Brain & Language 159 (2016) 60–73

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Brain & Language

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

L1 and L2 processing in the bilingual brain: A meta-analysis

of neuroimaging studies
Hengshuang Liu a, Fan Cao b,⇑
a
Division of Psychology, School of Humanities and Social Sciences, Nanyang Technological University, 14 Nanyang Drive, Singapore 637332, Singapore
b
Department of Communicative Sciences and Disorders, Michigan State University, 1026 Red Cedar Rd, Rm 104, East Lansing, MI 48823, United States

a r t i c l e i n f o a b s t r a c t

Article history: Neuroimaging studies investigating bilingual processes have produced controversial results in determin-
Received 14 July 2015 ing similarities versus differences between L1 and L2 neural networks. The current meta-analytic study
Revised 25 May 2016 was conducted to examine what factors play a role in the similarities and differences between L1 and L2
Accepted 28 May 2016
networks with a focus on age of acquisition (AOA) and whether the orthographic transparency of L2 is
Available online 10 June 2016
more or less transparent than that of L1. Using activation likelihood estimation (ALE), we found L2
processing involved more additional regions than L1 for late bilinguals in comparison to early bilinguals,
Keywords:
suggesting L2 processing is more demanding in late bilinguals. We also provide direct evidence that AOA
Meta-analysis
Bilingualism
of L2 influences L1 processing through the findings that early bilinguals had greater activation in the left
First/second language fusiform gyrus than late bilinguals during L1 processing even when L1 languages were the same in the
Age of acquisition two groups, presumably due to greater co-activation of orthography in L1 and L2 in early bilinguals. In
Orthographic depth/transparency addition, we found that the same L2 languages evoked different brain activation patterns depending
Assimilation on whether it was more or less transparent than L1 in orthographic transparency. The bilateral auditory
Accommodation cortex and right precentral gyrus were more involved in shallower-than-L1 L2s, suggesting a ‘‘sound-out”
strategy for a more regular language by involving the phonological regions and sensorimotor regions to a
greater degree. In contrast, the left frontal cortex was more involved in the processing of deeper-than-
L1 L2s, presumably due to the increased arbitrariness of mapping between orthography and phonology
in L2.
Ó 2016 Elsevier Inc. All rights reserved.

1. Introduction to understand what factors play a role in determining when L2 and

Second language acquisition is mediated by structural and func-
tional brain changes (Costa & Sebastián-Gallés, 2014; Li, Legault, &
Litcofsky, 2014; Stein, Winkler, Kaiser, & Dierks, 2014). A number
of neuroimaging studies have been conducted to understand how
the brain supports more than one language. The primary question
addressed in these studies was how different or similar brain acti-
vations for L1 and L2 are, and the findings are controversial. While
there is evidence that second language (L2) is processed using the
first language (L1) brain network (Illes et al., 1999; Nakada, Fujii, &
Kwee, 2001; Soles, 2011; Tan et al., 2003), there is also evidence
that L2 processing is related to the involvement of additional brain
areas (Huang, Itoh, Kwee, & Nakada, 2012; Liu, Dunlap, Fiez, &
Perfetti, 2007; Nelson, Liu, Fiez, & Perfetti, 2009; Ng, 2008; Park,
Badzakova-Trajkov, & Waldie, 2012). Research has been conducted
⇑ Corresponding author.
E-mail addresses: H120006@e.ntu.edu.sg (H. Liu), fcao@msu.edu (F. Cao).
L1 networks overlap and when they are different.
One such factor is age of acquisition (AOA) for L2. According to
the framework of critical period, later L2 onset age is often associ-
ated with lower L2 performance due to the maturational decay of
the ability to acquire language and/or the interference of the exist-
ing linguistic system (Archila-Suerte, Zevin, Bunta, & Hernandez,
2012; Archila-Suerte, Zevin, & Hernandez, 2015; Birdsong &
Molis, 2001; Newport, 1990). Although a decline in L2 capability
has been demonstrated in morphology, phonology, semantics,
and syntax through adulthood (Flege, Yeni-Komshian, & Liu,
1999; Johnson & Newport, 1989; Weber-Fox & Neville, 1996),
age six or seven is widely thought as the general turning point
and thus the cut-off between early and late bilinguals (Meisel,
2004, chap. 3; Perani et al., 2003). However, in the last few years,
new studies that use EEG measures have suggested much greater
plasticity for late bilinguals than previously thought. For example,
studies have found that ERPs in late L2 learners at high levels of
proficiency are not distinguishable from those of native speakers
in morpho-syntactic tasks, suggesting that AOA effects are not

http://dx.doi.org/10.1016/j.bandl.2016.05.013
0093-934X/Ó 2016 Elsevier Inc. All rights reserved.
 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73
driven by maturational constraints (Morgan-Short, Finger, Grey, &
Ullman, 2012; Morgan-Short, Steinhauer, Sanz, & Ullman, 2012;
Steinhauer, 2014). Another ERP study also found that adult second
language learners show rapid L2 word learning after a minimal
instruction of 14 h (McLaughlin, Osterhout, & Kim, 2004), suggest-
ing the great plasticity in late bilinguals.
Whether and how L2 AOA influences the similarities and differ-
ences between L1 network and L2 network is still debatable. Neu-
rocognitive studies have documented that early bilinguals give rise
to a language processing network that is activated homogeneously
by L1 and L2 (Bloch et al., 2009; Cherodath & Singh, 2015; Saur
et al., 2009; Soles, 2011), while late bilinguals produce a more dif-
ferentiated brain network for L2 (Huang et al., 2012; Nelson, Liu,
Fiez, & Perfetti, 2005; Park et al., 2012; Wang et al., 2011). In con-
trast, there is also evidence that early bilinguals are characterized
by a more differentiated network for L1 and L2 (Das, Padakannaya,
Pugh, & Singh, 2011), while late bilinguals show an overlap
between L1 and L2 processing (Cao et al., 2013; Chee, 2009; Tan
et al., 2003). Kim, Relkin, Lee, and Hirsch (1997) even compared
early and late bilinguals in one study and found late bilinguals
compared to early bilinguals had more separated L1 and L2 activa-
tions in frontal lobe while both bilingual groups showed little sep-
aration of L1 and L2 activity in temporal lobe. However, this
comparison was based on visual inspection instead of direct statis-
tical comparison between early and late bilinguals. To our knowl-
edge, no published studies have conducted direct statistical
comparison between early and late bilinguals on the contrast of
L2 > L1 to examine whether early bilinguals have a more over-
lapped network between L1 and L2 than late bilinguals or vice
versa. The current meta-analytic study aimed to address this issue
by examining the effect of L2 AOA on the contrast of L2 > L1 across
previously published studies.
Even though many efforts have been made to study how AOA
influences L2 brain activation patterns, far less attention has been
paid to how L2 AOA influences L1 processing. Research has sug-
gested that acquisition of L2 also influences L1 processing (Costa
& Sebastián-Gallés, 2014, for a review), probably due to three rea-
sons: (1), reduced usage of L1 (Baus, Costa, & Carreiras, 2013;
Gollan, Montoya, Cera, & Sandoval, 2008; Ivanova & Costa, 2008);
(2), continuous interaction of L1 and L2 which leads to linguistic
transfer from L2 to L1 (Malt, Li, Pavlenko, Zhu, & Ameel, 2015;
Pavlenko & Malt, 2011; Runnqvist, Gollan, Costa, & Ferreira,
2013); (3) the need to constantly control and monitor two lan-
guages (Green, 1998; Kroll, Bobb, Misra, & Guo, 2008). Recent neu-
roimaging evidence also suggests that L1 is more permeable than
previously thought, in the sense that after L2 is acquired, it is pos-
sible for L1 processing to evoke either stronger (Jones et al., 2011;
Mei et al., 2015) or weaker (Mei et al., 2014) activation than before.
Stronger activation has been found in the right fusiform gyrus in
reading English (L1) words after Chinese (L2) was learned in a
group of English speaking adults (Mei et al., 2015), suggesting that
brain areas that are associated with L2 processing (the right fusi-
form as a typical Chinese area; Bolger, Perfetti, & Schneider,
2005; Wu, Ho, & Chen, 2012) may also become involved in L1 pro-
cessing. Another study found that the left fronto-temporal cortex
was more activated in Greek (L1) articulation for Greek-English
bilinguals than for Greek monolinguals (Jones et al., 2011), pre-
sumably due to the greater co-activation and monitoring mecha-
nism involved in bilinguals than monolinguals even during L1
processing. On the other hand, weaker activation has also been
found in the bilateral frontal and occipital cortices during Chinese
(L1) word reading after the semantics of an artificial language (L2)
was learned in a group of Chinese speaking adults (Mei et al.,
2014), suggesting the orthographic and semantic processing of
L1, mediated by frontal and occipital activation, may be more vul-
nerable to the influence of a new language in comparison to the
61
phonological or syntactic processing of L1. Interestingly, one study
has found not only weaker activation in the classical left-
lateralized language areas (e.g., left middle temporal gyrus, left
inferior occipital gyrus), but also stronger activation in some
non-typical language areas (e.g., posterior cingulate cortex, right
superior temporal gyrus) for Spanish-Catalan early bilinguals as
compared to Spanish monolinguals during Spanish (L1) picture
naming (Palomar-García et al., 2015), suggesting early bilinguals
utilize a more distributed and less efficient network for L1 process-
ing than monolinguals. Recent ERP studies have also suggested
native language changes during L2 acquisition (Chang, 2013), even
during early stages of L2 learning (Bice & Kroll, 2015). Taken
together, a handful of studies have shown brain changes in several
regions during L1 processing after L2 is acquired. However, very
few studies have examined the effect of L2 AOA on L1 processing
in the brain.
Past behavioral studies have suggested L1 processing to be
more immune to the backward influence of L2 processes for late
bilinguals as compared to early bilinguals; possibly due to the
well-established L1 system in late bilinguals (Gathercole &
Moawad, 2010; Pavlenko, 2011). However, this behavioral finding
has not been ascertained by neuroimaging data. The only related
published neuroimaging study found that, compared to early bilin-
gual children, L1 sentence processing in late bilingual children was
associated with greater activation in the bilateral prefrontal cortex
and the bilateral superior temporal gyrus (Jasinska & Petitto, 2013).
This implies that L2-to-L1 influence is greater for later bilinguals;
which is opposite to the behavioral finding that early bilinguals
are subject to greater influence from L2 to L1 (Gathercole &
Moawad, 2010; Pavlenko, 2011). Hence, the research on how L2
AOA influences L1 processing is still very sparse and inconclusive,
and the current meta-analytic study attempted to directly investi-
gate how L1 network is mediated by L2 AOA by comparing L1 acti-
vation in early versus late bilinguals with L1 languages, L2
proficiency, and tasks matched between the two groups.
In comparison to L2 AOA, orthographic transparency has gained
surprisingly little attention in determining what brain regions are
involved in L2. Orthographic transparency refers to the regularity
of mapping from graphemes to phonemes (Liberman, Liberman,
Mattingly, & Shankweiler, 1980). An orthography in which one
grapheme consistently maps to one phoneme is orthographically
shallow/transparent (e.g., Italian). An orthography in which one
grapheme corresponds to more than one phoneme or one
phoneme can be written in multiple ways is orthographically
deep/opaque (e.g., English; Katz & Frost, 1992). Fig. 1 illustrates a
continuum of orthographic depth/transparency for some languages
(adapted from Fig. 2.4 in Perfetti & Dunlap, 2008; Table 1 in
Chiswick & Miller, 2005; Rogers, 2004), where these languages
are sequentially listed from the shallowest orthography (Finnish,
on the top) to the deepest orthography (Hebrew, at the bottom)
with the respective orders numerally specified. It has been pro-
posed by the orthographic depth hypothesis that shallower
orthographies can be decoded according to the orthography-to-
phonology correspondence to a greater degree, whereas deeper
orthographies need more lexical-level procedures to match the
spelling of a whole word with its phonology stored in memory
(Bolger et al., 2005; Fern-Pollak, 2008; Katz & Frost, 1992;
Seymour, Aro, & Erskine, 2003; Ziegler & Goswami, 2005). Cross-
linguistic studies have provided neurocognitive evidence that
orthographic depth influences brain activation. The shallower
orthography is associated with greater involvement of the left
temporo-parietal regions underlying phonological assembly,
whereas the deeper orthography is more related to lexical-level
processing regulated by the ventral prefrontal gyrus and left infe-
rior temporal gyrus (Paulesu et al., 2000). This cross-linguistic dif-
ference has also been replicated in bilingual studies. One study
62 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73
Fig. 1. Orthographic depth of some languages (adapted from Fig. 2.4 in Perfetti &
Dunlap, 2008; Table 1 in Chiswick & Miller, 2005; Rogers, 2004).
found that early Hindi-English bilinguals showed greater activa-
tion in left inferior parietal lobule when reading Hindi (shallower
than English) as compared with higher activity in left inferior tem-
poral gyrus for English processing (deeper than Hindi; Das et al.,
2011). Another study found Spanish-English bilinguals had greater
activation in the superior temporal gyrus during Spanish process-
ing (shallower than English) in contrast to increased involvement
of left middle frontal gyrus in English processing (deeper than
Spanish; Jamal, Piche, Napoliello, Perfetti, & Eden, 2012). There is
also one English-Chinese bilingual study which observed increased
activity in left superior temporal gyrus for English reading (shal-
lower than Chinese) and greater activation in left middle frontal
gyrus for Chinese processes (deeper than English; Nelson et al.,
2009). The relative orthographic depth of L1 and L2 appeared to
play an important role in determining what brain regions are
involved in L1 and L2.
Furthermore, it appears that the relative orthographic depth of
both L1 and L2 does not only influence brain activation of L1 and
L2, but also influences how similar or different L1 and L2 networks
are. We have observed that when L2 is more transparent than L1,
such as in the case of Chinese-English bilinguals, L2 could be pro-
cessed using the L1’s network without much accommodation
needed (Cao et al., 2013; Tan et al., 2003), while when L2 is more
opaque than L1 such as in the case of Spanish-English bilinguals,
L2 processing is associated with significant involvement of addi-
tional regions (Jamal et al., 2012). In order to directly test how
the relative orthographic depth influences brain activation of L2,
an ideal experiment would investigate two groups of bilinguals
with the same L2 language wherein, for one group of bilinguals,
the L2 is more transparent than their L1 and, in the other group
of bilinguals, the L2 is more opaque than their L1. If it is found that
the same L2 is related to different activation patterns depending on
whether it is more transparent or more opaque than L1, then there
is direct evidence that L2’s transparency in relation to L1 influences
how L2 is processed in the brain. In other words, the existing L1
mechanism determines what L2 network would be. To date, no
published studies have addressed this issue, probably due to the
challenge of finding such two bilingual groups in one study.
Meta-analytic studies serve as a great choice for such a purpose.
The current meta-analysis aimed to examine three questions:
(1) the effect of L2 AOA on the similarity between L2 and L1 net-
works, (2) the effect of L2 AOA on L1 activation, and (3) the effect
of orthographic depth of L2 in relation to that of L1 on L2 activa-
tion. For the first question, we chose a group of early bilingual
studies and a group of late bilingual studies controlling for the L2
proficiency level, participants’ age, and tasks used in the neu-
roimaging procedure. The relation between L1 and L2 in terms of
orthographic transparency was also controlled (see Table 2). We
then compared the two groups on brain activation in the contrast
of L2 > L1 to determine how L2 AOA influenced L2 processing in the
bilingual brain. For the second question, we also chose a group of
early bilingual studies and a group of late bilingual studies with
the same first languages in both groups. In addition, task, L2 profi-
ciency, participants’ age, and relative L2 to L1 transparency were
also matched between the two groups (see Table 2). We then com-
pared brain activation for L1 between early and late bilingual
groups to examine how L2 AOA impacted L1 activation. For the
third question, we chose a group of studies with L2 being more
transparent than L1 and a group of studies with L2 being more opa-
que than L1. The L2s were the same languages in the two groups,
with task, participants’ age, L2 AOA, and L2 proficiency all equiva-
lent between groups (see Table 2). We then compared brain activa-
tion of L2 between the two groups to test how relative
transparency of L2 to L1 influenced brain activation of L2.
2. Methods
2.1. Literature selection
With ‘‘(bilingual or bilingualism) and (brain imaging or fMRI or
PET)” as keywords, neuroimaging articles published from 1998 to
2014 were searched through the following databases: PsycINFO,
Scopus, ScienceDirect, Web of Science, and Google Scholar. The
preliminary inclusive criteria were as follows: (1) fMRI or PET stud-
ies reporting complete coordinates in Talairach (Talairach &
Tournoux, 1988) or Montreal Neurological Institute (MNI; Collins
et al., 1998) through whole-brain scanning; (2) studies recruiting
healthy early or late bilingual adults (18–50 years), with no restric-
tion on the two specific languages for each bilingual sample; (3)
studies examining the activation of the first language, the second
language, or the contrast between them. On the other hand, studies
meeting any of the following criteria were excluded: (1) ERP or
MEG studies (through abstract screening); (2) studies employing
bilinguals under 18 years or above 50 years, or employing
unhealthy bilinguals (through abstract screening); (3) studies
without coordinates reported on whole-brain activation (through
full-text article assessment); (4) studies without activation
reported for L2 > L1, L1, or L2 (through full-text article assess-
ment); (5) studies mainly evaluating non-linguistic processing
such as working memory or executive functioning (through full-
text article assessment). Following the inclusive and exclusive cri-
teria, studies reporting activation of L2 > L1, L1, or L2 were respec-
tively sorted into Set A, B, or C for final selection before further
investigating the three questions (Fig. 2).
Set A was used to study whether L2 AOA plays a role in deter-
mining how similar or different the brain activation of L2 was to
that of L1, which was based on the comparison between early-
and late-bilinguals on the ‘‘L2 > L1” activation. The relative depth
of L2 versus L1 (Table 1) was obtained by subtracting the trans-
parency order of L1 from that of L2 according to the ordering in
Fig. 1. Independent-sample T test was performed to ascertain that
the relative depth of L2 versus L1 was equal in early- and late-
bilingual studies. Tasks used in the selected studies were catego-
rized into articulatory, phonological, orthographic, semantic, and
sentential categories (Table 2). Chi-square test was conducted to
justify that different types of tasks were equally distributed in
early- and late-bilingual studies. L2 proficiency was graded into
‘‘high”, ‘‘moderate”, and ‘‘low” levels (see Table 1, the ‘‘L2 PL”
column), with those scored above 80% in the proficiency
 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73
Fig. 2. PRISMA flow chart of the literature search.
measurements defined as ‘‘high”, between 60% and 80% defined as
‘‘moderate”, and below 60% defined as ‘‘low”. For those without
quantitative measurements of L2 proficiency, categorization was
done according to the statement of the language proficiency in
the specific article. All studies included in Set A were with high
proficient L2. In total, 13 early-bilingual studies and 27 late-
bilingual studies were matched on participants’ age (t = 1.652,
p = 0.110), the relative depth of L2 versus L1 (t = 0.513, p = 0.611),
and the distribution of task types (v2 = 4.383, p = 0.223) (Table 2).
Thereby, any activation difference between the early and late bilin-
guals should be driven by L2 AOA.
Set B was used to detect whether the AOA of L2 would, in
reverse, influence the first language network based on the compar-
ison between early and late bilinguals in the L1 activation. 15 stud-
ies were chosen for early- and late-bilingual groups each with the
L1s exactly matched: 8 Chinese, 5 English, 1 German, and 1 Ladin.
In addition to L1s, participants’ ages (t = 1.348, p = 0.192), the
transparency difference between L2 and L1 (t = 0.687, p = 0.498),
the distribution of L2 proficiency levels (v2 = 2.360, p = 0.307),
and the distribution of task types (v2 = 2.863, p = 0.581) were also
equivalent between early- and late-bilingual groups (Table 2).
Thus, between-group differences in L1 activation should be attrib-
uted to L2 AOA.
Set C was used to assess whether the orthographic transparency
of L2 relative to L1 affects the neurocognitive activation of the
second language, in other words, Set C was to assess L2 activation
63
differences between the L2 with more transparent orthography-to-
phonology mapping than L1 (labeled as ‘‘shallower L2”) and the L2
with more opaque orthography-to-phonology mapping than L1
(labeled as ‘‘deeper L2”). 25 studies were chosen for each group
with the L2s exactly matched: 18 English, 2 Japanese, and 5 French.
The two groups differed in the depth of L2 relative to L1 (t = 10.077,
p < 0.001), but were homogeneous on participants’ ages (t = 0.392,
p = 0.698), L2’s AOA (v2 = 2.381, p = 0.123), L2 proficiency levels
(v2 = 1.690, p = 0.430), and task types (v2 = 3.759, p = 0.440)
(Table 2). Thus, L2 to L1 orthographical depth would likely be the
only contributor to the group difference in L2 activation in Set C.
Following the entire selection procedure as illustrated in the
Preferred Reporting Items for Systematic Reviews and Meta-
Analyses (PRISMA) flow chart (Fig. 2), a total of 81 studies (Table 1)
consisting of three sets were included in the meta-analyses. Among
them, four common studies were shared by Set A, B, and C; three
studies were common for Set A and B, 17 for Set B and C, and 10
for Set A and C; only 23 studies were exclusively included in Set
A, six in Set B, and 19 in Set C (Fig. 3). The key information of these
selected studies was summarized in Table 2.
2.2. Data analysis
The present meta-analyses were performed using the revised
algorithm of Activation-Likelihood Estimation (ALE) analyses
(Eickhoff, Bzdok, Laird, Kurth, & Fox, 2012; Eickhoff et al., 2009;
Table 1

64
Important information about studies selected for Set A, B, and C.

Studya L1 L2 L2 AOA L2 vs. L1 L2 Sample size Task

proficiency (subjects’ ageb)
Set A
Perani et al. (2003) Catalan Spanish Early Shallower High 5 (23.5 yrs) Overt word generation – rest
1
Klein, Watkins, Zatorre, English French Early Shallower High 10 (22 yrs) Overt nonword repetition – silence
and Milner (2006) 1
Hernandez (2009) Spanish English Early Deeper High 12 (21.4 yrs) Covert picture naming – rest
11
Perani et al. (2003) Spanish Catalan Early Deeper High 6 (23.5 yrs) Overt word generation – rest
1
Wartenburger et al. (2003) Italian German Early Deeper High 11 (26.9 yrs) Sentence semantic/grammatical violation judgment – fixation
9
Perani et al. (1998) Spanish/Catalan Catalan/Spanish Early Equal High 12 (23 yrs) Story listening – backward story listening
0
Saur et al. (2009) German/French French/German Early Equal High 12 (26.2 yrs) Auditory sentence judgment – silence
0
Nakamura et al. (2010) Japanese English Late Shallower High 24 (25 yrs) Word categorization judgment – word absent baseline
1

H. Liu, F. Cao / Brain & Language 159 (2016) 60–73

Saur et al. (2009) French German Late Shallower High 12 (41.6 yrs) Auditory sentence judgment – silence
5
Abutalebi (2008) German French Late Deeper High 12 (25.4 yrs) Overt picture naming – fixation
5
De Bleser and Kauschke (2003) Dutch French Late Deeper High 11 (19.5 yrs) Covert picture naming – fixation
3
Halsband (2006) Finnish English Late Deeper High 10 (27.3 yrs) Retrieval of high-imagery/low-imagery word pairs – non-sense words
17
Jeong, Sugiura, Sassa, Haji, et al. Korean English Late Deeper High 30 (28.5 yrs) Auditory sentence understanding judgment – white noise
(2007) 8
Park et al. (2012) Macedonian English Late Deeper High 8 (24.88 yrs) Letter case judgment – fixation; lexical decision – fixation
12
Saur et al. (2009) German French Late Deeper High 12 (29.3 yrs) Auditory sentence judgment – silence
5
Vingerhoets et al. (2003) Dutch French English Late Deeper High 12 (27.6 yrs) Covert word fluency – covert number counting; covert picture
3.5 naming – scratch drawing; silent text reading – pseudoword reading
Wartenburger et al. (2003) Italian German Late Deeper High 12 (29.2 yrs) Sentence semantic/grammatical violation judgment – fixation
9
Set B
Lin, Wu, Li, and Guo (2013) Chinese Japanese Early Shallower High 9 (23.2 yrs) Character size decision – rest
1
Ng (2008) English Chinese Early Deeper High 8 (26.63 yrs) Word passive viewing – checkerboard
2
Koyama, Stein, Stoodley, and English Japanese kana Late Shallower High 14 (26.2 yrs) One-back matching – Tibet control
Hansen (2013) 8
Stein et al. (2009) English German Late Shallower Low 10 (17 yrs) Noun meaning known judgment – fixation
6
Videsott et al. (2010) Ladin English Late Deeper Low 20 (27.3 yrs) Overt picture naming – fixation
14
Liu et al. (2007) English Chinese Late Deeper Low 23 (27.62 yrs) Word passive viewing – fixation
2
Set C
Soles (2011) English French Early Shallower High 15 (23.3 yrs) Overt sentence reading-unreadable sentence
1
Cao, Kim, Liu, and Liu (2014) Chinese English Late Shallower High 15 (22.9 yrs) Word rhyming – baseline
2
Table 1 (continued)

Studya L1 L2 L2 AOA L2 vs. L1 L2 Sample size Task

proficiency (subjects’ ageb)
Luke, Liu, Wai, Wan, and Tan (2002) Chinese English Late Shallower High 7 (25.5 yrs) Phrase syntactic/semantic plausibility judgment – font size judgment
2
Kovelman, Baker, and Petitto (2008) Spanish English Early Deeper High 11 (19 yrs) Sentence semantic plausibility judgment – fixation
11
Rao, Mathur, and Singh (2013) Hindi English Early Deeper High 15 (22.5 yrs) Concrete noun judgment – rest
7
Waldron and Hernandez (2013) Spanish English Early Deeper High 11 (21.5 yrs) Covert past sense generation – rest
11
Buchweitz, Shinkareva, Mason, Portuguese English Late Deeper High 11 (29.9 yrs) Dwellings/tools noun categorization – fixation
Mitchell, and Just (2012) 9
Golestani et al. (2006) French English Late Deeper Moderate 12 (24 yrs) Covert sentence generation – silence; covert words reading – silence
1
Halsband (2006) Finnish English Late Deeper High 10 (27.3 yrs) Encoding of high-imaginary/low-imaginary word pairs – non-sense words
17
Jeong, Sugiura, Sassa, Haji, et al. Korean Japanese Late Deeper High 30 (28.5 yrs) Auditory sentence understanding judgment – white noise
(2007) 9
Perani et al. (1998) Italian English Late Deeper Moderate 9 (34.5 yrs) Story listening – backward story listening
15

H. Liu, F. Cao / Brain & Language 159 (2016) 60–73

Vingerhoets et al. (2003) Dutch English/ Late Deeper High 12 (27.6 yrs) Covert picture naming – scratch drawing (only for Dutch-French bilinguals);
French 4/3 Covert word fluency – covert number counting; silent text reading – pseudoword
reading
(for both Dutch-English bilinguals and Dutch-French bilinguals)
Set A & B
Klein et al. (2006) English French Early Shallower High 10 (22 yrs) Overt word repetition – silence
1
Videsott et al. (2010) Ladin Italian Early Shallower High 20 (27.3 yrs) Overt picture naming – fixation
1
Chee et al. (2001) English Chinese Early Deeper High 10 (24 yrs) Pyramids and palm trees semantic relatedness judgment– size judgment
2
Set B & C
Abutalebi et al. (2013) German English Early Deeper Moderate 14 (23.5 yrs) Overt picture naming – baseline
6
Chan et al. (2008) Chinese English Early Shallower High 11 (26.5 yrs) Noun/verb lexical decision – fixation
2
Frenck-Mestre, Anton, Roth, English French Early Shallower High 6 (24 yrs) Overt words/sentences articulation – nonpronounceable consonant strings
Vaid, and Viallet (2005) 1
Tan, Rajapakse, Hong, Lee, Chinese English Early Shallower High 6 (17 yrs) Covert word reading – Indian-tamil fonts
and Sitoh (2004) 2
Ng (2008) Chinese English Early Shallower High 16 (32.63 yrs) Word passive viewing – checkerboard
2
Tham Wei Ping (2003) Chinese English Early Shallower High 6 (20.5 yrs) Homophone matching – fixation
2
Ng (2008) Chinese English Early Shallower Low 8 (29 yrs) Word passive viewing – checkerboard
2
Frenck-Mestre et al. (2005) English French Late Shallower High 6 (41 yrs) Overt words/sentences articulation – nonpronounceable consonant strings
1
Huang et al. (2012) Chinese Japanese Late Shallower High 14 (30 yrs) Covert sentence reading – unknown language
1
Tan et al. (2003) Chinese English Late Shallower High 12 (34 yrs) Word rhyming – font size
2
Wang, Wang, and Lee (2010) Chinese English Late Shallower Moderate 12 (23.5 yrs) Silent word reading – fixation
2
Set A & C
Jamal et al. (2012) Spanish English Early Deeper High 12 (22.67 yrs) Ascender letter detecting: real word-pseudofonts
11

65
(continued on next page)
 66 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73

Laird et al., 2005; Turkeltaub, Eden, Jones, & Zeffiro, 2002) imple-

Participants’ mean age was not reported in the following studies, where the minimal and maximal ages were averaged as the mean age (Chan et al., 2008; Chee et al., 2001; De Bleser and Kauschke, 2003; Golestani et al., 2006;
mented in GingerALE 2.3.2. This approach generates an estimate

Conjoined sentence comprehension – rest; embedded sentence comprehension – rest

of how likely the convergence of activation across studies is
located at specific coordinates on a voxel-by-voxel basis. For con-
trast analysis between groups, group discrepancy in study sizes

Pyramids and palm trees semantic relatedness judgment– size judgment

(e.g., Set A: 13 early bilingual studies, 27 late bilingual studies)
can be corrected by this approach (Eickhoff et al., 2011): Simulated

Huang et al., 2012; Jeong, Sugiura, Sassa, Haji, et al., 2007; Luke et al., 2002; Nakamura et al., 2010; Perani et al., 1998, 2003; Rao et al., 2013; Tan et al., 2003; Tham Wei Ping, 2003; Wang et al., 2010).
Orthographic search –fixation; semantic categorization – fixation
data is created by pooling the foci data sets and randomly dividing
them into two groups of the same size as the original data sets (e.g.,
for Set A, this step would generate two simulated groups: one
Auditory sentence understanding judgment – white noise

Auditory sentence understanding judgment – white noise

Auditory sentence understanding judgment – white noise

Covert picture naming (cognates/non-cognates) – fixation

group with 13 randomly-chosen studies, the other with the rest

27 studies). Subtraction is then conducted between the two simu-
Semantic plausibility judgment – 3 nouns or verbs

lated groups. After thousands of permutations, thousands of sub-

traction results jointly yield a null distribution which could serve
as a baseline to be compared with the true contrasted result
(e.g., for Set A, the true contrast between the 13 early bilingual
studies and the 27 late bilingual studies). A voxel-wise P value
image is then resulted, showing the significance level of the true
contrast distinguished from the null distribution in every specific
Covert word reading – rest

voxel.
In the current study, all relevant foci reported in the selected
studies were extracted for ALE analyses. Coordinates in MNI space
were first converted into Talairach space. Coordinates of the con-
trast ‘‘L2 > L1” were extracted for Set A; coordinates of the contrast
‘‘L1 > baseline” were extracted for Set B; coordinates of the con-
Task

trast ‘‘L2 > baseline” were extracted for Set C. Only clusters that
had been reported in no less than two studies (if the total number
(subjects’ ageb)

of studies <15) or no less than three studies (if the total number of
12 (22.3 yrs)

36 (21.1 yrs)

11 (19.5 yrs)

16 (22.9 yrs)

6 (22.67 yrs)
Sample size

studiesP15) were included in further analyses (Turkeltaub &

9 (28.5 yrs)
12 (26 yrs)

18 (27 yrs)

12 (26 yrs)

Branch Coslett, 2010; Wagner, Sebastian, Lieb, Tuscher, & Tadic,

2014). ALE analyses were performed for each subset separately
with a threshold at false discovery rate (FDR) p < 0.05 and a mini-
mum cluster volume of 150 mm3. Contrast analyses were then
proficiency

conducted between the two subsets within each set using a thresh-
Note: Studies with multiple tasks were used more than once in data analysis (e.g., Buchweitz et al., 2012).

old at p < 0.05 (uncorrected) with 10,000 permutations and a min-

High

High

High

High

High

High

High

High

High
L2

imum cluster volume of 150 mm3. This resulted in three pairs of

contrasts: early ‘L2 > L1’ vs. late ‘L2 > L1’ for Set A, early L1 vs. late
Shallower

Shallower

Shallower

Shallower

Shallower
L2 vs. L1

L1 for Set B, and shallower L2 vs. deeper L2 for Set C.

Deeper

Deeper

Deeper

Deeper
8/9
1

2
11

8
L2 AOA

Early

Late

Late

Late

Late

Late

Late

Late

Late

3. Results

3.1. The effect of L2 AOA on the activation of L2 > L1

Japanese

Japanese
English/

As shown in Fig. 4 and Table 3, the overlap regions for early and
English

English

English

English

English

English
French

late bilinguals on the contrast of L2 > L1 were the left middle fron-
L2

tal gyrus, the left inferior frontal gyrus, the left precentral, and the
left insula. A statistical comparison between the early and late
bilinguals revealed greater activation in the left superior frontal
gyrus (BA 6/8) for late bilinguals than early bilinguals and no
Japanese
Chinese

Chinese

Chinese

Chinese
Spanish

Korean

Korean
Dutch

greater activation for early bilinguals than late bilinguals.

L1

Meschyan and Hernandez (2006)

Jeong, Sugiura, Sassa, Yokoyama,

Jeong, Sugiura, Sassa, Yokoyama,

Jeong, Sugiura, Sassa, Yokoyama,

De Bleser and Kauschke (2003)

3.2. The effect of L2 AOA on L1 activation

Even when early bilingual and late bilingual groups had the
Yokoyama et al. (2006)

same languages as their L1s, we found a significant difference in

Chee et al. (2001)

Ding et al. (2003)

Suh et al. (2007)

their brain activation during L1 processing (Fig. 5 & Table 3). The
Table 1 (continued)

et al. (2007)

et al. (2007)

et al. (2007)
Set A & B & C

left fusiform gyrus (BA 37) was found to be more activated in early
bilinguals than late bilinguals during L1 processing, with age, tasks
Studya

and L2 proficiency matched between the two groups. However, no

region was observed to be more activated for late bilinguals than
a

early bilinguals.
 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73 67

Table 2
Factors that have been matched between the two subgroups within each set. Set A: transparency of L2 in relation to L1, L2 proficiency, participants’ age, and tasks; Set B: L1s,
transparency of L2 in relation to L1, L2 proficiency, participants’ age, and tasks; Set C: L2s, L2 AOA, L2 proficiency, participants’ age, and tasks.

Set Languages Relative L2 L2 Participants’ age Tasks

orthographic AOA proficiency averaged across
depth of L2 to L1 studies
A: L2 – L1 Early 2(L1 English, L2 French), 4 shallower, 7 13 13 high 23.73 yrs 7 1 Overt picture naming – fixation,
1(L1 Ladin, L2 Italian), deeper, 2 equal early Articulatory 1 Covert picture naming – rest,
1(L1 Catalan, L2 Spanish), Average depth of 2 Overt word/nonword repetition –
1(L1 Spanish, L2 Catalan), L2 vs. L1: 3.85 silence,
1(L1 English, L2 Chinese), 2 Overt word generation – rest,
3(L1 Spanish, L2 English), 1 Covert word reading – rest
2(L1 Italian, L2 German), 1 1 Ascender letter detecting: real
1(L1 Catalan/Spanish, L2 Orthographic word-pseudofonts
Spanish/Catalan), 1 Semantic 1 PPT semantic relatedness
1(L1 German/French, L2 judgment– size judgment
French/German) 4 Sentential 1 Sentence semantic violation
judgment – fixation,
1 Sentence grammatical violation
judgment – fixation,
1 Auditory sentence judgment –
silence,
1 Story listening – backward story
listening
Late 2(L1 Japanese, L2 English), 7 shallower, 20 27 late 27 high 26.22 yrs 6 1 Overt picture naming – fixation,
4(L1 Chinese, L2 English), deeper Articulatory 4 Covert picture naming – fixation/
1(L1 Chinese, L2 Japanese), Average depth of scratch drawing,
1(L1 French, L2 German), L2 vs. L1: 4.83 1 Covert word fluency – covert
2(L1 German, L2 French), number counting
3(L1 Dutch, L2 French), 2 1 Letter case judgment – fixation,
2(L1 Finnish, L2 English), Orthographic 1 Orthographic search –fixation
4(L1 Korean, L2 English), 6 Semantic 1 PPT semantic relatedness
1(L1 Korean, L2 Japanese), judgment– size judgment,
2(L1 Macedonian, L2 2 Semantic categorization –
English), fixation,
3(L1 Dutch, L2 French 2 Word pairs retrieval – non-sense
+ English), words,
2(L1 Italian, L2 German) 1 Lexical decision – fixation
13 Sentential 2 Sentence comprehension
judgment– rest,
7 Auditory sentence understanding
judgment– white noise/silence,
2 Sentence semantic violation
judgment – fixation/3 nouns or
verbs,
1 Sentence grammatical violation
judgment – fixation,
1 Silent text reading – pseudoword
reading
B: L1 Early L1s: 8 Chinese, 5 English, 1 13 shallower, 2 15 14 high, 1 24.69 yrs 10 2 Overt picture naming – fixation,
German, 1 Ladin deeper early low Articulatory 2 Overt words/sentence
Average depth of articulation – nonpronounceable
L2 vs. L1: 1.6 consonant strings,
1 Overt word repetition – silent,
5 Covert word reading – Indian-
tamil fonts/checkerboard
1 1 Character size decision –rest
Orthographic
1 1 Homophone matching – fixation
Phonological
3 Semantic 1 PPT semantic relatedness
judgment– size judgment,
2 Lexical decision – fixation
Late L1s: 8 Chinese, 5 English, 1 12 shallower, 3 15 late 11 high, 3 27.62 yrs 7 2 Overt picture naming – fixation,
German, 1 Ladin deeper low, 1 Articulatory 2 Overt words/sentence
Average depth of moderate articulation – nonpronounceable
L2 vs. L1: 0.6 consonant strings,
3 Covert word reading – fixation
2 1 One-back matching – Tibet
Orthographic control,
1 Orthographic search – fixation
1 1 Word rhyming – font size
Phonological
3 Semantic 1 PPT semantic relatedness
judgment– size judgment,
1 Semantic categorization –
fixation,

(continued on next page)

68 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73

Table 2 (continued)

Set Languages Relative L2 L2 Participants’ age Tasks

orthographic AOA proficiency averaged across
depth of L2 to L1 studies
1 Noun meaning known judgment
– fixation
2 Sentential 1 Auditory sentence understanding
judgment– white noise,
1 Sentence silent reading –
unknown language
C: L2 Shallower L2s: 18 English, 2 Japanese, 25 shallower 10 22 high, 1 26.06 yrs 11 4 Overt words/sentence
5 French Average depth of early, low, 2 Articulatory articulation – nonpronounceable
L2 vs. L1: 1.68 15 late moderate consonant strings,
1 Overt sentence reading –
unreadable sentence,
6 Covert word reading – Indian-
tamil fonts/checkerboard/fixation
1 1 Orthographic search – fixation
Orthographic
3 1 Homophone matching – fixation,
Phonological 2 Word rhyming – font size
4 Semantic 1 PPT semantic relatedness
judgment– size judgment,
2 Lexical decision – fixation,
1 Semantic categorization –fixation
6 Sentential 2 Auditory sentence understanding
judgment– white noise,
2 Sentence semantic plausibility
judgment – 3 nouns or verbs/font
size judgment,
1 Sentence syntactic plausibility
judgment – font size judgment,
1 Sentence silent reading –
unknown language
Deeper L2s: 18 English, 2 Japanese, 25 deeper 5 21 high, 4 25.39 yrs 10 1 Overt picture naming – baseline,
5 French Average depth of early, moderate Articulatory 3 Covert picture naming – fixation/
L2 vs. L1: 7.64 20 late scratch drawing,
2 Covert word fluency – covert
number counting,
2 Covert word reading – rest,
1 Covert past sense generation –
rest,
1 Covert sentence generation –
silence
1 1 Ascender letter detecting: real
Orthographic word-pseudofonts
5 Semantic 3 Noun categorization – fixation,
2 Word pairs encoding – non-sense
words
9 Sentential 3 Auditory sentence understanding
judgment– white noise,
1 Sentence semantic plausibility
judgment – fixation,
2 Sentence comprehension
judgment – rest,
2 Silent text reading – pseudoword
reading,
1 Story listening – backward story
listening

3.3. The effect of relative orthographic transparency on L2 activation 4. Discussion

We found that the same L2 languages evoked different brain
activations in the shallower L2 group and the deeper L2 group after
task, age, L2 AOA, and L2 proficiency were matched (Fig. 6 &
Table 3). The right precentral (BA 6), the left middle temporal gyrus
(BA 21), and the right superior temporal gyrus (BA 21) were more
activated in the shallower L2 group than the deeper L2 group. In
contrast, the left middle frontal gyrus (BA 46), the left inferior fron-
tal gyrus (BA 45/47), the left insula (BA 13), and the left medial
frontal gyrus (BA 8) were more activated in the deeper L2 than
the shallower L2 group.
In this meta-analytic study, we examined how L2 AOA influ-
enced L2 and L1 processing networks in the brain. We found that
L2 processing involved more additional regions than L1 process-
ing in late bilinguals than in early bilinguals, suggesting more
neural resources are needed in the L2 processing of late than
early bilinguals. We found that L1 processing involved the left
fusiform to a greater degree for early than late bilinguals, sug-
gesting greater co-activation of orthography in L1 and L2 in early
bilinguals. In this study we also examined how orthographic
transparency of L2 in relation to L1 influences L2 processing in
 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73 69

Table 3
ALE results for comparisons in Set A, B, and C.

Volume Region BA Coordinates Max

(mm3) ALE
x y z
[A] Early ‘‘L2 > L1”
800 Left insula 13 30 12 14 0.008
440 Left precentral gyrus 6 52 2 26 0.010
368 Left middle frontal gyrus 10 38 38 14 0.009
264 Left inferior frontal gyrus 9 36 8 30 0.009
[A] Late ‘‘L2 > L1”
4864 Left middle frontal gyrus, left 9, 44 10 26 0.030
precentral gyrus 44
2216 Left superior frontal gyrus, left medial 6, 8 4 12 52 0.022
frontal gyrus
1768 Left insula 47 32 24 2 0.021
544 Left inferior frontal gyrus 46 52 30 8 0.013
[A] Early ‘‘L2 > L1” vs. late ‘‘L2 > L1”: non-significant
[A] Late ‘‘L2 > L1” vs. early ‘‘L2 > L1”
952 Left superior frontal gyrus 8, 6 7 17 53 1.970
[B] Early L1 > late L1
1880 Left fusiform gyrus 37 42 56 8 3.353
[B] Late L1 > early L1: not significant
Fig. 3. Demonstration of studies included in Set A, B, and C.
[C] Shallower L2 > deeper L2
2024 Right precentral gyrus 6 46 8 34 3.891
960 Left middle temporal gyrus 21 62 28 5 2.543
560 Right superior temporal gyrus 21 47 28 5 3.719
the brain. We found that L2 processing recruited bilateral supe- [C] Deeper L2 > shallower L2
rior/middle temporal and precentral gyri to a greater degree 3672 Left middle frontal gyrus, left inferior 46, 44 18 20 3.432
when L2 was more transparent than L1, whereas the left frontal frontal gyrus 45
gyri were more involved in L2 processing if L2 was less transpar- 2456 Left inferior frontal gyrus, left insula 47, 30 18 2 3.432
13
ent than L1. These findings suggest that the same L2s are pro- 1064 Left medial frontal gyrus 8 8 22 48 2.260
cessed by different regions in the brain depending on the
existing L1 system. This is the first evidence that the relative L2
to L1 orthographic transparency plays an important role in deter- 4.1. The effect of L2 AOA on the activation of L2 > L1
mining the brain activation pattern of L2.
We found that a number of regions were more involved in L2
than in L1 and that this was especially true in late bilinguals com-
pared to early bilinguals. First, we found that several regions in the
left frontal cortex were more involved in L2 than in L1 for both
early and late bilinguals. Namely, the left middle frontal gyrus, left
inferior frontal gyrus, left precentral gyrus, and left insula. A prior
meta-analytic study (Indefrey, 2006) also found that these regions
were more involved in L2 than L1, suggesting a general L2 effect. It
is likely that, because the proficiency level of L2 is usually lower
than that of L1, more neural resources are needed for L2 in lexical
integration at left middle frontal gyrus (Siok, Perfetti, Jin, & Tan,
2004; Wu et al., 2012), in speech production and preparation at left
insula and left inferior frontal gyrus (Price, 2010; Vigneau et al.,
2011), and also in sensori-motor coordination during language
processing at left precentral gyrus (Richlan, Martin, Schurz, &
Kronbichler, 2014). It is also noted that for both early and late
bilinguals, no temporal region was seen in the ‘‘L2 > L1” compar-
ison, replicating the past finding that L1 and L2 activity had little
separation in temporal lobe for both early and late bilinguals
(Kim et al., 1997).
Second, we found that the left superior frontal gyrus (BA6/8)
had significantly greater activity for late than early bilinguals in
the contrast of L2 > L1. It suggests that more planning is needed
for coordinated movements in L2 than in L1 especially for late
bilinguals than for early bilinguals, given that the left superior
frontal gyrus (BA6/8) has been found to be involved in planning
Fig. 4. The effect of L2 AOA on the activation of L2 > L1: (a) for the contrast of of complex, coordinated movements in previous studies (Price,
L2 > L1, early bilinguals activated the left inferior frontal gyrus (L IFG, BA9), the left 2010; Talati & Hirsch, 2005; Tremblay & Gracco, 2010; Vigneau
middle frontal gyrus (L MFG, BA10), the left insula (BA13), and the left precentral et al., 2011). Meanwhile, this SFG (BA8) in our study could also
gyrus (BA6), (b) for the contrast of L2 > L1, late bilinguals showed activation in the be identified as the supplementary motor area (SMA). Together
left inferior frontal gyrus (L IFG, BA46), the left middle frontal gyrus (L MFG, BA9),
the left insula (BA47), the left precentral gyrus (BA44), and the left superior/medial
with the anterior cingulate cortex (ACC), this SMA has been found
frontal gyri (L SFG/MeFG, BA6/8), (c) late bilinguals > early bilinguals for the to be associated with language switching, especially for L2 to L1
contrast of L2 > L1 revealed the left superior frontal gyrus (L SFG, BA6/8). switching in the bilingual literature (Abutalebi et al., 2013;
70 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73

Fig. 5. The effect of L2 AOA on L1 activation. (a) L1 processing of early bilinguals was significant in the left fusiform gyrus (L FG, BA37), the left precentral gyri (BA4/6), the left
superior temporal gyrus (L STG, BA22), the left superior parietal lobule (L SPL, BA7), and the left medial frontal gyrus (L MeFG, BA6). (b) L1 processing of late bilinguals was
significant in the left precentral gyri (BA4/6), the left superior temporal gyrus (L STG, BA22/38), the left precuneus (BA7), and the left superior/medial frontal gyrus (L SFG/
MeFG, BA6). (c) L1 processing engaged the left fusiform (L FG, BA37) to a greater degree for early than late bilinguals.

observed from the left superior frontal activation in the present

study.

4.2. The effect of L2 AOA on L1 network

We found that early bilinguals showed greater activation than

Fig. 6. The effect of relative orthographic transparency of L2 to L1 on L2 activation.
(a) Shallower L2 > deeper L2 for L2 processing was significant in the right precentral
gyrus (BA6), the left middle temporal gyrus (L MTG, BA21), and the right superior
temporal gyrus (R STG, BA21). (b) Deeper L2 > shallower L2 for L2 processing was
significant in the left middle/inferior frontal gyri (L M/IFG, BA46/45), the left insula
(BA13), and the left medial frontal gyri (L MeFG, BA8).
Garbin et al., 2011). Its greater activation during L2 than L1 pro-
cessing for late bilinguals but not for early bilinguals may imply
that late bilinguals engage more translation/switching to L1 during
L2 processing than early bilinguals. Thus, more neural resources
were needed in L2 than L1 processing for late bilinguals, while
early bilinguals had reduced cortical differentiation between L2
and L1. This is consistent with previous findings that early bilin-
guals used a relatively overlapped and shared network to process
L1 and L2 (Bloch et al., 2009; Cherodath & Singh, 2015; Saur
et al., 2009; Soles, 2011) while comparably dissociated L1 and L2
networks were recruited by late bilinguals (Huang et al., 2012;
Nelson et al., 2005; Park et al., 2012; Wang et al., 2011), and this
is especially true for frontal activation (Kim et al., 1997) as
late bilinguals in the left fusiform gyrus (BA37) for L1 processing
with the L1s exactly the same across early and late bilingual
groups. This is one of the first evidences that L2 AOA influences
the L1 brain network. Prior behavioral studies (Gathercole &
Moawad, 2010; Pavlenko, 2011) indicated that L1 processing may
get more interference from early- rather than late-exposed L2s,
since early-exposed L2s are acquired almost simultaneously as
L1s and thus are unavoidably interweaved with L1s in develop-
ment, making it difficult to establish an independent L1 lexical sys-
tem. L1 processing, therefore, is more likely to be accompanied by
the incidental processing of L2 for early bilinguals than late bilin-
guals (Gathercole & Moawad, 2010; Pavlenko, 2011). The left fusi-
form gyrus has been found to be sensitive to orthographic
processing in L1 and L2 (Chan et al., 2008; Chee, Hon, Lee, &
Soon, 2001; Ng, 2008; Tham Wei Ping, 2003) and more activated
if orthographies of both languages are processed (Garbin et al.,
2011; Guo, Liu, Misra, & Kroll, 2011; Price, Green, & Von
Studnitz, 1999). Therefore, it is possible that early bilinguals co-
activated L1 and L2 orthographies during L1 processing and thus
elicited more orthographic activation in the left fusiform gyrus in
comparison to late bilinguals. These results are also supported by
an anatomical finding that learning a second language early in life
increased white matter integrity at the inferior fronto-occipital fas-
ciculus (Mohades et al., 2012). By virtue of the closer connection
with other parts of the brain (e.g., frontal area), the left fusiform
activity was likely to be enhanced in early bilinguals during their
L1 processing. However, our finding appears to be inconsistent
with Jasinska and Petitto’s (2013) findings that bilateral prefrontal
cortex and bilateral superior temporal gyrus were more activated
for late than early bilingual children during sentence judgment
task in L1 (Jasinska & Petitto, 2013). Probably, differentiated ages
 H. Liu, F. Cao / Brain & Language 159 (2016) 60–73
of samples (9 year-old children in their study vs. 20 + year-old
adults in this study), differentiated language aspects assessed (sen-
tential in their study vs. articulatory, orthographic, phonological,
semantic, and sentential in this study), and differentiated defini-
tions of early bilinguals (before 4 year-old in their study vs. before
7 year-old in this study) are all contributable to the incongruent
findings in terms of whether early or late bilinguals need more
neural resources for L1 processing (late in their study vs. early in
this study) and which specific cortical areas are differentially acti-
vated between early and late bilinguals during their L1 processing
(B PFC and B STG in their study vs. L FG in this study).
4.3. The effect of relative orthographic transparency on L2 network
We found the same L2s were associated with differential brain
activation patterns depending on whether L2 was more or less
transparent than L1. Activation in the right primary motor cortex
(BA6, right precentral gyrus), left middle temporal gyrus (BA21),
and right superior temporal gyrus (BA21) was stronger when L2
was more transparent than L1 compared to when L2 was more
opaque than L1. In contrast, greater activation was observed in
the left middle, inferior, and medial frontal gyri (BA46/47/8) and
the left insula (BA13) when L2 was more opaque than L1 compared
to when L2 was more transparent than L1. This finding is in con-
gruent with the broad concept that L1 influences how L2 is
acquired (Elston-Güttler, Paulmann, & Kotz, 2005; Flege, Frieda, &
Nozawa, 1997; Perani & Abutalebi, 2005). When learning a L2 that
is more transparent than L1, the primary sensori-motor cortex and
phonological processing areas in the temporal lobe are more
involved, probably because the brain adapts to the higher regular-
ity of grapheme-phoneme-correspondence in L2 and engages the
‘‘sound out” strategy to a greater degree. When the same L2 is
more opaque than L1, the higher-order frontal regions are more
involved as the brain adapts to the increased arbitrariness of map-
ping from orthography to phonology in L2. The left middle/inferior
frontal gyri underlying lexical integration (Hu et al., 2010; Price,
2010) were more recruited probably due to the greater need to
coordinate the relatively disparate sound, spelling, and meaning
for deeper L2s; The elevated activation in the articulation-related
left insula (Ackermann & Riecker, 2004; Dronkers, 1996) may have
enabled those harder-to-pronounce L2s to be better verbalized;
The BA 8 subserving L2 to L1 switching (Abutalebi et al., 2013;
Garbin et al., 2011) was likely to support L2 processing by translat-
ing those deeper L2s back to the more regular native languages.
Overall, our finding is accordant with past findings that increased
spelling-to-sound consistency from L1 to L2 allowed L2 processing
to rely more on premotor and temporal regions underlying phono-
logical assembly, while reduced consistency from L1 to L2 required
more lexical or semantic mediation in frontal areas to help resolve
the more conflicting pronunciation in L2 (Carreiras, Mechelli,
Estévez, & Price, 2007; Nosarti, Mechelli, Green, & Price, 2009;
Paulesu et al., 2000; Plaut, McClelland, Seidenberg, & Patterson,
1996). Therefore, learning of the same L2 may involve different
strategies and brain changes depending on the existing L1 system.
This finding fits well with the adaptive control hypothesis which
argues that brain networks that support bilingual processing
change with the nature of the bilingualism (Green & Abutalebi,
2013).
One limitation of the current study is that a larger number of
late than early bilingual studies were included in both the shal-
lower L2 and deeper L2 groups, which may imply that the current
finding was slightly more driven by the late bilingual studies.
Future research is needed to examine any possible interaction
between L2 AOA and orthographic depth of L2 in relation to L1.
Another limitation is that even though the type of tasks was
matched in the two comparing groups for each question, the tasks
71
were still not exactly the same, therefore, there is a possibility that
the group differences observed in the current study were due to
task difference. Future research should be conducted with more
carefully matched tasks.
5. Conclusions
The current meta-analytic study was conducted to examine
which factors played a role in determining brain activation pat-
terns of L1 and L2 with a focus on L2 age of acquisition (AOA)
and whether the orthography of L2 was more or less transparent
than L1. Three major findings have been reported: (a) L1 and L2
networks were more divergent for late bilinguals than for early
bilinguals, with the coordination areas (left SFG) more involved
in L2 processing in late bilinguals than in early bilinguals, indicat-
ing more neural resources are needed for late than early bilinguals’
L2 processes; (b) L1 processes engaged the left fusiform to a
greater degree for early bilinguals than for late bilinguals, presum-
ably indicating that there is a greater co-activation of L2 orthogra-
phy with L1 orthographic processing in early bilinguals; (c) the
shallower-than-L1 L2s were related to greater activity in the bilat-
eral temporal regions and the right primary motor cortex, possibly
because the phonological assembly was more relied on for the
more transparent L2s, whereas the left frontal gyri were more acti-
vated in the processing of deeper-than-L1 L2s, probably to cope
with the increased arbitrariness of orthography-phonology map-
ping from L1 to L2. Taken together, the current study has extended
our understanding of how the L2 onset age and the relative trans-
parency of L2 to L1 shape the functional representations of the
bilingual brain.
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