, Software, Formal
analysis & Visualization, Investigation, Writing - Original
Draft, Writing - Review & Editing, Project administration,
Funding acquisition. I.R.: Conceptualization, Supervision,
Funding acquisition, Writing - Original Draft, Writing - Review
& Editing. Note: Lenka Hapa kova , Petra Petra

sova
 , and Dra-
homı́r Michalko helped with data collection under supervision
of the authors.
Research transparency
We report how we determined our sample size, all data ex-
clusions, all inclusion/exclusion criteria, whether inclusion/
exclusion criteria were established prior to data analysis, all
manipulations, and all measures in the study. Data analyzed
during the current study as well as supplementary digital
study materials and codes are stored on accessible repository
304 c o r t e x 1 3 4 ( 2 0 2 1 ) 2 9 6 e3 0 6
(OSF: https://osf.io/82rw3/). No part of the study procedures or
analyses was pre-registered prior to the research being
conducted.
Open practices
The study in this article earned Open Materials and Open Data
badges for transparent practices. Materials and data for the
study are available at https://osf.io/82rw3/.
Declaration of competing interest
None.
The authors declare that the research was conducted in the
absence of any commercial or financial relationships that
could be construed as a potential conflict of interest.
Acknowledgements
This work was supported by the Scientific Grant Agency of the
Ministry of Education, Science, Research and Sport of the
Slovak Republic (projects no. VEGA 2/0059/20 and VEGA 2/
0170/19) and the Slovak Research and Development Agency
(projects no. APVV-19-0570 and APVV-16-0202). We thank
Lenka Hapa  kova
, Petra Petra

sova
, and Drahomı́r Michalko for
help with data collection.
Supplementary data
Supplementary data to this article can be found online at
https://doi.org/10.1016/j.cortex.2020.11.001.
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