Brain & Language 211 (2020) 104879

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Brain and Language

journal homepage: www.elsevier.com/locate/b&l

Thinking outside the box: The brain-bilingualism relationship in the light of

early neurobiological variability
Nicola Del Maschio a, Simone Sulpizio a, b, Jubin Abutalebi a, c, *
a
Centre for Neurolinguistics and Psycholinguistics (CNPL), Faculty of Psychology, University Vita-Salute San Raffaele, Milano, Italy
b
Department of Psychology, University of Milano-Bicocca, Milano, Italy
c
The Arctic University of Norway, Tromsø, Norway

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

Keywords: Bilingualism represents a distinctive way to investigate the interplay between brain and behaviour, and an
Bilingualism elegant model to study the role of environmental factors in shaping this relationship. Past neuroimaging research
Brain-behavior relationship has mainly focused on how bilingualism influences brain structure, and how eventually the brain accommodates
Structure-leading-function
a second language. In this paper, we discuss a more recent contribution to the field which views bilingualism as
Brain anatomy
Sulcal pattern
lens to understand brain-behaviour mappings from a different perspective. It has been shown, in contexts not
Language related to bilingualism, that cognitive performance across several domains can be predicted by neuroanatomical
Structural MRI variants determined prenatally and largely impervious to postnatal changes. Here, we discuss novel findings
indicating that bilingualism modulates the predictive role of these variants on domain-specific cognition. The
repercussions of these findings are potentially far-reaching on multiple levels, and highlight the need to shape
more complex questions for progress in cognitive neuroscience approaches to bilingualism.

1. Introduction relationship. The continuation of research must therefore be ensured –

It is well established that the ability to acquire and use more than one
language acts as a catalyst for change in the human brain. Bilingualism-
related adaptations have been found, for example, in auditory cortical
structures (e.g., Heschl’s gyrus), portions of the language network (e.g.,
Broca’s area), and executive systems which support language control
and code-switching (e.g., anterior cingulate cortex, dorsal striatum) (see
Abutalebi & Green, 2016; Krizman & Marian, 2015). However, despite
the considerable amount of studies accumulated in recent years, there is
limited agreement on the specific brain areas that are structurally
different in bilinguals and monolinguals, and difficulties persist in
reconciling neuroanatomical findings with (the lack of) behavioral ef­
fects on cognitive task performance (e.g., Leivada et al., 2020). This is
arguably due to a combination of factors which pertain, on the one hand,
to bilingualism as an incremental, multifactorial and non-uniform object
of inquiry, and on the other hand, to the inherent limitations of the
correlative methods used to study it, which typically lead to under­
determined results.
Bilingualism represents, however, a distinctive way to investigate
how brain and behaviour affect one another, and an elegant model to
study the role of environmental factors in modulating this intriguing
we argue – in order to shed light on fundamental questions at the core of
cognitive neuroscience. For this to happen, though, it is critical to depart
from oversimplified main-effect-type questions about whether bi­
linguals outperform monolinguals on a certain task, or show structural
differences in a given region of the brain. It is increasingly recognized,
indeed, that concrete progress in the field requires appreciating the
multidirectional interactions between brain, behaviour and environ­
ment over time, as well as the range of variation which inherently
characterizes second language (L2) learning and bilingualism (see
Birdsong, 2018; Hernandez et al., 2019; Li, 2015). Past anatomical MRI
research has focused on the brain’s capacity to alter its structure in order
to successfully engage with a bilingual environment. The aim of this
article is to present a parallel, more recent contribution to the field
which views bilingualism as a lens to understand brain-behaviour
mappings from a different angle. In particular, rather than exploring
the consequences of bilingualism for mind and brain, attempting to link
a specific environmental experience to neurocognitive changes, the
presently-illustrated line of research aims to investigate whether bilin­
gualism interacts with longitudinally stable markers of early brain
development in shaping cognition. Otherwise stated, the goal is to test
whether a specific environmental experience such as bilingualism

* Corresponding author at: Vita-Salute San Raffaele University, Via Olgettina, 58, 20132 Milan, Italy.
E-mail address: abutalebi.jubin@hsr.it (J. Abutalebi).

https://doi.org/10.1016/j.bandl.2020.104879
Received 14 May 2020; Received in revised form 1 August 2020; Accepted 6 October 2020
Available online 17 October 2020
0093-934X/© 2020 Elsevier Inc. All rights reserved.
N. Del Maschio et al.
affects cognitive functions in different ways depending on the neuro­
anatomical substrata that predate this experience. This shift of perspec­
tive stems from a growing body of evidence which shows that cognitive
performance across several higher-order functions is partly constrained
by individual differences in regional patterns of cortical morphology
such as gyro-sulcal patterns (e.g., Borst et al., 2016; Cachia et al., 2018;
Fornito et al., 2004). Differently from typical markers of neuroplasticity,
these patterns are determined in utero and largely impervious to post­
natal changes (Cachia et al., 2016; Tissier et al., 2018), thereby
providing the opportunity to track the directionality of brain-behaviour
interactions. In this article, we will present data which indicate that
bilingual language background interferes with the predictive role of
these markers on cognitive functioning. The general finding is that living
with two languages modulates the functional consequences of local
neuroanatomical variants, with bilinguals’ performance benefiting from
specific patterns of cortical morphology that are detrimental to mono­
linguals and vice-versa. In what follows, we will first synthesize previous
findings on anatomic trait markers of cognition; then, we will discuss the
interaction between some of these markers and bilingual experience in
shaping cognitive abilities, with special reference to the domain of ex­
ecutive control functions. We will conclude by emphasizing the poten­
tial repercussions of these novel findings for bilingualism research.
2. Neuroanatomic trait markers of cognition
A wealth of information about individuals is encoded in their brain
anatomy and can be studied in vivo using structural MRI. The brain’s
architecture results from the dynamic interplay between genes and
environment, and both of these factors contribute to neurostructural
variations across individuals (see Gu & Kanai, 2014). Brain imaging
studies of twins and genome-wide association studies (GWAS) of imag­
ing phenotypes from unrelated subjects point to a moderate-to-strong
heritability of brain structure across gray and white matter tissue
(Thompson, Martin, & Wright, 2010; Strike et al., 2015). Recently,
large-scale multivariate analyses have linked the heritability of brain
structure with the heritability of general cognitive traits such as
response speed, fluid intelligence and numerical reasoning in healthy
populations (e.g., Davies et al., 2018; Zhao et al., 2019). These findings
suggest that the heritability of a cognitive function may partly depend
on the extent to which the neural substrata of that function are influ­
enced by genetic factors. The same findings also hint at the possibility
that individual differences in a cognitive function may be partly
explained by genetically determined variations in the function’s sub­
strata. Importantly, heritability estimates in the studies aforementioned
also show that a proportion of the observed variance in brain and
cognitive phenotypes is not due to genes, indicating that gene-by-
environment or gene-by-age interactions contribute to individual dif­
ferences in these phenotypes.
In addition to twin and GWAS imaging studies, a distinct line of
research has examined the relationship between local brain morphology
and domain-specific cognition. In particular, this research aimed to test
whether cognitive differences among individuals can be partly attrib­
uted to the naturally occurring variation in gyro-sulcal patterns of spe­
cific cortical areas (e.g., Borst et al., 2016; Cachia et al., 2014, 2018;
Fornito et al., 2004). Unlike quantitative indices of cortex structure,
gyro-sulcal patterns are determined prenatally and largely unaffected by
brain maturation and postnatal experiential factors (Cachia et al., 2016;
Sun et al., 2012; Tissier et al., 2018). Measures of sulcal anatomy such as
sulcal depth or length can obviously change with age and experience (e.
g. Kochunov et al., 2005; Li et al., 2010), but the inter-hemispheric
distribution and continuity/interruption of cortical folds have been
proven to be longitudinally stable (Cachia et al., 2016; Tissier et al.,
2018). Gyro-sulcal patterns also vary considerably in human pop­
ulations. Whereas the earlier-forming sulci are robustly conserved across
individuals, thereby suggesting a deterministic imprinting (e.g. Le Guen
et al., 2018), a greater variability in number, positioning, and
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Brain and Language 211 (2020) 104879
relationship to functional areas characterizes secondary and tertiary
sulci (Fischl et al., 2008; Hasnain, Fox, & Woldorff, 2006), which
develop sequentially after the 30th week of gestation. It has been pro­
posed, on these grounds, that biomechanical, environmental or sto­
chastic factors must play a role in the patterning of secondary and
tertiary folding at later stages of corticogenesis (see Borrell, 2018;
Quezada, Castillo-Melendez, Walker, & Tolcos, 2018; White, 2019).
Conceivably emerging from the co-occurrent action of genetic and more
random variables, gyro-sulcal patterns have been identified as
early-determined factors which can contribute to shape subsequent
cognitive abilities. At the structural level, the extant evidence concerns
the cognitive effects of sulcal pattern variability in three macro-areas: a)
Anterior cingulate cortex (ACC); b) Superior temporal gyrus (STG); c)
Occipito-temporal cortex (OTC). In the rest of this section, we will
succinctly review evidence for each macro-area, before moving on to
discuss the modulatory effect of bilingualism on neuroanatomic trait
makers of cognition.
2.1. Anterior cingulate cortex (ACC)
The ACC – a key component of the executive network (Botvinick,
Braver, Barch, Carter, & Cohen, 2001; Petersen & Posner, 2012) – is
located bilaterally in the medial frontal lobe and shows high variability
in gyro-sulcal architecture (Vogt & Palomero-Gallagher, 2012). Rele­
vant for the present work, the cingulate sulcus (CS) may be flanked by an
additional secondary sulcus, known as paracingulate sulcus (PCS),
which runs dorsal and parallel to the CS and separates the medial di­
vision of the superior frontal gyrus from the paracingulate gyrus. The
PCS is present in 30–60% of the normal population, with greater prev­
alence in the left hemisphere (Paus et al., 1996; Wei et al., 2017).
Variability in the ACC sulcal pattern has been found to be associated
with individual differences in cognitive functions tapping executive
control (EC), from working memory (Fornito et al., 2004) to inhibition
(Borst et al., 2014; Huster et al., 2009). Specifically, both children and
adults with an asymmetrical ACC sulcal pattern (e.g., a PCS present in
the left hemisphere only) relative to those with a symmetrical sulcal
pattern (e.g., a PCS absent in both hemispheres) have been reported to
exhibit higher EC efficiency on tasks requiring conflict monitoring and
interference suppression (e.g., Cachia et al., 2014; Tissier et al., 2018).
For example, by using structural MRI and Stroop interference scores in a
longitudinal design, Borst et al. (2014) found that children with an
asymmetrical ACC sulcal pattern had better EC efficiency both at age 5
and age 9 than children with a symmetrical pattern. A number of non-
mutually exclusive mechanistic hypotheses have been proposed to ac­
count for this advantage, including the effects of sulcal pattern on
functional brain activity (e.g., Amiez et al., 2013; Fedeli, Del Maschio,
Caprioglio, Sulpizio, & Abutalebi, 2020), and the association between
differences in sulcal pattern and distinct profiles of underlying white
matter architecture (e.g., Borst et al., 2014; Tissier et al., 2018; Van
Essen, 1997).
2.2. Superior temporal gyrus (STG)
Morphologic STG variability has been extensively described across
hemispheres and brains (e.g., Galaburda, Corsiglia, Rosen, & Sherman,
1987; Ochiai et al., 2004). A prime example is represented by Heschl’s
gyrus (HG), which lies transversely across the superior temporal plane
and hosts primary auditory areas. The HG is most frequently identified
as a single gyrus, but HG duplications/multiplications have also been
reported in either one or both hemispheres (e.g., Marie, Maingault,
Crivello, Mazoyer, & Tzourio-Mazoyer, 2016). Distinct lines of evidence
point to a specific role of HG duplications/multiplications associated
with advanced auditory, musical, and phonetic skills (e.g., Benner et al.,
2017; Golestani, Price, & Scott, 2011; Warrier et al., 2009). For example,
a group of studies by Golestani and colleagues explored the relationship
between variability in the gross morphology of the auditory cortex and
N. Del Maschio et al.
phonetic learning in phoneticians and non-experts (e.g., Golestani &
Pallier, 2007; Golestani, Molko, Dehaene, LeBihan, & Pallier, 2007;
Golestani et al., 2011). Among non-experts, faster phonetic learners had
a larger left HG than slower phonetic learners, and more frequently
exhibited multiple or split left transverse gyri (Golestani & Pallier,
2007). When comparing phoneticians with non-experts, the former
more frequently displayed multiple or split left transverse gyri, and the
degree of left transverse gyrus splitting within the phoneticians group
was not predicted by the amount of phonetic training (Golestani et al.,
2011). When interpreting this latter finding, the authors suggested that
neuroanatomical predispositions might have biased career choices.
Studies on musical aptitude and professional musicianship provide
complementary evidence that HG morphotypes are more likely to have
their origin in predisposition and early neurodevelopmental processes
rather than training-related plasticity (e.g., Benner et al., 2017; Zatorre,
2013).
2.3. Occipito-temporal cortex (OTC)
The left ventral OTC – a critical part of the reading circuitry (Lochy
et al., 2018; Price, 2012) – shows extensive anatomical variability in
normal populations. The left lateral occipito-temporal sulcus (OTS), in
particular, which runs horizontally on the inferior surface of the tem­
poral lobe, may appear as continuous or interrupted (e.g., Alves, Ribas,
Parraga, & de Oliveira, 2012). The posterior part of the left lateral OTS
hosts the “visual word form area” (VWFA) (Cohen et al., 2000), a region
which displays selective activation to written forms of words modulated
by reading experience (e.g., Dehaene & Cohen, 2011; Dehaene, Cohen,
Morais, & Kolinsky, 2015). Morphologic variability of the left OTS may
possibly affect the organization of the VWFA, with repercussions on
reading skills. This hypothesis has been recently tested on healthy adults
by Cachia et al. (2018), who also explored whether the effect of sulcal
variability might be modulated by the age of reading acquisition. Results
indicated that the variability of the left posterior OTS predicted reading
fluency in literate adults (i.e., participants who learned to read during
childhood). Furthermore, reading fluency was significantly better in
literate adults with a posterior interruption in the left OTS compared to
literate adults with a continuous posterior left OTS, while no significant
difference was detected between ex-illiterates (i.e., adult participants
who learned to read during adulthood) with or without interruption in
the posterior part of the left OTS. Note that the distribution of the sulcal
interruption of the left posterior OTS did not differ between groups. In
mechanistic terms, the relationship between left OTS variability and
reading fluency has been proposed to be mediated by differences in
structural connectivity between VWFA and the anterior and medial
temporal lobe (Cachia et al., 2018). Crucially, these findings also suggest
that morphologic variants per se do not necessarily contribute to shape
cognitive differences. Indeed, an effect of OTS variability was detected
only when coupled with a relevant environmental input within a specific
time-window.
Overall, albeit still scattered, the experimental evidence reviewed in
this section is interesting in many respects. Previous understanding of
brain-behaviour relationships within a structure-leading-function
perspective was mainly related to neuropathological conditions. It was
well established, for instance, that pathological alterations in brain size
or folding lead to gross defects in cognitive development and perfor­
mance (for review: Fernández, Llinares-Benadero, & Borrell, 2016). The
findings presented herein suggest that cognitive functions can also be
constrained by neuroanatomical variants in non-clinical populations, as
exemplified by the effect of regional gyro-sulcal patterns on domain-
specific cognition. On the other hand, it is worth remarking that
neuroanatomic trait markers of cognition should not be interpreted
within a strong deterministic framework. Normal morphologic variants
may predict, at least in part, cognitive development and performance,
but multiple other variables are expected to intersect with patterns of
brain structure in shaping cognitive and behavioral outcomes. We tested
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Brain and Language 211 (2020) 104879
this hypothesis by investigating whether the gyro-sulcal architecture of
the ACC predicts EC performance differently when individuals have
variable linguistic experience.
3. Bilingualism modulates neuroanatomic trait markers of
cognition
To investigate a potential modulatory role of bilingualism on brain-
behaviour relationships, we collected structural MRI data from bilingual
and monolingual groups, and compared their performance on a classic
measure of conflict monitoring (i.e., the Flanker task). In bilinguals,
activation of the ACC (and, in particular, of the dACC/pre-SMA com­
plex) is typically observed during resolution of conflict in both verbal
and non-verbal contexts (for reviews: Abutalebi & Green, 2016; Cala­
bria, Costa, Green, & Abutalebi, 2018; Sulpizio, Del Maschio, Fedeli, &
Abutalebi, 2020). In a first exploratory study on young adults, Cachia
et al. (2017) found differential effects of ACC sulcal pattern on bilinguals
and monolinguals’ EC performance, suggesting that bilingual experience
modulates the effects of local morphologic variants on cognitive effi­
ciency. In particular, a reversed pattern of structure-function relation­
ship between ACC sulcation and conflict effect induced by the Flanker
task was reported for bilingual and monolingual groups. Asymmetrical
versus symmetrical sulcal patterns were associated with a performance
advantage in monolinguals and a performance detriment to bilinguals
and vice versa. This preliminary result was replicated by Del Maschio,
Sulpizio, Fedeli, et al. (2019) with a larger sample varying on age,
gender and ethnicity. In this study, a paracingulate sulcus (PCS) on the
medial surface of the ACC was detected in ~50% of participants, with a
proportionally higher occurrence in the left hemisphere. The inter-
hemispheric distribution of the ACC sulcal pattern was not affected by
gender or ethnicity. We found that EC deterioration with increasing age
was modulated by language group, with monolinguals outperforming
bilinguals at a young age, and bilinguals being more resistant to decline
at an old age. Independent of language group membership, the conflict
resolution capacity of participants across age was modulated by ACC
sulcal pattern, with a performance advantage associated with an
asymmetrical sulcal pattern (a PCS present in the left hemisphere only),
and a detriment to performance associated with a symmetrical pattern (a
PCS absent in both hemispheres). Independent of age, asymmetrical and
symmetrical sulcal patterns were associated – respectively – with higher
and poorer EC efficiency in monolinguals, whereas the reverse was true
for bilinguals. The reversed pattern of structure-function relationship
between ACC sulcal pattern and EC efficiency for bilinguals and
monolinguals is represented in Fig. 1. Fig. 2 represents, for illustrative
purposes, a 3D rendering of a cingulate sulcus surmounted by a para­
cingulate sulcus on the medial surface of the ACC.
The key finding of the two studies aforementioned is represented by
the reversed pattern of structure-function relationship between ACC
sulcal pattern and EC efficiency for bilinguals and monolinguals. This
finding suggests that the long-term exposure to an L2 may overcome the
known correlation between sulcal asymmetry and better conflict reso­
lution capacity, pointing to a compensatory effect of bilingualism over
early and invariant neurodevelopmental factors. As a tentative mecha­
nistic explanation, one may speculate that bilinguals would rely more
extensively on a bilateral recruitment of the EC network relative to
monolinguals, with a greater contribution of inter-hemispheric activity
to executive functioning, which could account for the positive associa­
tion between ACC symmetrical patterns and EC efficiency in bilinguals.
Support to this interpretation comes from functional MRI studies
reporting a bilateral recruitment of the ACC in bilinguals engaged in
non-verbal EC tasks (e.g., Abutalebi et al., 2012; De Baene, Duyck, Brass,
& Carreiras, 2015) and diffusion tensor imaging (DTI) studies showing
enhanced structural connectivity in a bilateral cluster of cingulo-frontal
regions as a result of L2 training (e.g., Schlegel, Rudelson, & Tse, 2012)
or associated with L2 usage and immersion (Del Maschio, Sulpizio, Toti,
et al., 2019; Pliatsikas, Moschopoulou, & Saddy, 2015). Further
N. Del Maschio et al. Brain and Language 211 (2020) 104879

Fig. 1. Sulcal asymmetry of the anterior cingulate

cortex (ACC) and cognitive control efficiency in
bilinguals and monolinguals. Top: ACC sulcal
patterns. The cingulate sulci (aquamarine) and
paracingulate sulci (red) are represented on the
medial cortical surface. L = Left Hemisphere; R =
Right Hemisphere. Middle: Flanker task condi­
tions and procedure. Bottom: Conflict effect
scores (differential RTs: Incongruent minus
Congruent trials) in bilinguals and monolinguals
with different ACC morphology (adapted from
Cachia et al., 2017). (For interpretation of the
references to colour in this figure legend, the
reader is referred to the web version of this
article.)

4
N. Del Maschio et al.
Fig. 2. 3D rendering of a right hemisphere’s brain showing a cingulate sulcus
(red) and a paracingulate sulcus (blue) on the medial surface of the anterior
cingulate cortex (ACC). (For interpretation of the references to colour in this
figure legend, the reader is referred to the web version of this article.)
experimental indication is needed to substantiate this interpretation, but
the repercussions of the findings presented in this section are potentially
far-reaching.
In general, these findings contribute to an increasing body of work in
the cognitive neuroscience of bilingualism which underlines the
importance of appreciating both the complexity of neural development
and the interactions that take place between patterns of brain structure
and patterns of behaviour at different time-scales. Preconditions related
to normal variation in brain structure and morphology are not usually
numbered among the potential sources of inter-individual variability in
bilingual language processing. Recently, a neurogenetic approach to
individual differences in the context of L2 has begun to be explored,
though with a specific focus on the interplay between the dopaminergic
system, neural activity in the fronto-striatal pathway, and task-related
performance (Vaughn & Hernandez, 2018; Wong, Morgan-Short, Ett­
linger, & Zheng, 2012). Vaughn et al. (2016) reported that activity in the
inferior frontal gyrus during picture naming in L2, as well as activity in
the ACC during a shape-colour switching task, could be predicted by
variation in a gene encoding the D2 subtype of the dopamine receptor (i.
e., DRD2). Based on these findings, the authors concluded that poly­
morphisms of dopamine-related genes such as DRD2 could mediate, at
least in part, aspects of bilingual language processing associated with
fronto-striatal functions. We believe that the study of bilingual language
processing would largely benefit from an integrated approach contem­
plating both neurogenetics and the novel findings presented in this
section.
Of equal interest, neuroanatomical preconditions are not usually
numbered even as potential predictors of cognitive differences between
bilinguals and monolinguals when making group inferences. Shaping a
more complex scenario – we argue – could prove particularly useful also
in light of the controversies in the behavioral literature concerning a
putative cognitive advantage of bilinguals relative to monolinguals (for
review: Antoniou, 2019). A multimodal approach, contemplating the
interactive effects of early neuroanatomical constraints and bilingual
experience on task performance, could be profitably used to frame data
inconsistencies within a more comprehensive perspective (for a similar
proposal, see also, Sulpizio & Abutalebi, 2019). Finally, the findings
presented herein may also disclose new opportunities for further un­
derstanding the neural architecture of language processing in dual or
multiple language contexts. Topics such as individual differences in
5
Brain and Language 211 (2020) 104879
cortical development, structural brain asymmetry, and functional
hemispheric specialization are active areas of research in system
neuroscience, and yet thus far critically understudied in bilingualism
research. We suggest that neurocognitively plausible theories of bilin­
gual language processing might benefit from an increased attention to
these topics, possibly as a result of multi- and cross-disciplinary efforts.
4. Conclusions
Predominantly informed by a function-leading-structure perspective,
past structural MRI research in bilingualism has mainly focused on the
brain’s capacity to change in order to adapt to an enriched communi­
cative environment. In this paper, we discussed novel findings according
to which bilingual experience modulates neuroanatomic trait markers of
cognition, sketching a non-deterministic scenario wherein both neuro­
anatomical predispositions and experiential factors contribute to shape
behaviour. The data here presented add to an increasing body of
research which require simultaneously asking about patterns of brain
structure and patterns of behaviour at different time-scales, and possibly
provide new opportunities to begin to understand the neural basis of
behavioural differences in ways that exceed mere correlations. The
implications of these findings appear to go beyond the domain of
bilingualism, extending to other forms of experience-based plasticity
which may modulate the effect of neuroanatomical predispositions on
cognitive functions. Future research should test to what extent the
modulatory role of bilingual experience on brain-behaviour relation­
ships may be replicated by other cognitively stimulating factors or
experiential demands more in general.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
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