Received: 29 May 2022 Revised: 28 December 2022 Accepted: 20 April 2023

DOI: 10.1002/hbm.26347

RESEARCH ARTICLE

A new perspective on the role of the frontoparietal regions in

Stroop-like conflicts

Noga Oren 1 | Donna Abecasis 1 | Edna Inbar 2,3 | Amir Glik 3,4,5 |
3,5 1,3,6
Israel Steiner | Irit Shapira-Lichter

1
Functional MRI Center, Beilinson Hospital,
Rabin Medical Center, Petach Tikva, Israel Abstract
2
Imaging Department, Beilinson Hospital, Humans are goal-directed; however, goal-unrelated information still affects us, but
Rabin Medical Center, Petach Tikva, Israel
3
how? The Stroop task is often used to answer this question, relying on conflict
Sackler Faculty of Medicine, Tel Aviv
University, Tel Aviv, Israel (incongruency) between attributes, one targeted by the task and another irrelevant
4
Cognitive Neurology Clinic, Beilinson to the task. The frontal regions of the brain are known to play a crucial role in proces-
Hospital, Rabin Medical Center, Petach Tikva,
Israel
sing such conflict, as they show increased activity when we encounter incongruent
5
Department of Neurology, Beilinson Hospital, stimuli. Notably, the Stroop stimuli also consist of conceptual dimensions, such as
Rabin Medical Center, Petach Tikva, Israel semantic or emotional content, that are independent of the attributes that define the
6
Sagol School of Neuroscience, Tel Aviv
conflict. Since the non-targeted attribute usually refers to the same conceptual
University, Tel Aviv, Israel
dimension as the targeted-attribute, it is relevant to the task at hand. For example,
Correspondence
when naming the emotion of an emotional face superimposed by an emotional word,
Irit Shapira-Lichter, Functional MRI Center,
Beilinson Hospital, Rabin Medical Center, both the targeted-attribute and the non-targeted attribute refer to the conceptual
Petach Tikva 49100, Israel.
dimension “emotion”. We designed an fMRI paradigm to investigate how conflicts
Email: iritlichter@yahoo.com
between different conceptual dimensions impact us. Even though the conflict was
task-irrelevant, incongruent stimuli resulted in longer reaction times, indicating a
behavioral congruency effect. When examining the neural mechanisms that underlie
this effect, we found that the frontal regions exhibited repetition suppression, while
the bilateral intraparietal sulcus (IPS) showed a congruency effect linked to the
behavioral effect. Taken together, these findings suggest that individuals are unable
to completely ignore task-irrelevant information, and that the IPS plays a crucial role
in processing such information.

KEYWORDS
attention, conflict adaptation, conflict monitoring, congruency effect, intraparietal sulcus,
repetition suppression

1 | I N T RO DU CT I O N but still affects us. One of the most prevalent methods for studying
this phenomenon is the Stroop task. The stimuli used in the Stroop
In everyday situations, we are constantly bombarded with information task are defined by the relationship between two attributes, such as
that is not related to our current tasks, responsibilities, or objectives, the word form or the color in color-word. Participants are instructed

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© 2023 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.

4310 wileyonlinelibrary.com/journal/hbm Hum Brain Mapp. 2023;44:4310–4320.

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OREN ET AL. 4311

to attend to one attribute, referred to as the targeted attribute in this
article, and ignore the other attribute, which is typically considered
task-irrelevant. The attributes can be compatible (congruent) or
incompatible (incongruent) with each other. Congruency is known to
affect performance (MacLeod, 1991): judging the color of a word is
harder when the word itself spells another color (color-word Stroop);
judging the content of a picture is harder when a superimposed word
indicates a different content (picture-word Stroop); judging emotional
facial expression is harder when a superimposed word indicates a dif-
ferent emotion (emotional picture-word Stroop).
Furthermore, the Stroop stimuli also consist of conceptual dimen-
sions such as semantic content, emotional content, and so forth
(Hatukai & Algom, 2017). Conceptual dimensions differ from the sti-
muls attributes. Hence, the non-targeted, so-called task-irrelevant
attribute in standard Stroop tasks is related to the task at hand, since
it refers to the same conceptual dimension as the targeted attribute
and the task itself (Kornblum, 1992). For example, in the emotional
picture-word Stroop, both the targeted attribute (emotional expres-
sion) and the non-targeted attribute (emotional word) provide infor-
mation about the conceptual dimension the task refers to (emotion;
Figure 1a). Here, we define task-irrelevant as an attribute referring to
a different conceptual dimension than the task. Can a task-irrelevant
conflict between two attributes in one conceptual dimension influ-
ence our ability to process information in another conceptual dimen-
sion of the same stimulus? That is, would incongruence between
emotional expression and an emotional word affect an animacy judg-
ment of the picture? Below, we present a functional magnetic reso-
nance imaging (fMRI) task involving Stroop stimuli in which the
conflicting attributes refer to one conceptual dimension while the task
refers to another conceptual dimension. We used both traditional

F I G U R E 1 Paradigm.
(a) Uncoupling the conflict from the
task: The current study breaks the
coupling between the conflict and
the task by asking participants to
make an animacy judgment, thus
making the conflict task-irrelevant.
Note that in the figure, the term
dimension refers to the conceptual
dimension, defined as the semantic
field. (b) The event-related fMRI
paradigm presented three types of
stimuli: color-word, picture-word,
and emotional picture-word. Each
stimulus was either congruent or
incongruent, depending on the
relationship between its attributes.
Every word (in the color-word type)
and picture (in the picture-word
types) appeared at least twice, once
in a congruent and once in an
incongruent stimulus. The second
appearance was always in a different
run than the first appearance. In the
figure, this is represented by the
three dots in the timeline.

4312
comparisons between congruent and incongruent conditions and a
repetition suppression approach not yet applied, to the best of our
knowledge, to the study of the neural mechanisms underlying a
Stroop-like task (see below).
Congruency effect, manifested by lower accuracy and longer reac-
tion time (RT) in incongruent compared with congruent trials, is evi-
dent in the color-word (Stroop, 1935), picture-word (MacLeod, 1991)
and emotional picture-word (Etkin et al., 2015) Stroop versions. The
brain regions that are usually more activated for incongruent than
congruent trials include the superior and inferior lateral prefrontal cor-
tices, dorsal anterior cingulate cortex (ACC)/pre-supplementary motor
area (pre-SMA), and the intraparietal sulcus (IPS) (Xu et al., 2016).
Contemporary neuroscientific models highlight the role of the frontal
regions in the Stroop task (Banich, 2019; Botvinick et al., 2001). The
conflict-monitoring model claims that the ACC detects the presence
of a conflict and engages the lateral prefrontal cortices to impose cog-
nitive control (Botvinick et al., 2001). The cascade of control model
postulates that the superior and inferior lateral prefrontal cortices
determine the attentional set and bias the selection towards task-
relevant information; the dorsal ACC/pre-SMA selects and evaluates
the response (Banich, 2019). These models pay relatively little atten-
tion to the IPS and largely ignore behavioral models that emphasize
the role of data-driven selective attention mechanisms (Algom &
Chajut, 2019; MacLeod, 1992), learning, and memory biases
(Schmidt, 2019) or the impact of emotions (Hatukai & Algom, 2017) in
the Stroop task. Since the non-targeted attributes refer to the same
conceptual dimension as the task, the results and models describe the
neural response to task-relevant conflicting information. The current
study aims to portray the neural response to task-irrelevant conflict.
One approach to studying the response to task-irrelevant infor-
mation is the emotional analog of the Stroop task, in which patients
are slower to name the color of a word associated with their clinical
condition (Williams et al., 1996). These results show that even without
a conflict, task-irrelevant information is processed and affects perfor-
mance. Another approach was taken by Hatukai and Algom (2017),
who examined RT for incongruent stimuli in tasks that referred to a
conceptual dimension unrelated to the conflict. For example, partici-
pants judged whether a color word written in a congruent or incon-
gruent ink color was positioned at the top or bottom of the screen
(Experiment 3) or written in a wide or narrow font (Experiment 5); or
judged whether the contour of an outline drawing of a cat or a dog,
with the randomly congruent or incongruent words “cat” or “dog”
embedded within it, was continuous or dashed (Experiment 4). Con-
trary to the traditional congruency effect, they found shorter RT in
the incongruent versus the congruent conditions. They claimed that
incongruent stimuli in their tasks provoked negative affect, driving
people to avoid or terminate them as quickly as possible. Alterna-
tively, it is possible that the avoidance of incongruent stimuli in these
experiments stemmed from the focus of the task on a low-level fea-
ture of the stimulus (e.g., position, width, or outline characteristics).
Indeed, in responding to a higher-level feature that necessitates in-
depth processing and semantic conceptualization, a congruency effect
was evident, although the task referred to another conceptual
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OREN ET AL.
dimension, possibly reflecting the unavoidability of the conflict
(Zhao & Wang, 2013). For example, Zhao and Wang (2013) instructed
participants to name geometric shapes containing color-word Stroop
stimuli. They found a congruency effect manifested as less accurate
responses for incongruent trials. Notably, in Zhao and Wang's (2013)
task, the Stroop stimulus was outside the focus of attention.
In our approach, animacy judgments were made on either congru-
ent or incongruent stimuli in the Stroop paradigm (Figure 1a). Since
the conceptual dimension of the conflicting attributes of the stimuli
(e.g., emotion, color, etc.) and the task (animacy judgment) differed,
according to our definition, the conflict was task-irrelevant. Notably,
an animacy judgment requires in-depth processing of a conceptual
dimension of the stimulus that is not related to the conflict. It should
also be noted that if a congruency effect is found despite the conflict
being task-irrelevant, this effect may still be specific to a certain stim-
ulus type. Therefore, to allow generalization of the results, different
types of stimuli were used in our Stroop-like paradigm.
Using this paradigm, we aimed at characterizing the behavioral
and neural effects of task-irrelevant conflict processing. On top of the
established congruency effect analysis, a repetition suppression analy-
sis was performed as an independent complementary indicator of
brain response to conflict. Repetition suppression—a robust phenome-
non observed across the brain—is typically used to study the neural
representation of information (Barron et al., 2016). Diminished activa-
tion to repeated presentations of a stimulus or information is an index
of the insensitivity of that brain region to that information (Grill-
Spector & Malach, 2001). In the current study, finding decreased brain
activation in response to a change in the relationship between the
attributes of a stimulus from congruent to incongruent indicates
insensitivity to congruency, and vice versa. Although repetition sup-
pression is reliable, valid, and widely used in animal and human studies
of visual perception, and although many Stroop-related regions show
repetition suppression in a variety of tasks and stimuli (Kim, 2017), to
our knowledge, this is the first attempt to use the repetition effect in
the context of a Stroop-like task.
To sum up, our study is original in that it aimed at characterizing
the response to conflicting information related to a different concep-
tual dimension than a demanding task. Behavioral and neural analyses
examined both the standard congruency effect and the repetition sup-
pression effect, applied for the first time in this context. We expected
to find a behavioral congruency effect (Kornblum, 1992; Zhao &
Wang, 2013), and repetition suppression. In frontal brain regions—
known to represent goal-related information (Sreenivasan
et al., 2014), and in the context of the Stroop task, to process task-
related aspects of the conflict (Banich, 2019; Botvinick et al., 2001)—
we expected to find a significant repetition suppression but not nec-
essarily a congruency effect. The IPS was anticipated to demonstrate
a congruency effect, but not necessarily repetition suppression since
it is known to represent information about non-target stimuli unre-
lated to the task (Chao et al., 2011; Geng & Mangun, 2009; Mazaheri
et al., 2011), carrying the potential to represent task-irrelevant conflict
in the present paradigm as well. The present study tested these
hypotheses.

OREN ET AL.
2 | MATERIALS AND METHODS
2.1 | Participants
Young, healthy participants were recruited from the community
between November 2016 and September 2017 using standard ads.
Inclusion criteria: aged between 18 and 55 years, healthy, native
Hebrew speakers. Exclusion criteria: large tattoos, permanent makeup,
metal implants, claustrophobia, a history of neurological or psychiatric
disorders, and a background of learning disabilities, such as attention
deficit hyperactivity disorder (ADHD) and dyslexia.
Thirty-six healthy participants were recruited. One participant
was excluded from the study due to an incidental finding of a brain
abnormality. Seven others were discarded prior to statistical analyses
due to extensive head movements (>2 mm) in more than two runs.
These eight participants were excluded from all behavioral and fMRI
analyses (behavioral results remained the same when the 7 participants
were included in the analyses, see supplementary information S1).
Based on these criteria, analyses were conducted on 28 participants
(17 females; age range: 18–55 years; mean age (standard deviation
[SD]): 31.96 years (2.38); mean education (SD): 14.96 years (2.5); right-
handed: 25 participants were right-handed, 2 left-handed and 1 did not
report). A diverse age range was chosen in order to enhance the gener-
alization of the results while excluding minors and avoiding the poten-
tial slowing and cognitive effects of aging. Handedness was not an
exclusion criterion since, based on the literature (Xu et al., 2016), we
assumed that the neural response to Stroop stimuli is bilateral. This
assumption was later supported by the data. The experimental proce-
dures were approved by the Beilinson Hospital Institutional Review
Board, and all participants provided written informed consent and
received nominal monetary remuneration for their participation.
2.2 | MRI acquisition
MRI scans were performed on a 3.0 Tesla MRI scanner (Magnetic Reso-
nance Imaging system, Ingenia 3 T 517, Philips, Best, The Netherlands)
using a 32-channel head coil. Blood-oxygen-level-dependent (BOLD)
functional MRI was acquired with T2*-weighted sequence: repetition
time (TR) = 2000 ms; echo time (TE) = 35 ms; flip angle (FA) = 90 ;
field of view (FOV) of 230 mm  251 mm  105 mm; 35 axial slices of
3 mm thickness, 0 gap. A high-resolution anatomical T1-weighted fast
spoiled gradient-echo imaging was acquired, with FOV dimensions of
240 mm  240 mm  166 mm, TR of 8.2 ms, and TE of 3.8 ms. The
anatomical scan was used for cortical surface reconstruction. The
experiment was performed using Presentation software
(Neurobehavioral Systems, Inc., Berkeley, CA; Version 17.0, www.
neurobs.com) and presented via an LCD projector to a tilted (45 ) mir-
ror positioned over the participant's forehead.
2.3 | Stimuli, paradigm, and procedure
The experimental paradigm was a visual Stroop-like task. It consisted
of eight runs with 24 trials per run, summing up to 192 trials per
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4313
subject. There were three types of stimuli (Figure 1b): (1) color-word
(color name written in color ink; 24 stimuli across 8 runs, 12.5% of the
trials); (2) picture-word (a picture of an emotionally neutral object or
person with a neutral adjective word superimposed on it; 120 stimuli
across 8 runs, 62.5% of the trials); (3) emotional picture-word
(a picture of an emotional face with an emotional word superimposed
on it; 48 stimuli across 8 runs, 25% of the trials). For more details
regarding the stimuli and how they were chosen, see supplementary
information S1.
This paradigm manipulated congruency (congruent/incongruent
condition), as the word could either be congruent or incongruent with
the color ink (in the color-word type) or with the picture meaning
(in the picture-word and emotional picture-word types). Repetition
was included as an independent two-level factor (first/non-first) since
each word (in the color-word type) and picture (in the other two
types) appeared at least twice (once as congruent and once as incon-
gruent), with the order counterbalanced across participants. The par-
ticipants' task was to determine whether a visual stimulus was
animate or inanimate (Figure 1b). Therefore, the conflict between the
Stroop attributes was not relevant to the task and referred to a differ-
ent conceptual dimension. It is worth noting that the three types of
stimuli differed in terms of animacy: the color-word type had only
inanimate stimuli, the picture-word type had both animate and inani-
mate stimuli, and the emotional picture-word type had only animate
stimuli. Importantly, diverse stimuli were used solely to enhance gen-
eralizability rather than to test the effect of stimulus type, so the three
types were collapsed together in all analyses. Moreover, confirmatory
analyses were conducted to ensure there was no bias due to the dif-
ferences in stimulus types, and these results are presented in detail in
supplementary information S1.
The design was slow event-related, divided into eight runs. Each
run started with an 18 s fixation cross followed by repeated
presentations of stimuli, each appearing for 2 s and a fixation cross for
10 s. The order of the runs and stimuli was randomized, with the fol-
lowing restrictions: (1) stimuli from the same type (e.g., color-word)
were not presented in consecutive trials; and (2) for each stimulus, its
congruent appearance was presented in a different run than the
incongruent appearance, in order to minimize the visibility of the rep-
etition manipulation. As a result, stimuli that appeared for the second
time (non-first stimuli) tended to be presented in later runs. A control
analysis was specifically designed to dismiss the possibility that prac-
tice interfered with the hypothesized repetition effect, as described in
supplementary information S1. Participants responded to the animacy
task by pressing a button on an MRI-compatible response box.
2.4 | fMRI preprocessing
fMRI preprocessing was performed using SPM8 software (http//:
www.fil.ion.ucl.ac.uk/spm, The Wellcome Centre for Human Neuro-
imaging, UCL, London, UK). The first six volumes in each run were dis-
carded to allow for steady-state magnetization. The remaining
volumes underwent preprocessing, including slice scan time correc-
tion, 3D motion correction, coregistration, normalization to the

4314
Montreal Neurological Institute (MNI) coordinate system, and spatial
smoothing using an 8 mm full-width at half-maximum Gaussian kernel.
A high-pass filter with a 128 s cutoff was applied to the voxel-wise
time courses, which were then globally normalized using session-
specific grand mean scaling. In addition to the seven participants
excluded because of head movements, another four participants had
one or two runs discarded due to head movements larger than 2 mm
or technical problems.
2.5 | Defining regions of interest
We aimed to examine the fMRI activations that would emerge in the
current Stroop-like task relative to the established knowledge regard-
ing the standard Stroop patterns. Therefore, the fMRI analysis was
performed on predefined regions of interest (ROIs) of the frontoparie-
tal network, using coordinates obtained from a meta-analysis that
focused on the Stroop task. We chose the Xu et al. (2016) meta-
analysis since it included studies of non-emotional color-word Stroop
and emotional picture-word Stroop, similar to the stimuli in our para-
digm. Since we were interested in studying the effect of interference
regardless of emotionality, we focused on the analysis conducted
across those stimuli (first section of Table 3, page 10 in Xu et al.'s
(2016) paper).
ROIs were defined as follows: (1) A sphere of 10 mm was drawn
around each of the 13 coordinates identified by Xu et al. (2016)
(Table 1); (2) adjacent spheres with overlap > 10 voxels were united
into a single ROI; (3) homologous regions were united after confirming
that the correlation between them was significantly higher than the
averaged correlation between all possible pairs of regions. This was
determined in the following way: (a) the time course was extracted
from each region and run (6 to 8 runs per participant); (b) for every
participant, correlation was calculated between every possible pair of
T A B L E 1 Coordinates taken from the Xu et al. (2016) meta-
analysis were used to define a 10 mm sphere.
Name Coordinates (MNI, x,y,z)
Bilateral intraparietal sulcus 28, 64,46
32, 48,44
38, 50,46
Bilateral middle frontal gyrus 48,18,22
40,10,26
46,18,32
48,22,20
Bilateral pre-SMA 6,20,42
6,8,54
Right inferior frontal gyrus/insula 36,22, 12
32,28,0
38,18,6
42,16,0
Note: Each ROI is composed of several coordinates because overlapping
and homologous spheres were united.
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OREN ET AL.
regions, Fisher transformed, and averaged across runs (i.e., resulting in
a single correlation per participant per pair of regions); (c) for every
participant, the averaged correlation of all possible pairs of regions
was calculated; (d) across participants, the difference between the
correlation of a pair of homologous regions and the averaged correla-
tion of all possible pairs of regions was examined using paired samples
t tests. Three pairs of homologous regions were examined. In all of
them, the correlation between the homologous regions was signifi-
cantly higher than the average correlation of all possible pairs: left and
right IPS (t[27] = 11.71, p < .0001), left and right middle frontal gyrus
(MFG) (t[27] = 6.01, p < .0001), and left and right pre-SMA
(t[27] = 12.55, p < .0001). Therefore, all the homologous pairs were
united, so including the right inferior frontal gyrus (IFG) there were
four ROIs.
Bilateral regions were used, although the Stroop-like task has ver-
bal elements, because we were interested in the effect of interfer-
ence. The neuroscientific models of the Stroop do not distinguish the
left and right regions (Banich, 2019; Botvinick et al., 2001), and the
data shows bilateral activation (Xu et al., 2016). Moreover, the ROIs
are not core linguistic regions, and were united only after confirmation
that they are highly correlated. Hence, separating the ROIs would
unnecessarily increase the number of comparisons, require correc-
tions, and thereby diminish the statistical power.
2.6 | Analyses
Behavioral and fMRI analyses were conducted across all stimuli. Prior
to statistical analyses, the trials were pooled together in order to opti-
mize generalization. That is, a dependent variable (i.e., accuracy, RT or
PSC) was averaged across all the congruent or incongruent trials,
regardless of stimulus type. Behavioral analyses were conducted after
discarding: five stimuli with average accuracy levels lower than 0.6
across all participants; the color-word type of four participants that
did not respond to that stimulus type; in the reaction time
(RT) analysis, trials with wrong, lack of response, or RT higher than
three SD from the mean RT of that subject. Trials excluded based on
behavioral criteria were not removed from the fMRI analysis.
The measures for behavioral analyses were accuracy level and
reaction time (RT). The measure for fMRI analyses was percent signal
change (PSC) from baseline, which was calculated for each ROI and
condition after the standard preprocessing and first-level analysis.
PSC is defined by the following formula: (condition baseline)/base-
line * 100. To determine which TRs should be used for the condition
and which for the baseline, we analyzed the actual hemodynamic
response in each ROI (see supplementary information S1). We found
that in all ROIs, the actual response shape resembled the canonical
hemodynamic response function. Additionally, we observed that the
highest hemodynamic response to the stimulus occurred during the
third and fourth TRs (5–8 s after stimulus onset), while the lowest
response was observed during the first and sixth TRs (0–2 and 11–
12 s after stimulus onset, respectively). Thus, we calculated the PSC

OREN ET AL.
by using the average signal during the third and fourth TRs as the con-
dition and the average signal during the first and sixth TRs as the
baseline.
The first behavioral and fMRI analyses examined the congruency
effect. They tested the hypothesis that the incongruent condition
compared to the congruent condition would have lower accuracy
levels, a longer RT and a higher PSC. Follow-up analyses established
the functional significance of congruency-related modulation in the
ROIs by calculating the relationship between PSC and RT. The second
set of behavioral and fMRI analyses provided an independent method
for inferring responsiveness to congruency via repetition suppression.
The hypothesis was that the non-first compared to the first condition,
would have higher accuracy level, a shorter RT and a lower PSC. Both
hypotheses were tested using directional paired samples t tests
(1-tailed). fMRI analysis was corrected for multiple comparisons across
all ROIs using the Holm-Bonferroni method (Holm, 1979), separately
for each hypothesis.
3 | RESULTS
3.1 | Behavioral results
Twenty-eight participants judged whether Stroop-like stimuli repre-
sented an animate or an inanimate object. Three types of stimuli were
used (Figure 1b): color-word (a word referring to a color written in
color ink), picture-word (a picture of an emotionally neutral object or
person with an emotionally neutral adjective superimposed on it), or
emotional picture-word (a picture of an emotional face with an emo-
tional word superimposed on it; see supplementary information S1 for
validation of the modified paradigm). To test the hypothesis of a
retained behavioral congruency effect despite the modification to the
Stroop task, we compared the congruent and incongruent conditions
across all three types, using directional paired samples t tests
(1-tailed). The difference between the conditions in accuracy was
non-significant (t[27] = 0.32, p = .37, 1-tailed; congruent mean SD:
0.91 (0.086), incongruent: 0.9 (0.085)). In line with our hypothesis, RT
was slower for incongruent (0.93 s [0.2]) than congruent (0.91 s
[0.17]) conditions (t(27) = 2.06, p = .024, 1-tailed, Cohen's d = 0.39),
confirming that conflicting information, unrelated to the animacy task,
was indeed processed and interfered with performance (see an addi-
tional behavioral analysis related to a congruency sequence effect in
supplementary information S1).
To test the hypothesis of behavioral repetition suppression, direc-
tional paired samples t tests (1-tailed) were carried out. As in the con-
gruency effect, repetition did not affect accuracy level (t[27] = 0.92,
p = .18, 1-tailed; first: 0.9 (0.094), non-first: 0.91 (0.091)). In line with
our hypothesis, repetition decreased RT, with faster responses to the
non-first presentation of stimuli (t(27) = 6.96, p < .001, 1-tailed,
Cohen's d = 1.15; first: 1.01 s (0.23), non-first: 0.85 s (0.15)). Thus,
performance in the animacy task was improved by repetition (see the
control analysis that ruled out a potential interaction between repeti-
tion and practice in supplementary information S1).
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4315
3.2 | fMRI results
Focusing on the frontoparietal regions that typically emerge in the
Stroop task (Xu et al., 2016), we tested the hypothesis that only the
IPS would exhibit the congruency effect when a task-irrelevant con-
flict referring to a different conceptual dimension than the task is pre-
sented. The frontoparietal regions were defined based on a meta-
analysis of Stroop studies by Xu et al. (2016). Four regions of interest
were included in our analyses, and overlapping and homologous ROIs
were combined, as explained in the Materials and Methods section:
bilateral IPS, bilateral MFG, bilateral pre-SMA and right IFG
(Figure 2a). In each ROI, we computed the PSC from baseline for both
congruent and incongruent conditions. As stated above, we com-
pared the conditions using directional paired samples t tests (1-tailed),
corrected for multiple comparisons using the Holm-Bonferroni
method (Holm, 1979). As expected, the bilateral IPS exhibited more
activation in the incongruent condition than the congruent condition
(Figure 2b; see statistics in Table 2). The bilateral MFG exhibited a
similar pattern, but it did not survive the correction for multiple com-
parisons. In the bilateral SMA and right IFG, the difference was non-
significant.
To clarify the functional significance of the congruency-related
modulation in the bilateral IPS, we examined the relationship between
activity in this region and behavior. We tested whether higher activa-
tions were related to longer RT in the incongruent versus congruent
conditions (Figure 3a). Since each stimulus appeared as incongruent
and as congruent, the difference (Δ) between the incongruent and
congruent conditions was calculated per stimulus in terms of RT and
PSC (Δ = incongruent congruent). To test whether trials with
behavioral congruency effect also showed activation congruency
effect in the IPS, we performed a linear repeated measures analysis
(where the participants were repeated) using a linear mixed model
with the maximum likelihood estimation method. The dependent vari-
able was ΔRT and the fixed effect was ΔPSC. Results showed a signif-
icant main effect for ΔPSC (F[11954] = 17.134, p < .001), indicating
that the greater the neural effect of incongruent stimuli in the IPS, the
greater their behavioral impact. This result was confirmed in a comple-
mentary analysis that compared the average ΔPSC of trials with posi-
tive ΔRT and negative ΔRT using paired-samples t-test. Averaging
across trials avoids the noise potentially associated with single-trial
analysis. Here too, there was a significant difference in the ΔPSC
(t[27] = 3.18, p = .002 1-tailed, Cohen's d = 0.6; Figure 3b). These
complementary results indicate there is an association between
slower responses and higher bilateral IPS activations for the incongru-
ent versus the congruent conditions, corroborating the role of the IPS
in processing task-irrelevant conflicting information. In other words,
longer responses to incongruent stimuli (behavioral congruency effect)
were associated with higher activations in the IPS.
Finally, repetition suppression was examined as a complemen-
tary method to identify brain regions that responded to task-
irrelevant conflicting information. In line with our hypothesis, repeti-
tion suppression was evident in the bilateral MFG, bilateral pre-SMA,
and right IFG but not in the bilateral IPS (Figure 2c, Table 2). Our
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4316 OREN ET AL.

F I G U R E 2 Regions of interest (ROI)

and activations. (a) Four ROIs were used
for the analysis based on Xu et al. (2016).
Each ROI is composed of several spheres
because overlapping and homologous
spheres were united and are presented in
the same color. (b) The difference (Δ) in
PSC between the incongruent and
congruent conditions (incongruent—
congruent). In the bilateral IPS activation
was significantly higher in the incongruent
than the congruent condition, manifested
by a positive Δ. (c) The difference in PSC
between the first and non-first conditions
(first—non-first). In all regions, except the
bilateral IPS, activation was significantly
lower for the non-first than the first
condition, manifested by a positive Δ. The
ROIs showed different sensitivity to
congruency and repetition in the parietal
and frontal regions. For each effect,
correction for multiple comparisons
across the four regions was carried out
using the Holm-Bonferroni method. Error
bars are standard errors. IFG, inferior
frontal gyrus; IPS, intraparietal sulcus; L,
left; MFG, middle frontal gyrus; R, right;
SMA, supplementary motor area; Δ,
difference (incongruent—congruent) or
(first—non-first).

TABLE 2 Congruency effect and repetition suppression in each region.

Congruency effect Repetition suppression

PSC PSC PSC non-

Name congruent incongruent Statistics PSC first first Statistics
Bilateral intraparietal 0.30 (0.15) 0.32 (0.15) t(27) = 2.4*, Cohen's 0.32 (0.16) 0.30 (0.14) t(27) = 1.37
sulcus d = 0.46
Bilateral middle frontal 0.24 (0.13) 0.26 (0.12) t(27) = 1.87 0.3 (0.14) 0.20 (0.11) t(27) = 6.9*, Cohen's
gyrus d = 1.3
Bilateral pre-SMA 0.31 (0.13) 0.32 (0.13) t(27) = 0.45 0.36 (0.14) 0.28 (0.12) t(27) = 6.05*, Cohen's
d = 1.14
Right inferior frontal 0.17 (0.09) 0.17 (0.11) t(27) = 0.4 0.20 (0.11) 0.14 (0.09) t(27) = 5.64*, Cohen's
gyrus/insula d = 1.06

Note: Percent signal change (PSC; means [SD]) and statistics for the congruency effect and repetition suppression.
*p < .05, 1-tailed, corrected.

results thus showed different response patterns in the parietal and repetition. Altogether, our findings provide converging evidence for
frontal regions to task-irrelevant information. The bilateral IPS the representation of task-irrelevant information in the parietal
responded to congruency, while the frontal ROIs responded to rather than the frontal regions.
 10970193, 2023, 11, Downloaded from https://onlinelibrary.wiley.com/doi/10.1002/hbm.26347 by Cochrane Mexico, Wiley Online Library on [06/07/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
OREN ET AL. 4317

F I G U R E 3 Relationship between IPS

activation and behavior. (a) The
calculation procedure. For each stimulus,
the difference (Δ) between incongruent
and congruent conditions
(Δ = incongruent—congruent) was
calculated in terms of the RT and PSC of
the IPS. The averaged ΔPSC of the IPS
was computed for each participant for
stimuli with positive ΔRT (incongruent
RT > congruent RT) and negative ΔRT
(incongruent RT < congruent RT). (b) A bar
graph presenting the result of the ΔPSC
comparison between the positive and
negative ΔRT. Across participants, a
higher ΔPSC was evident for stimuli with
positive ΔRT than negative ΔRT, implying
a functional significance of the bilateral
IPS activation in the congruency effect.
PSC, percent signal change; RT, reaction
time; Δ, difference (incongruent—
congruent); *p < .05.

4 | DISCUSSION
The present study asked whether information unrelated to the current
goal still affects us and, if so how. Previous studies attempted to
answer this question using the Stroop task. However, the non-
targeted attribute in a typical Stroop refers to the same conceptual
dimension as the targeted attribute and the task, making it
task-relevant (Figure 1a). To avoid such task-relevance we used a des-
ignated Stroop-like task consisting of conflicts that referred to a dif-
ferent conceptual dimension than the task (Figure 1b). Using this task,
a behavioral congruency effect still emerged, indicating that relevance
to current goals is not an essential requirement for a conflict to affect
human behavior. We further portrayed the neural correlates of a task-
irrelevant conflict using two complementary contrasts: congruency
and repetition. As hypothesized, a congruency effect was evident only
in the IPS and repetition suppression only in the frontal regions—the
bilateral MFG, bilateral pre-SMA, and right IFG—suggesting that parie-
tal, rather than frontal regions subserve the processing of task-
irrelevant conflicts.
4.1 | Typical behavioral congruency effect in our
Stroop-like task
At the behavioral level, our findings demonstrated the congruency
effect of longer RT in incongruent condition as compared with con-
gruent condition, in line with typical Stroop studies (MacLeod, 1991).
Regarding previous Stroop studies that separated the conflict from
the task, our results fit the findings of Zhao and Wang (2013) but con-
tradict those of Hatukai and Algom (2017). The later inconsistency
could be explained by the fundamental difference in the processing
depth needed to accomplish the task: semantic in-depth processing in
our study and perceptual low-level processing in the Hatukai and
Algom (2017) study. Hence, though it may be possible to avoid the
conflict when focusing on a low-level dimension, as Hatukai and
Algom (2017) interpreted their results, avoidance might become
impossible when the task requires extensive processing of the
conflict-irrelevant conceptual dimension, as in the present study.
Future studies are needed in order to directly test this explanation.
The diverse stimulus types used in the current study differed in
the proportion of animate or inanimate stimuli and difficulty level, as
stated above. Since every stimulus was presented both as congruent
and incongruent and the comparison was between these two condi-
tions, any potential bias from the proportion of animate or inanimate
stimuli or difficulty level was controlled for. This claim was supported
by empirical evidence presented in supplementary information S1.
We further found the congruency sequence effect, manifested as
longer RT in congruent trials that follow incongruent trials compared
with those following congruent trials (see supplementary information
S1), in accordance with previous studies (Egner, 2007, 2014). Finding
a congruency sequence effect, which measures the influence of a cer-
tain trial on the subsequent trial, in a design with different stimulus
types presented in a mixed order, expands our present understanding
of congruency sequence effects. Specifically, it indicates that a

4318
general sense of congruency underlies the effect, regardless of spe-
cific stimulus characteristics such as stimulus type. Overall, the repli-
cation of two hallmark behavioral effects of the Stroop task, namely
congruency and congruency sequence effects (Egner, 2007;
MacLeod, 1991; Schmidt, 2019), suggests our Stroop-like task has
much in common with the original Stroop task.
4.2 | Different activation patterns in parietal and
frontal regions
The parietal and frontal regions showed different response patterns
with regard to these contrasts (Figure 2b,c). The bilateral IPS main-
tained the typical congruency effect (Figure 2b); further, this effect in
the brain was related to the behavioral effect, so that increased bilat-
eral IPS activation for incongruent trials was seen in trials in which
incongruence led to larger RT delays (Figure 3). This finding is consis-
tent with the notion formulated by Banich et al. (2001) that activation
in these regions could constitute the neural correlate of the behavioral
elongation of RT for incongruent stimuli. In contrast, the frontal
regions showed repetition suppression (Figure 2c), implying that the
parameter that changed across repeated presentations, namely con-
gruency, was not represented in these regions. Notably, we did not
detect a repetition suppression effect in the IPS or a congruency
effect in the frontal regions at the conventional 1-tailed significance
threshold following correction for multiple comparisons. Altogether,
the results suggest the IPS is sensitive to task-irrelevant conflict asso-
ciated with a different conceptual dimension than the task, while fron-
tal regions are not. Therefore, taking into account the relationship
between the conflict and the task in terms of their conceptual dimen-
sion may be beneficial to future studies.
4.3 | Task-irrelevant information represented in
the IPS
The IPS maintained the congruency effect even when the conflict
task-irrelevant. The IPS is part of the parietal regions that impose top-
down and bottom-up attention (Shomstein, 2012). The left IPS has
been implicated in attention selection, allocating greater attentional
resources to incongruent than congruent stimuli, reflecting an
increased need for attentional control (Banich et al., 2001). The pre-
sent findings indicated a greater allocation of attentional resources in
this region, regardless of the task relevance of the conflict. This may
explain previous findings showing an even greater Stroop effect in this
region for multimodal as compared to unimodal congruency (Fitzhugh
et al., 2019).
Incongruent stimuli can be considered more salient than congru-
ent stimuli (Guido, 1998). From this perspective, greater recruitment
of the IPS for incongruent stimuli might reflect greater saliency, as
shown in non-Stroop visual tasks. For instance, the IPS responded to
the perceptual saliency of a task-irrelevant stimulus placed at a differ-
ent spatial location than the target (Geng & Mangun, 2009; Mazaheri
et al., 2011), a response correlated with the behavioral manifestation
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OREN ET AL.
(RT) (Geng & Mangun, 2009). Notably, IPS responsiveness to saliency
might manifest not only at a perceptual level but also at higher levels
of processing, as has previously been shown in the context of self-
association (Sui et al., 2015) or, as in the current study, in the context
of congruency.
To summarize, the Stroop literature and more general attention
research indicate modulation of IPS activity by task-irrelevant infor-
mation, in line with the current findings. Moreover, the fact that the
IPS was the key region that maintained the congruency effect when
the conflict was task-irrelevant, corresponds with the essentiality of
the posterior parietal regions to attentional control.
4.4 | When the task is irrelevant, the conflict is not
processed by the frontal regions
While task relevance in not essential for IPS recruitment, it seems to
be a prerequisite for the recruitment of the frontal regions tested in
our study. Though Stroop models focus mainly on the frontal regions
(Banich, 2019; Botvinick et al., 2001), in the current study, the repeti-
tion suppression effect showed limited involvement of Stroop-related
frontal regions for task-irrelevant conflict. This claim fits the current
understanding that the anterior prefrontal cortex receives diverse
inputs, integrates them, and controls and coordinates processing in
the pursuit of a behaviorally meaningful goal (Fuster et al., 2000;
Miller & Cohen, 2001; Ramnani & Owen, 2004).
Further, conflicting information that relates to a different concep-
tual dimension than the task may need fewer control mechanisms
than conflicting information that relates to the same conceptual
dimension as the task. Possibly, when it is task-irrelevant, the conflict
is processed by the IPS. Frontal regions are recruited only when addi-
tional control mechanisms are required due to task relevance.
4.5 | Repetition suppression in the service of
studying congruency effect
We used repetition suppression as a complementary means to infer
regional sensitivity to congruency. At least three factors can influence
repetition suppression. The first is the practice effect, which may be
disguised as repetition suppression, especially in a long task such as
ours. Yet, as presented in depth in supplementary information S1, in
the present study, a practice effect did not contribute to the repeti-
tion effect. The second is stimulus expectations, which can diminish
repetition suppression (Summerfield et al., 2008). Possibly, we are
prone to expect consistencies in the world, so incongruent stimuli
might violate our expectations. The third factor is saliency, so that
response to salient stimuli distinct from their background strongly
decreases in response to their repeated presentation (e.g., Ishai
et al., 2004). As mentioned before, incongruent stimuli may be
regarded as salient (Guido, 1998). The design of the current study
ruled out any potential distortion of the results due to the latter two
factors. The congruent and incongruent stimuli were equally distrib-
uted across the first and second positions, preventing any potential

OREN ET AL.
systematic bias in our results due to expectations or saliency. Hence,
we believe that repetition suppression does provide a valid measure
of neural tuning (Larsson & Smith, 2012) and faithfully reflects respon-
siveness to task-irrelevant conflicting information. Violating expecta-
tions and saliency probably played a significant role in the congruency
effect and the congruency sequence effect.
4.6 | Limitations
The study has several limitations. First, directional paired samples
t tests (1-tailed) were used in the statistical analyses. This choice was
backed up by firm and solid hypotheses regarding every contrast.
Moreover, in the fMRI analysis, the relatively liberal statistical thresh-
old was ameliorated by a strict correction for multiple comparisons.
Second, the study used a moderate sample of 28 participants. None-
theless, this sample size corresponds to the suggestions of Desmond
and Glover (2002), which are 24 participants, in order to maintain suf-
ficient statistical power. Third, as mentioned above, the design had
several potential sources for biases, including diverse stimulus types, a
different proportion of animate vs. inanimate stimuli, and practice.
However, the analyses described in depth in the supplementary ruled
out any bias due to these factors.
5 | C O N CL U S I O N
In summary, studies that examined the processing and response to
Stroop conflicts that refer to the same conceptual dimension as the
task implicated both frontal and parietal regions (Xu et al., 2016). Our
results suggest that the parietal regions, rather than the frontal
regions, play a key role in mediating the effect when conflicts arise
from a different conceptual dimension than the task. It seems that the
relevance of a conflict to a task, defined by its conceptual dimensions,
influences the recruitment of the frontal regions and not the IPS.
Future studies are needed to determine the role of control processes
in mediating this effect. The present study provides a new perspective
on how humans process and respond to information that is unrelated
to their current goal and advances understanding through the use of a
novel paradigm.
ACKNOWLEDGMENTS
The authors wish to thank Dr. Vered Kronfeld-Duenias for her helpful
comments regarding the manuscript, the Beilinson hospital manage-
ment for supporting our research, and the Beilinison hospital MRI
technicians for assisting throughout the MRI scans.
CONF LICT OF IN TE RE ST ST AT E MENT
The authors declare no competing financial interests.
DATA AVAI LAB ILITY S TATEMENT
Data subject to third party restrictions.
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4319
OR CID
Noga Oren https://orcid.org/0000-0002-0469-4180
Irit Shapira-Lichter https://orcid.org/0000-0002-4734-7744
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SUPPORTING INF ORMATION
Additional supporting information can be found online in the Support-
ing Information section at the end of this article.
How to cite this article: Oren, N., Abecasis, D., Inbar, E., Glik,
A., Steiner, I., & Shapira-Lichter, I. (2023). A new perspective
on the role of the frontoparietal regions in Stroop-like
conflicts. Human Brain Mapping, 44(11), 4310–4320. https://
doi.org/10.1002/hbm.26347