Neuropsychologia 169 (2022) 108204

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Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Nonverbal cognitive control training increases the efficiency of

frontal-subcortical collaboration for bilingual language control
Junjie Wu a, b, c, d, Man Zhang b, Qinpu Dang b, Qianwen Chang b, Qiming Yuan b, Zhaoqi Zhang b,
Guosheng Ding b, e, Chunming Lu b, e, Taomei Guo b, e, *
a
Key Research Base of Humanities and Social Sciences of the Ministry of Education, Academy of Psychology and Behavior, Tianjin Normal University, Tianjin 300387,
PR China
b
State Key Laboratory of Cognitive Neuroscience and Learning & IDG/McGovern Institute for Brain Research, Beijing Normal University, Beijing 100875, PR China
c
Faculty of Psychology, Tianjin Normal University, Tianjin 300387, PR China
d
Tianjin Social Science Laboratory of Students’ Mental Development and Learning, Tianjin 300387, PR China
e
Center for Collaboration and Innovation in Brain and Learning Sciences, Beijing Normal University, Beijing 100875, PR China

A R T I C L E I N F O A B S T R A C T

Keywords: Domain-general cognitive control is closely related to language control during bilingual language production.
Bilingualism Previous neural imaging studies have revealed a highly overlapped but rewired brain network for language
Brain network control and nonverbal cognitive control. In the present study, we examined this issue from a training perspective.
Domain-general cognitive control
Two groups of participants performed the language switching task at pre-and post-tests during functional
fMRI
Language control
magnetic resonance imaging (fMRI) scanning. After the pre-test, the experimental group received 8-day training
Training in a non-verbal switching task, while the control group performed an unrelated color judgement task. We found
that only the experimental group but not the control group showed decreased strength of connectivity from the
ventral lateral frontal cortex to the left caudate nucleus and from the medial surface of the frontal lobe to the left
thalamus. These results indicate an increased efficiency after nonverbal training for the frontal cortex to
implement domain-general suppression and monitoring in a domain-specific conflict context during bilingual
language and lexical selections. This study is the first to investigate the transfer effects of nonverbal cognitive
control on the brain network of bilingual language control and shed light on the mechanisms of how domain-
general cognitive control may underpin bilingual language control.

1. Introduction
Domain-general cognitive control is an ability for people to regulate
their behaviors in accordance with internal goals (Braver, 2012; Miller,
2000). For bilinguals, language control is recruited to inhibit the
non-target language for the successful selection and production of the
intended language (Christoffels et al., 2007; Guo et al., 2011; Meuter
and Allport, 1999). In the past decade, the relationship between bilin­
gual language control and domain-general cognitive control has been a
hot research topic (e.g., Blanco-Elorrieta et al., 2018; Blanco-Elorrieta
and Pylkkanen, 2016; Branzi et al., 2016a; Branzi et al., 2016b; De Bruin
et al., 2014; Timmer et al., 2018).
Behavioral studies have found that bilinguals with a higher level of
domain-general cognitive control showed better performance in lan­
guage control tasks (e.g., Declerck et al., 2017; Linck et al., 2012;
Woumans et al., 2015; but see Branzi et al., 2016a; Calabria et al., 2013;
Calabria et al., 2011). For example, Li et al. (2021) recruited participants
with various inhibitory control (IC) abilities to perform a language
switching task, which has been used to examine bilingual language
control (e.g., Christoffels et al., 2007; Costa and Santesteban, 2004; Guo
et al., 2011; Meuter and Allport, 1999). In this task, participants are
required to name pictures in the appropriate language according to some
sort of cue (e.g., a red frame around the picture, naming in the first
language [L1]; blue, naming in the second language [L2]). Unpredict­
able cues result in switch trials (where a different language is required to
name a picture from the preceding trial; i.e., L1→L2, L2→L1) and
non-switch trials (where the same language is required to name a picture
from the preceding trial; i.e., L1→L1, L2→L2). It has been found that
participants produce longer response times (RTs) and more errors on
switch trials than on non-switch trials, which is termed as switch cost.

* Corresponding author. State Key Laboratory of Cognitive Neuroscience and Learning, Beijing Normal University, Beijing, 100875, PR China.
E-mail address: guotm@bnu.edu.cn (T. Guo).

https://doi.org/10.1016/j.neuropsychologia.2022.108204
Received 2 October 2021; Received in revised form 21 February 2022; Accepted 3 March 2022
Available online 3 March 2022
0028-3932/© 2022 Elsevier Ltd. All rights reserved.
J. Wu et al. Neuropsychologia 169 (2022) 108204

Furthermore, larger switch costs have been observed in L1 than in L2 in improved neural efficiency or/and capacity (indexed by less connec­
unbalanced bilinguals, which is termed as asymmetric switch cost and tivity at comparable/better performance levels or/and more connec­
interpreted as evidence of inhibition on the dominant language in un­ tivity with better performance levels, respectively; see Ghazi Saidi et al.,
balanced bilinguals (e.g., Linck et al., 2012; Ma et al., 2016; Meuter and 2013; Giehl et al., 2020; Heinzel et al., 2014; Karim et al., 2017) in the
Allport, 1999; but see Christoffels et al., 2007; Costa et al., 2006). In the brain network for bilingual language control.
study of Li et al. (2021), larger asymmetric switch costs of RTs were
found in the bilinguals with higher IC ability, suggesting (at least partly) 2. Methods
shared mechanisms of bilingual language control and domain-general
cognitive control. 2.1. Participants
With regard to the neural activity, some imaging studies have
revealed a highly overlapped but rewired brain network for language Forty-seven Chinese-English bilinguals were recruited in the present
control and nonverbal cognitive control in bilinguals (Abutalebi & study. All participants were right-handed and had normal or corrected-
Green, 2008, 2016; Anderson et al., 2018; Dash and Kar, 2020; De Baene to-normal vision. None of them reported any neurological disorders.
et al., 2015; Tao et al., 2021; Weissberger et al., 2015; Wu et al., 2019). Participants were randomly assigned to the experimental group or the
For example, De Baene et al. (2015) found highly similar activation in control group. After excluding one participant because of excessive head
the prefrontal cortex, parietal cortex, and subcortical areas when par­ motion during fMRI scanning, twenty-four participants were left for the
ticipants performed language switching as well as nonverbal switching experimental group and 22 in the control groups. Due to the potential
tasks. In one of our recent studies (Wu et al., 2019), we further inves­ impact of demographic and background measures on behavioral per­
tigated the brain connectivity among the brain regions conjunctively formance and brain connectivity (Dash et al., 2022; DeLuca et al., 2019;
activated across these two tasks. We found a brain network with a Grundy, 2020; Wang et al., 2022), we collected or measured age, Raven
similar core-periphery structure and rewired connections from frontal to scores, College English Test Band-4 (CET-4, a compulsory test of college
subcortical regions for language control compared to that for nonverbal students in China, full score = 720), age of L2 acquisition, Length of L2
cognitive control. Overall, these studies have provided insight into the learning and using, self-rating scores (from 1 to 10, a higher score cor­
relationship between bilingual language control and domain-general responds to a higher level of proficiency) of listening, speaking, reading,
cognitive control from the perspective of neural activity. and writing in L1 or L2 for participants of both groups. As shown in
Recently, some studies have also investigated the impact of domain- Table 1, we further conducted two-sample t-tests on these measures and
general cognitive control training on language control (Liu et al., 2016; revealed no significant difference, ts (44) < 1.56, ps > 0.126. Neither did
Prior and Gollan, 2013; Wu et al., 2021). For example, Liu et al. (2016) Chi-square test in gender, χ2 = 2.14, p = 0.543. We performed 2 (Group:
conducted a study with a 6-day training of nonverbal cognitive control control group, experimental group) × 2 (Language: L1, L2) ANOVAs on
on bilingual participants with low IC ability (i.e., below the median). For self-rating scores of listening, speaking, reading, writing. Results showed
the behavioral performance of the language switching task, the asym­ significant main effects of Language, Fs(1, 44) > 62.94, ps < 0.001, ηp2
metric language switch costs at the pre-test were turned symmetric at = 0.589, indicating that participants were unbalanced bilinguals with
the post-test. For the event-related potentials (ERPs), the comparable higher proficiency in Chinese (overall = 8.3 ± 1.2, listening = 8.5 ± 1.2,
late positive components for switch trials of L2 and L1 at the pre-test speaking = 8.1 ± 1.4, reading = 8.5 ± 1.4, writing = 8.0 ± 1.6) than
became significantly larger for that of L2 than L1 at the post-test, that in English (overall = 5.3 ± 1.2, listening = 4.8 ± 1.7, speaking =
which were more like those of the bilinguals with high IC ability. 4.9 ± 1.5, reading = 6.1 ± 1.4, writing = 5.5 ± 1.4). Moreover, the
However, due to the low spatial resolution of ERP, this study failed to main effect of Group and the interaction between Language and Group
reveal the neural changes in specific brain regions. A subsequent neu­ were not significant, Fs < 1.207, ps > 0.278, ηp2 < 0.002, indicating that
roimaging study (Wu et al., 2021) examined the brain activation in bi­ these two groups were also well-matched in language dominance.
linguals who were performing the language switching task after an
8-day inhibitory control training. The study found improved activa­
tion in the left dorsal lateral prefrontal cortex (DLPFC) and no measur­
able changes in the left caudate nucleus (LCN) and anterior cingulate
cortex (ACC). Based on these results, it was proposed that inhibitory Table 1
control training may have reorganized cortical resources for bilingual Demographic and background measures for experimental and control groups.
language production by enhancing conflict resolution but not conflict Characteristics Experimental group Control group p
detection or lexical selection. However, this study investigated the (n = 24) (n = 22) values
activation in each brain region separately, leaving it an open question as Gender (male, female) 15, 9 9, 13 0.543
to how the collaboration among brain regions for language control Age (years) 22.6 (1.9) 22.3 (1.9) 0.542
adapts to the improvement of domain-general cognitive control. Raven scores 56 (3.8) 57 (3.4) 0.676
CET-4 scores 524 (51) 521 (54) 0.897
In the present study, we aimed to explore how nonverbal cognitive
Age of L2 acquisition 9.9 (3.0) 8.6 (2.5) 0.126
control training would alter the brain connectivity that underpins Length of L2 learning and 11.5 (3.3) 11.5 (1.7) 0.996
bilingual language control. Specifically, using a similar pre-post test using (years)
design and tasks of Wu et al. (2021), we recruited two groups of Overall Self-rating scores in 8.2 (1.3) 8.4 (1.2) 0.555
L1
Chinese-English bilinguals to perform a language-switching task during
Listening 8.3 (1.2) 8.7 (1.1) 0.222
functional magnetic resonance imaging (fMRI) scanning at both the Speaking 8.2 (1.4) 8.0 (1.5) 0.778
pre-and post-tests. After the pre-test, the experimental group received Reading 8.3 (1.6) 8.8 (1.2) 0.307
8-day nonverbal cognitive control training, whereas the control group Writing 7.9 (1.5) 8.0 (1.7) 0.794
performed an unrelated color judgement task during the same period. Overall Self-rating scores in 5.5 (1.3) 5.2 (1.2) 0.536
L2
Furthermore, based on our recent study (Wu et al., 2019), we adopted
Listening 5.0 (1.8) 4.7 (1.5) 0.655
extended unified structure modeling (euSEM) in a confirmatory way to Speaking 5.1 (1.6) 4.6 (1.4) 0.241
estimate the strength of the connections in the language control network Reading 6.1 (1.4) 6.2 (1.3) 0.816
for each participant at the pre-and post-tests. According to the (at least Writing 5.7 (1.3) 5.4 (1.6) 0.555
partly) shared mechanisms for bilingual language control and Note: L1, Chinese; L2, English. The difference in gender was examined by a Chi-
domain-general cognitive control documented in the literature, we ex­ square test, and the differences among other characteristics were examined by
pected that only participants in the experimental group would show two-sample t-tests. The standard deviations (SDs) are shown in parentheses.

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J. Wu et al.
2.2. Materials
We adopted the same materials as in Wu et al. (2021). Specifically,
forty-eight black-in-white line drawings were selected as formal stimuli
or fillers for the language switching task. Colored arrows and rectangles
were used in the training tasks respectively for the experimental and
control groups.
2.3. Procedure
This study was conducted with approval from the Institutional Re­
view Board of Beijing Normal University. All participants signed
informed consent before the experiment. The procedure and tasks in the
present study were the same as the protocol of Wu et al. (2021).
Therefore, we only provided a brief description in this section.
At the pre-and post-tests, participants performed the same language-
switching task while undergoing fMRI scanning. They were instructed to
name a picture in the appropriate language according to a cue indicated
by the frame color of the picture; i.e., red or blue. In each test session,
there were two runs of the language switching task. Each run included
82 trials with the first two being fillers. Each trial began with a fixation
cross of 300 ms and was followed by a blank screen of 200 ms. Then, a
picture with a colored frame was presented for 1000 ms, which was
replaced by a blank screen of 1, 2, 3, or 4 s (with equal probability).
Participants were required to name the picture in a soft voice to reduce
movement-related artifacts in the scanner. Due to loud noise during
scanning, we failed to record naming responses in the scanner. There­
fore, participants were invited to perform the same language-switching
task again outside the scanner to collect behavioral data (see also Kang
et al., 2017; Wu et al., 2019).
There was an 8-day interval between the pre-and post-tests. On each
consecutive day, the experimental group received nonverbal cognitive
control training for about 30 min, whereas the control group did a task
without any specific training for a similar amount of time. Specifically, a
nonverbal switching task was adopted as the training task to enhance the
domain-general cognitive control (precisely, nonverbal cognitive con­
trol) for participants in the experimental group (see Wu et al., 2021 for
details). In this task, participants were required to respond to an arrow,
pointing to the left or right according to a cue (i.e., the color of the
arrow). For example, if the arrow was red, participants were required to
press the button in the same direction as the arrow, which is termed
congruent rule or easy rule and thought to match the dominant language
in bilinguals. If the arrow was blue, participants had to press the button
in the opposite direction to the arrow, which is termed incongruent rule
or hard rule and thought to match the nondominant language in bi­
linguals (Wu et al., 2019). When the color of the arrow changed or
remained constant across two consecutive trials, participants had to
switch into another rule or repeat the same rule, respectively, which
intensively recruited domain-general cognitive control and resulted in
switch trials and non-switch trials. In contrast, the control group adop­
ted an unrelated color judgement task as a training task. Red and blue
rectangles without pointing directions were used as stimuli. Participants
were instructed to judge the color of the rectangle by pressing a left or
right button. The color judgement task was not expected to train any
components related to domain-general cognitive control or language
control for the control group. The general parameter settings were
identical across training tasks for the experimental and control groups.
2.4. Imaging data acquisition
The fMRI data were collected by a Siemens TRIO 3-T whole-body
MRI scanner, using a T2*-weighted gradient echo-planar images (EPI)
sequence with repeated time (TR) = 2000 ms, echo time (TE) = 20 ms,
flip angle (FA) = 90◦ , field-of-view (FOV) = 200 × 200 mm, matrix size
= 64 × 64, resolution within slices = 3.125 × 3.125 mm. All functional
images were interleaved from bottom to top, covering the entire brain
3
Neuropsychologia 169 (2022) 108204
with 33 slices, the thickness and gap of which were 4 mm and 0.8 mm,
respectively. After the fMRI scanning, T1-weighted anatomical images
were also obtained for coregistration with TR = 2530 ms, TE = 3.39 ms,
FA = 7◦ , FOV = 256 × 256 mm, matrix size = 256 × 256, resolution
within slices = 1.0 × 1.0 mm, slice thickness = 1.33 mm, number of
slices = 144.
2.5. Behavioral data analysis
For the training data, we first removed blocks with an accuracy
below 80% (1.48% and 0.86% of the data in experimental and control
groups, respectively). Trials with an incorrect response (3.84% and
2.57%, respectively for two groups), correct trials with a response
shorter than 200 ms or longer than 2000 ms (0.01% and 0.41%,
respectively), trials with a response time 2.5 SDs above or below the
mean (2.42% and 2.71%, respectively) were eliminated before the an­
alyses of RTs. With error rate (ER, 1 – accuracy) in each block as a
covariable, we separately fitted a linear mixed-effects model (LMEM) to
the RTs for the experimental and control groups. For the experimental
group, the Congruency (incongruent = 1, congruent = − 1), Trial type
(switch = 1, non-switch = − 1), and centered Training day and their
interactions were taken as fixed effects, with random intercepts for
participants, stimuli, and blocks. Model formula: RT ~ Congruency *
Trial type * Training day + ER + (1 | Subject) + (1 | Stimuli) + (1 |
Block). For the control group, the Color (blue = 1, red = − 1), Trial type
(switch = 1, non-switch = − 1), and centered Training day and their
interactions were taken as fixed effects, with random intercepts for
participants, stimuli, and blocks. Model formula: RT ~ Color * Trial type
* Training day + ER + (1 | Subject) + (1 | Stimuli) + (1 | Block). It
should be noted that we centered Training day to obtain an interpretable
baseline for each LMEM. The original Training day (1–8) would make
LMEMs take the “zeroth day” (which does not actually exist) as the
baseline, resulting in that the effects of regressors without Training day
(e.g., the main fixed effect of Trial type, the interaction between Trial
type and Congruency/Color) represent effects (e.g., switch cost, asym­
metry of switch cost) on the “zeroth day”. We centered Training day so
that the baseline of the LMEM was the grand average of all 8 training
days and so were the above-mentioned effects. We focused on the main
fixed effect of Trial type, the interaction between Trial type and Training
day, and the interaction between Trial type, Training day, and Con­
gruency/Color.
For the pre-and post-tests of the language switching task, data for one
participant in the control group was unavailable due to an equipment
malfunction. We removed incorrect trials and trials after language errors
or no responses (7.18%), absolute outliers (less than 200 ms and greater
than 2000 ms, 0.58%), and relative outliers (2.5 SDs above or below the
mean, 2.17%) for the analysis of RTs. Then, an LMEM was fitted to the
RTs with the ER in each run as a covariable and the Group (Experi­
mental = 1, Control = − 1), Test session (pre = 1, post = − 1), Language
(L1 = 1, L2 = − 1), Trial type (switch = 1, non-switch = − 1) and their
interactions as fixed effects. Random intercepts for participants and
pictures were included in the LMEM to increase the generalizability of
the study findings to a wider population of participants and language
materials (Linck and Cunnings, 2015). Model formula: RT ~ Group *
Test session * Language * Trial type + ER + (1 | Subject) + (1 | Picture).
We focused on the interaction among Group, Test session, and Trial type,
and the interaction among Group, Test session, Trial type, and Language
that respectively reflects the training effects on language switch cost and
asymmetry of switch cost.
The reason why we adopted LMEMs rather than traditional analyses
of variance (ANOVAs) was due to the missing data in the training ses­
sion. The traditional ANOVA deals with the missing data through list­
wise deletion. That means only participants with complete data sets are
kept for further analysis, which may lead to potential bias and reduce
statistical power. However, LMEM outperforms the ANOVA in robust­
ness against missing data, assuming that the data are missing at random
J. Wu et al. Neuropsychologia 169 (2022) 108204

(Cunnings, 2012; Linck and Cunnings, 2015). Furthermore, in LMEM, < 0.05, standardized root mean square residual (SRMR) < 0.05 (Beltz
the Training day is conveniently taken as a continuous variable, rather et al., 2013; Gates et al., 2011). The connectivity strength values
than a categorical variable in ANOVA which is followed by complex represent the intensity of collaboration between two brain regions,
post-hoc pairwise comparisons. In the present study, the analyses of which were extracted from the individual models at the pre-and post-­
LMEM were conducted by the lme4 package (Bates et al., 2014) and the tests for the following analyses.
significances (p-values) were determined based on Satterthwaite’s ap­ LMEMs were fitted to the strength values of all connections to
proximations using the lmerTest package (Kuznetsova et al., 2017). The evaluate the transfer effects of the nonverbal switching task on the brain
emmeans package was adopted for simple-effect analysis when necessary network for language control. Using the lme4 package (Bates et al.,
(Lenth et al., 2018). All these packages are implemented in R statistical 2014), we included random intercepts for subjects and fixed effects for
computing environment (R Development Core Team, 2010). the Test session (pre = 1, post = − 1), Group (experimental group = 1,
control group = − 1), and their interaction in the LMEM. Model formula:
2.6. Imaging data preprocessing Connectivity strength ~ Test session * Group + (1 | Subject). It is worth
noting that, unlike the analyses on behavioral data, Trial type and
We adopted Data Processing Assistant for Resting-state fMRI Language were not included in the model for the connectivity data.
(DPARSF 2.3; Yan and Zang, 2010), a MATLAB toolbox based on Sta­ Because the euSEM approach is based on continuous time series and
tistical Parametric Mapping (SPM version 8, http://www.fil.ion.ucl.ac. does not provide separate connectivity values for each experimental
uk/spm), to preprocess the imaging data. For an equilibrium state of condition (e.g., switch or non-switch) in the event-related fMRI data (see
the magnetic field, we discarded the first four images in each run, cor­ also Chen et al., 2021; Wu et al., 2019; Wise Younger et al., 2017).
responding to the two filler trials at the beginning. We performed slice Therefore, connectivity values, which collapsed languages and trial
timing correction for the remaining images using the middle slice as the types, were taken as the dependent variable, and only the Test session
reference slice. Next, all images were realigned to the first image for and Group were included as the independent variables. We focused on
head motion correction. One participant was excluded due to excessive the connections with significant interaction between Test session and
head motion (i.e., 3.0 mm and 3.0◦ ). Then, the anatomical image of each Group, which were determined based on Satterthwaite’s approxima­
participant was co-registered to the corresponding mean functional tions using the lmerTest package (Kuznetsova et al., 2017) and survived
image generated during the realignment. After that, we normalized all Bonferroni correction for 14 tests of all connections. Finally,
images in Montreal Neurological Institute (MNI) space and resampled simple-effect analyses were conducted on these connections using the
them to 3-mm cubic voxels. Finally, all images were smoothed with a emmeans package (Lenth et al., 2018).
Gaussian kernel of 6 mm full width at half-maximum. Additionally, we also conducted exploratory analyses of correlations,
especially between brain connectivity and behavioral performance, to
2.7. Brain network analysis approach the whole profile of the data in the present study (for details,
see the Supplementary Materials).
In the present study, a confirmatory euSEM was conducted to re-
estimate the connectivity strength based on the nodes and connectiv­ 3. Results
ity pattern revealed in Wu et al. (2019). As shown in Table 2, the
bilateral DLPFC, dACC/pre-SMA, left IPL, bilateral anterior insu­ 3.1. Behavioral results during training
la/inferior frontal gyrus (LAI/IFG and RAI/IFG), caudate nucleus (LCN),
bilateral thalami (LThal and RThal), and left cerebellum (LCereb) were Due to the high accuracy during training for the experimental group
selected as nodes and their time series were extracted for the fMRI data (95.95% ± 2.62%) and the control group (97.29% ± 2.15%), we only
of pre-and post-tests. focused on the results of RTs. For the experimental group, we found a
With the time series of these brain regions, we adopted the GIMME significant main fixed effect of Training day, β = − 15.35, t = − 49.03, p
(Group Iterative Multiple Model Estimation; Gates and Molenaar, 2012) < 0.001, indicating that the overall RTs decreased with training. There
toolkit to conduct a confirmatory model for each individual based on the was a significant main fixed effect of Trial type, β = 16.20, t = 52.13, p <
connectivity matrix of language control network from Wu et al. (2019). 0.001, indicating slower responses were made for switch trials (523 ±
The models were considered as good fit when at least two of the four 56 ms) than for non-switch trials (490 ± 47 ms). The interaction be­
criteria were met: comparative fit index (CFI) > 0.95, and nonnormed fit tween Trial type and Training day was marginally significant, β =
index (NNFI) > 0.95, root mean square error of approximation (RMSEA) − 0.61, t = − 1.96, p = 0.050, suggesting that switch costs tended to
decrease with training. The interaction between Trial type and Con­
Table 2 gruency was significant, β = − 5.86, t = − 18.85, p < 0.001, indicating
Information on the brain regions for language control network. that smaller switch costs were found in the incongruent trials than those
in the congruent trials (20 ± 21 ms and 45 ± 21 ms, respectively). The
Brain areas Abbreviations Peak coordinates
(MNI) interaction between Trial type, Congruency, and Training day was sig­
nificant, β = 0.74, t = 2.37, p = 0.018, demonstrating that the differ­
x y z
ences between switch costs in the incongruent trials and that in the
Left dorsal lateral prefrontal cortex LDLPFC − 39 45 18 congruent trials decreased with training (Fig. 1A).
Right dorsal lateral prefrontal cortex RDLPFC 36 39 30
For the control group, we also found a significant main fixed effect of
Dorsal anterior cingulate cortex/pre- dACC/pre- − 3 12 51
supplementary motor area SMA Training day, β = − 6.54, t = − 19.70, p < 0.001, indicating that the
Left inferior parietal lobe LIPL − 42 − 48 45 overall RTs decreased with training. There was a significant main fixed
Left anterior insula/inferior frontal gyrus LAI/IFG − 42 12 0 effect of Trial type, β = 1.67, t = 5.08, p < 0.001, indicating slower
Right anterior insula/inferior frontal RAI/IFG 39 21 0 responses were made for switch trials (441 ± 64 ms) than for non-switch
gyrus
Left caudate nucleus LCN − 12 9 6
trials (438 ± 65 ms). The interaction between Trial type and Color was
Left thalamus LThal − 9 − 6 3 marginally significant, β = 0.61, t = 1.84, p = 0.065. More importantly,
Right thalamus RThal 12 − 3 12 the interaction between Trial type and Training day and the three-way
Left cerebellum LCereb − 21 − 72 − 27 interaction between Trial type, Color, and Training day were not sig­
Note: MNI refers to the standard stereotaxic space from the Montreal Neuro­ nificant, βs = 0.44, ts < 1.34, ps > 0.182, indicating nonsignificant
logical Institute, which is the average of 152 normal magnetic resonance im­ training effects on the switch costs and the differences between switch
aging (MRI) scans. costs in red and blue trials (Fig. 1B).

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J. Wu et al. Neuropsychologia 169 (2022) 108204

Fig. 1. Switch costs across eight training days for the experimental group (A) and the control group (B).
Note: Con, congruent; InCon, incongruent.

3.2. Behavioral results for pre-and post-tests
Accuracy for the experimental group and control group at both pre-
and post-tests was also very high (96.06% ± 4.16% and 97.17% ±
2.65%, respectively). Therefore, only the results of the RTs were re­
ported. We found a significant main fixed effect of Trial type, β = 18.94,
t = 11.51, p < 0.001, indicating slower responses in switch trials (925 ±
164 ms) than in non-switch trials (887 ± 157 ms). There was no sig­
nificant interaction between Trial type and Test session, β = 1.53, t < 1.
The interaction among Group, Test session, and Trial type did not reach
significant, β = 0.98, t < 1. Neither did the interaction among Group,
Test session, Trial type, and Language, β = 0.48, t < 1. These results

Fig. 2. The training effect of the nonverbal switching task on the language control network.
Note: Each line between nodes represents the contemporaneous relation after controlling for the time-lagged relation and the modulating effect. Error bars stand for
one standard error. ***, p < 0.05 (Bonferroni corrected); *, p < 0.05 (uncorrected). The nodes labeled with numbers respectively are 1 dorsal anterior cingulate
cortex/pre-supplementary motor area (dACC/pre-SMA), 2 left dorsal lateral prefrontal cortex (LDLPFC), 3 right dorsal lateral prefrontal cortex (RDLPFC), 4 right
anterior insula/inferior frontal gyrus (RAI/IFG), 5 left anterior insula/inferior frontal gyrus (LAI/IFG), 6 left caudate nucleus (LCN), 7 right thalamus (RThal), 8 left
thalamus (LThal), 9 left inferior parietal lobe (LIPL), 10 left cerebellum (LCereb).

5
J. Wu et al.
suggested no significant training effects on the behavioral performance
of the language switching task indexed by the switch cost and asym­
metry of switch cost.
3.3. Results of brain connectivity at pre-and post-tests
The results of euSEM showed that the data for all participants were
fitted successfully, CFI = 0.986 ± 0.005 (range: 0.970–0.990), NNFI =
0.963 ± 0.011 (range: 0.910–0.980), SRMR = 0.031 ± 0.006 (range:
0.016–0.052), RMSEA = 0.050 ± 0.003 (range: 0.050–0.080).
As shown in Fig. 2, the interaction between Test session and Group
was only found in LAI/IFG→LCN, β = 0.04, t = 3.78, p = 0.007 (Bon­
ferroni corrected), and dACC/pre-SMA→LThal, β = 0.03, t = 2.11, p =
0.040 (uncorrected). Two separate simple-effect analyses on the
connection strength of LAI/IFG→LCN and dACC/pre-SMA→LThal
revealed significant reductions for the experimental group, ts > 3.32, ps
< 0.002, but not for the control group, ts < 1.
4. Discussion
The present study investigated the issue of how the brain network for
bilingual language control adapts to the short-term training of nonverbal
cognitive control. Specifically, the experimental group undertook an
eight-day nonverbal cognitive control training, whereas the control
group went through the same amount of training with an unrelated color
judgement task. We found that only the experimental group but not the
control group showed decreased strength of the connection from the
LAI/IFG to the LCN and the connection from the dACC/pre-SMA to the
LThal. We interpreted these findings as indicators of increased efficiency
after nonverbal training for the frontal cortex to implement domain-
general suppression and monitoring in a domain-specific conflict
context during bilingual language and lexical selections.
First of all, we found decreased strength of the connection from the
LAI/IFG to the LCN only in the experimental group. The LAI/IFG is a
crucial region of the salience network, whose major function is to detect
and filter behaviorally relevant stimuli (Dosenbach et al., 2006; Menon,
2015). A training study showed reduced activity in the LAI/IFG when
participants were performing an inhibitory control task (specifically,
Go/NoGo task) after 2 weeks of training with the same task (Chavan
et al., 2015; Wu et al., 2020). Furthermore, Chavan et al. (2017) found
that, compared with healthy controls, experts in fencing, who inten­
sively recruit inhibitory control, showed greater fractional anisotropy
(FA) of white matter tracts in the left IFG and left basal ganglia
(including the LCN), and a speeding-up in the inhibitory control task.
Additionally, shorter RTs in the inhibitory control task were shown in
the experts with higher FA values in the left basal ganglia. The basal
ganglia, especially the LCN, is considered as a role of language selection
(Abutalebi et al., 2008, 2013; Crinion et al., 2006). Lesions in this region
may lead to pathological and unintentional language switching (Abu­
talebi et al., 2000; Mariën et al., 2005). The neuroplasticity was also
shown in the basal ganglia for the bilinguals who underwent training of
language switching (Kang et al., 2017) or simultaneous interpretation
(Hervais-Adelman et al., 2015). Here, we found reduced connection
strength of the LAI/IFG → LCN in bilinguals who underwent the training
of cognitive control, suggesting improved efficiency of domain-general
salient cue detection and suppression serving for the domain-specific
language selection.
In addition, we also found a reduction in the strength of the
connection from the dACC/pre-SMA to the LThal only in the experi­
mental group. The dACC/pre-SMA is on the medial surface of the frontal
lobe and documented as a typical brain region for domain-general
cognitive control, especially for conflict monitoring (Botvinick et al.,
1999; Carter et al., 1998). In contrast, the LThal is a subcortical brain
region associated with lexical selection and semantic representation
during language production (Abutalebi, 2008; Abutalebi and Green,
2016; Wu et al., 2019). Studies on bilingualism have consistently
6
Neuropsychologia 169 (2022) 108204
revealed activation of the dACC/pre-SMA and LThal in the language
switching task (Abutalebi et al., 2008, 2013; Wang et al., 2007). Our
recent study has revealed the role of the dACC/pre-SMA as a hub both in
the language control network and nonverbal control network, and a
specific connection in the language control network from the
dACC/pre-SMA to the LThal (Wu et al., 2019). The present study
confirmed the connection from the dACC/pre-SMA to the LThal and
further revealed a training-induced reduction of strength in this
connection. This finding may suggest an improved neural efficiency of
domain-general monitoring participated in the selection of lexical and
semantic representations.
It is worth noting that we did not find significant improvement in
behavioral performances as measured by the comparison between
switch cost or its asymmetry at the post-test and that at the pre-test,
although the effectiveness of the training was demonstrated by the
reduction of switch cost and its asymmetry in the nonverbal cognitive
control training. One possible reason is that the relationship between
behavioral and neural gains is more complex than the simple linear (e.g.,
Hu et al., 2017; McLaughlin et al., 2004). For example, McLaughlin et al.
(2004) revealed neural changes (as measured by event-related poten­
tials, specifically the N400 amplitude) without any significant behav­
ioral improvement in participants who learned new words of a second
language for just 14 h. In other words, the training-induced improve­
ment in neural activity might be a prelude to the improvement in
behavioral performance. Additionally, the length of 8-day training in the
nonverbal switching task in the present study might not be sufficient to
induce transfer effects on language control in behavioral performance.
Future studies may adopt more intensive training for a longer term.
Taken together, the present study revealed reduced strength of
connections from the ventral or medial frontal lobe to the subcortical
areas for bilinguals who underwent nonverbal cognitive control
training, indicating possible improved neural efficiency for bilinguals to
employ domain-general cognitive control in the context of bilingual
language production.
5. Caveats and limitations
Although our results offer an insight for the transfer effects of
nonverbal cognitive control on the brain network of bilingual language
control, it should be noted that the strength of the connection of dACC/
pre-SMA to the LThal differed between the two groups at the pre-test.
One possible reason is that the two groups actually differed in some­
what aspects that could not be detected by our measures taken to collect
their demographic and background information. If this were true, we
indeed would not know what these aspect(s) of properties were. How­
ever, the training-induced reduction for each group could be treated as
the reduced connectivity strength at the post-test compared to the
connectivity strength at the pre-test as the baseline. From this perspec­
tive, it is appropriate to claim that, compared to its own baseline, only
the experimental group but not the control group showed a reduction in
the connectivity strength of dACC/pre-SMA→LThal. Anyway, further
confirmations with more covariables controlled or matched are
appealed for in future studies.
6. Conclusion
In the present study, we adopted a pre-post-training design to
investigate the transfer effects of nonverbal cognitive control on the
brain network for bilingual language control. Only in the experimental
group (but not in the control group), we found improved behavioral
performance during the training of nonverbal cognitive control and
reduced strength of connection from the frontal lobe to the subcortical
areas (specifically, LAI/IFG→LCN, dACC/pre-SMA→LThal) at the post-
test of language control. These results indicate that nonverbal cogni­
tive control training enhanced the neural efficiency for the frontal-
subcortical collaboration to implement domain-general suppression
J. Wu et al.
and monitoring in resolving the dilemma of selecting languages and
lexical representations. The present study provides the first evidence for
the transfer effects of nonverbal cognitive control on bilingual language
control from the perspective of the brain network.
Funding
This work was supported by the National Natural Science Foundation
of China (32100854 to J.W. and 31871097 to T.G.), the Interdiscipline
Research Funds of Beijing Normal University, and the Fundamental
Research Funds for the Central Universities (2017XTCX04).
Declaration of competing interest
None.
CRediT authorship contribution statement
Junjie Wu: Conceptualization, Data curation, Formal analysis,
Investigation, Methodology, Software, Visualization, Writing – original
draft, Writing – review & editing. Man Zhang: Conceptualization, Data
curation, Investigation, Visualization. Qinpu Dang: Data curation,
Visualization. Qianwen Chang: Investigation, Visualization. Qiming
Yuan: Investigation, Methodology, Software. Zhaoqi Zhang: Investi­
gation, Methodology, Software. Guosheng Ding: Conceptualization.
Chunming Lu: Resources. Taomei Guo: Conceptualization, Funding
acquisition, Supervision, Writing – review & editing.
Acknowledgments
The authors would like to thank Qiping Wang, Alex Titus, Jack Bode,
and Dr. Simone Conceição for proofreading.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.neuropsychologia.2022.108204.
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