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Distinct cortical areas associated with native and second languages [Abstract–Electronic version]

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The ability to acquire and use several languages selectively is a unique and essential human capacity. Here we investigate the fundamental question of how multiple languages are represented in a human brain. We applied functional magnetic resonance imaging (fMRI) to determine the spatial relationship between native and second languages in the human cortex, and show that within the frontal-lobe language-sensitive regions (Broca's area), second languages acquired in adulthood ('late' bilingual subjects) are spatially separated from native languages. However, when acquired during the early language acquisition stage of development ('early' bilingual subjects), native and second languages tend to be represented in common frontal cortical areas. In both late and early bilingual subjects, the temporal-lobe language-sensitive regions (Wernicke's area) also show effectively little or no separation of activity based on the age of language acquisition. This discovery of language-specific regions in Broca's area advances our understanding of the cortical representation that underlies multiple language functions.
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Acknowledgements. We thank D. Krakauer and K. Sigmund for discussion. This work was supported by
the Wellcome Trust (M.A.N.) and the European Community (M.C.B.).
Correspondence should be addressed to M.A.N. (e-mail: martin.nowak@zoo.ox.ac.uk).
Distinct cortical areas
associated with native
andsecondlanguages
Karl H. S. Kim*
, Norman R. Relkin
, Kyoung-Min Lee*
&
Joy Hirsch*
Department of Neurology, * Memorial Sloan-Kettering Cancer Center,
1275 York Avenue, New York, New York 10021, USA
Department of Neurology and Neuroscience, Cornell University Medical College,
1300 York Avenue, New York, New York 10021, USA
.........................................................................................................................
The ability to acquire and use several languages selectively is a
unique and essential human capacity. Here we investigate the
fundamental question of how multiple languages are represented
in a human brain. We applied functional magnetic resonance
imaging (fMRI) to determine the spatial relationship between
native and second languages in the human cortex, and show that
within the frontal-lobe language-sensitive regions (Brocas
area)
1–3
, second languages acquired in adulthood (‘late’ bilingual
subjects) are spatially separated from native languages. However,
when acquired during the early language acquisition stage of
development (‘early’ bilingual subjects), native and second lan-
guages tend to be represented in common frontal cortical areas. In
both late and early bilingual subjects, the temporal-lobe language-
sensitive regions (Wernicke’s area)
1–3
also show effectively little or
no separation of activity based on the age of language acquisition.
This discovery of language-specific regions in Broca’s area
advances our understanding of the cortical representation that
underlies multiple language functions.
Indirect evidence for topographic specialization within the
language-dominant hemispheres of multilingual subjects has been
provided by clinical reports of selective impairments in one or more
of several languages as a result of surgery involving the left
perisylvian area
4
. Multilingual patients with complex partial seizure
disorders of temporal lobe origin have been reported to shift from a
primary to a second language together with ictal progression
5
.
Different languages have also been selectively disrupted in polyglots
by electrical stimulation of discrete regions of the neocortex of the
dominant hemisphere
6,7
. Changes in the topography of background
electroencephalogram (EEG) coherence obtained during transla-
tion tasks also suggest spatial separation of cortical regions involved
in multiple languages
8
. Although these reports are consistent with
the existence of spatially separate representations for each language,
such functions have not been localized.
Silent, internally expressive linguistic tasks were performed in two
languages by subjects who either acquired conversational fluency in
their second languages as young adults (‘late’ bilinguals) or who
acquired two languages simultaneously early in their development
(‘early’ bilinguals) (Table 1). As Brocas and Wernicke’s areas are
known to perform central roles in human language functions
13,912
,
we have focused our observations on these cortical areas.
The main findings for a typical ‘late’ bilingual subject (subject
(A)) are shown in Fig. 1. The anterior language area is highlighted
by the green box and shown expanded in the inset. Red indicates
significant activity during the native language task (English),
whereas yellow indicates activity associated with the second lan-
guage task (French). Two distinct but adjacent centres of activation
(+) separated by ,7.9 mm were evident within the inferior frontal
gyrus, suggesting that two specific regions served each of the two
languages. In the posterior language area of the same subject (Fig.
2), the same tasks yielded centroids of activity with a centre-to-
centre spacing of 1.1 mm, less than the width of a voxel, suggesting
that similar or identical cortical regions served both languages in
this posterior area.
For all six late bilingual subjects, distinct areas of activation were
observed for the native and second languages in Brocas area (Table
2a and Fig. 3). The separation between centroids of activity ranged
from ,4.5 mm to 9.0 mm within one slice, and the number of voxels
for each language was similar for each subject. On the other hand,
activity in Wernicke’s area (Table 2b) showed centre-to-centre
distances between the centre-of-mass centroids ranging from 1.1
to 2.8 mm. The mean centroid distance between the anterior
Figure 1 A representative axial slice from a ‘late’ bilingual subject (A) shows all
voxels that pass the multistage statistical criteria at P # 0:0005 as either red
(native language) or yellow (second acquired language). An expanded view of the
pattern of activity in the region of interest (inferior frontal gyrus, Brodmann’s area
44 (refs 2, 3,18), corresponding to Brocas area
1–3
) indicates separate centroids (+)
of activity for the two languages. Centre-of-mass calculations indicate that the
centroids are separated on this plane by 7.9 mm. The green line on the upper right
mid-sagittal view indicates the plane location. R indicates the right side of the
brain.
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language areas was 6.43 (61.83) mm and exceeded that of the
posterior language areas, which was 1.88 (60.62) mm, for these
subjects (t ¼ 5:43, d:f: ¼ 5, P # 0:004).
The overall stability of the centre-of-mass centroids with varia-
tion in the level of statistical stringency (probability of a false-
positive result, P) is illustrated for subject (A) in Fig. 4. In the case of
the anterior language area, the centre-to-centre distance between
the centroids of activity associated with each language remained
within the range of 7.8 to 9.1 mm over statistical stringency levels
from P # 0:0002 to P # 0:02. In the case of the posterior language
area, the centroids remained within the approximate width of one
voxel, 1.6 mm, over the same range of stringency levels. Similar
results were obtained for all subjects, confirming that threshold
criteria do not account for the centre-to-centre distances between
centroids of activity associated with each language task.
Figure 5 illustrates the main findings for a typical ‘early’ bilingual
subject (subject (G)) for whom the centre-to-centre distance
between the activity centroids (+) of the two activity patterns was
2.3 mm, less than 1.5 voxels (Table 2a). This represents the general
pattern for all six early bilingual subjects, where the mean separation
was 1.53 (60.78) mm. In the case of the posterior language area,
Wernicke’s area, the mean separation for all early bilingual subjects
was 1.58 (60.79) mm, similar to the anterior area.
Our findings are summarized by an analysis of variance (Table 3)
in which language area (Broca’s and Wernicke’s) was compared with
bilingual type (early and late) with respect to the centre-to-centre
distance in millimetres between the two language centroids. Sig-
nificant main effects for language area (P # 0:000059) and bilingual
type (P # 0:000084) with an interaction effect (P # 0:000067) show
that activation sites for the two different languages tend to be
spatially distinct in Brocas area when the second language was
obtained late in life and not when acquired in early childhood; and
Table 1 Subject information
Subject Age Gender Native language(s) Second language Handedness Laterality quotient
24
...................................................................................................................................................................................................................................................................................................................................................................
A 31 M English French Right 60
B 32 M Korean English Right 100
C 28 M Korean English Right 86
D 26 M English Japanese Ambidextrous 20
E 27 F Spanish English Right 100
F 32 F German English Right 60
...................................................................................................................................................................................................................................................................................................................................................................
G 38 F Turkish/English NA Ambidextrous 27
H 27 M English/Hebrew NA Right 85
I 23 M English/Spanish NA Right 100
J 24 F Croatian/English NA Right 89
K 31 M Italian/German NA Right 90
L 32 M Chinese/English NA Ambidextrous 24
...................................................................................................................................................................................................................................................................................................................................................................
Figure 2 Similar to Fig. 1, an expanded view of the pattern of activity within the
superior temporal gyrus (Brodmanns area 22 (refs 2, 3, 18), corresponding to
Wernicke’s area
1–3
) indicates centroids of activity for the two languages in this
posterior language region. Orange indicates that the voxels that passed all
statistical criteria during both the native and acquired language tasks. Centre-
of-mass calculations indicate that the centroids are separated on this plane by
1.1 mm, less than the diameter of a single voxel.
Figure 3 Expanded views of the activity patterns within Brodman’s area 44 (and 46
(refs 2, 3,18), subject B) for each ‘late’ bilingual subjects (AF) indicate the active
regions during the native language task (red) and the second acquired language
task (yellow). The level of statistical stringency (probability of a false positive
result, P) was #0.0005 for each subject. The centre-of-mass is indicated by a plus
sign for each language; area and centre-to-centre (CC) distances are listed in
Table 2a.
Table 3 ANOVA
Source of variation SS d.f. MS F P
A (language area) 52.01 1 52.01 25.70 0.000059
B (bilingual type) 48.80 1 48.80 24.11 0.000084
AB 50.79 1 50.79 25.10 0.000067
Within cell 40.47 20 2.02
.............................................................................................................................................................................
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that Wernicke’s area showed little or no separation of activity
regardless of age of acquisition.
The observation that the anatomical separation of the two
languages in Broca’s area varies with the time at which the second
language was acquired, suggests that age of language acquisition
may be a significant factor in determining the functional organiza-
tion of this area in human brain. Human infants, initially capable of
discriminating all phonetically ‘relevant’differences, may eventually
modify the perceptual acoustic space, based on early and repeated
exposure to their native languages
13
. It is possible that representa-
tions of languages in Broca’s area that are developed by exposure
early in life are not subsequently modified. This could necessitate
the utilization of adjacent cortical areas for the second language
learned as an adult.
The difference between the results of this investigation and a
positron emission tomography (PET) study in which multiple
languages were found to generate overlapping regions of activation
within the inferior frontal gyrus
14
may be reconciled in part by the
higher effective resolution of this fMRI technique. The intrinsic
resolution of the PET H
2
15
O cerebral-blood-flow technique was
5 3 5 3 6 mm
3
and the results from several subjects were combined
and averaged. Individual variability in both the locations of the
language areas and the diversity of brain shapes and gyral patterns of
the subjects averaged together further reduce the effective resolution
of this approach
9,15
, which could account for the discrepancy.
However, on the basis of our findings, the distinction between
native and second languages may be less for younger ages of
exposure to a second language. The average age of initial exposure
to the second language in the PET study was 7.3 years, younger than
that of the late bilingual subjects in our study, and could therefore be
consistent with our observation for early bilinguals.
To render our findings independent of particular languages or
cultural background, our study made use of simple expressive tasks
with similar semantic content across multiple languages and
Table 2 Areas of activation
(a) Anterior language area
Talairach &Tournoux
sectors
18
*
Subject Bilingual type Hemisphere Gyrus
Brodmann’s
area X Y Z
Area L1
(mm
2
)
Area L2
(mm
2
)
CC distance
(mm)
Common voxels
L1 and L2
(%)
...................................................................................................................................................................................................................................................................................................................................................................
A Late Left Inferior frontal 44 d D 12 34.2 34.2 7.9 0
B Late Left Inferior frontal 44, 46 c D, C 12 14.6 12.2 4.5 0
C Late Left Inferior frontal 44 c D 12 12.2 14.6 9.0 0
C9 Late Left Inferior frontal 44 c D 12 15.4 24.2 7.5 0
D Late Left Inferior frontal 44 c B 1 12.2 17.1 4.7 0
E Late Left Inferior frontal 44 c D 12 12.2 12.2 7.0 0
F Late Left Inferior frontal 44 c D 12 33.1 19.8 11.2 0
...................................................................................................................................................................................................................................................................................................................................................................
G Early Left Interior frontal 44 c D 12 107.4 75.7 2.3 50
H Early Left Interior frontal 44 c D 12 12.2 7.3 0.5 60
I Early Left Interior frontal 44 c B 1 41.9 28.6 0.7 46
J Early Left Interior frontal 44 c D 12 8.8 4.4 1.7 20
K Early Left Interior frontal 44 c D 12 123.4 154.2 2.2 45
L Early Left Interior frontal 44 c D 12 44.1 17.6 1.8 22
...................................................................................................................................................................................................................................................................................................................................................................
(b) Posterior language area
A Late Left Superior temporal 22 d G 20 39.1 65.9 1.1 48
B Late Left Superior temporal 22 d G 20 9.8 19.5 1.5 33
C Late Left Superior temporal 22 d F 8 31.7 34.2 2.2 35
C9 Late Left Superior temporal 22 d G 20 19.8 33.1 3.3 26
D Late Left Superior temporal 22 d E 1 43.9 19.5 2.2 30
E Late Left Superior temporal 22 d F 8 22.0 9.8 1.5 18
F Late Left Superior temporal 22 d G 16 83.7 46.3 1.0 18
...................................................................................................................................................................................................................................................................................................................................................................
G Early Left Superior temporal 22 d G 16 4.9 31.7 1.7 15
H Early Left Superior temporal 22 d F 16 56.2 39.1 1.5 56
I Early Left Superior temporal 22 d E 4 94.7 22.0 3.0 23
J Early Left Superior temporal 22 d F 16 15.4 125.6 1.4 12
K Early Left Superior temporal 22 d G 20 121.2 96.9 1.3 62
L Early Left Superior temporal 22 d F 12 48.5 88.1 0.6 48
...................................................................................................................................................................................................................................................................................................................................................................
C9 is a replication of C after 16 months on a different scanner. All observations of area and distance are made at a common level of significants (P # 0:0005).
* See ref.18 where the three-dimensional proportional grid system is described. Columns X, Yand Z indicate coordinates of an orthogonal parallelogram, the dimensions of which vary with
the principal axes of the brain. Each of these volumes is defined by its three dimensions indicated by a capital letter (Y), a lower-case letter (X), and a number (Z).
L1 and L2 refer to the native and second acquired languages for late bilingual subjects, respectively, and the two languages acquired during the early developmental period for the early
bilingual subjects in the respective order as listed on Table 1.
CC distance refers to the centre-to-centre spacing between the centre-of-mass centroids of significant voxels associated with the two languages.
Figure 4 All voxels that pass the statistical criteria during the native (red) and
acquired (yellow) language tasks (subject (A)) are shown for the anterior region of
interest, left, and the posterior region of interest, right, over approximately 3 orders
of statistical stringency (P # 0:0002 to P # 0:02). The centre-to-centre (CC)
distance between the centroids of activity in the anterior area (based on the
centre-of-mass, +), ranges from 7.8 mm at P # 0:0002 to 9.1 mm at P # 0:02,
suggesting that the centroid is nearly independent of cluster size (radial expansion).
Similarly, for the posterior language area, the CC distance ranges from 1.1 to 1.3 mm
(less than the width of a single voxel) over the same level of statistical stringency.
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included subjects with various combinations of languages (Table 1).
Our findings are consistent with distinct roles for the anterior and
posterior language areas in the processing of human language, and
raise further questions regarding the role of Brocas area in proces-
sing the phonetic structures of different languages.
M
. .. . . . . . . . .. . . .. .. . . . . . . . . . .. .. .. . . . . . . . . . .. .. .. .. . . . . . . . .. . . .. .. . . . . . . . . . .. .. .. . . . . . . . . . .. .. .. .. . . . . . . . .. . . .. .. . . . . . . . . .
Methods
Imaging. A 1.5-tesla magnetic resonance scanner (General Electric) retrofitted
(advanced NMR Instascan) for echoplanar imaging and subsequently
upgraded to the GE echoplanar system was used to obtain T2*-weighted
images with a gradient echo pulse sequence (echo time, 60 ms; repetition time,
3,000 ms; flip angle, 308) which is sensitive to magnetic resonance signal
changes caused by alteration in the proportion of deoxyhaemoglobin in the
local vasculature accompanying neuronal activation
16
. Either a volume-
optimized 5 3 5-mesh dome resonator
17
or a General Electric head coil was
employed. The in-plane resolution was 1.6 mm by 1.6 mm. Slice thickness was
4.54.7 mm and 16 contiguous slices of brain were obtained parallel to a
reference line through the superior edge of the anterior commissure and the
inferior edge of the posterior commissure
18,19
. These slices covered the inferior
frontal gyrus (the anterior language region, ‘Brocas’ area including Brod-
mann’s areas 44 and 46) and the posterior superior temporal gyrus (the
posterior language region, ‘Wernicke’s’ area including Brodmanns area
22)
13,18,19
. Thirty images were taken, one every 3 s; thus, an entire run lasted
90 s. The first 10 images (30 s) were acquired during a baseline period, followed
by a stimulation or task period of 10 images (30 s), and a final (30 s) baseline
period also consisting of 10 images. A fixation cross-hair was provided during
the baseline epochs to help the subject to maintain a stable head position.
Analysis. Two identical runs were performed in each language. Before
statistical analysis, all brain images were computationally aligned to allow
direct spatial comparisons between different language tasks for individual
subjects
20
, and a two-dimensional gaussian filter (approximately 3 volume
elements, voxels, at half-height) was applied to the data. Significant signal
changes were identified by a multistage statistical analysis which compared
average baseline and stimulation signal intensities and required significant
signal changes on two runs (coincidence)
21,22
. The rate of false-positive voxels,
p, was empirically determined from images of a copper sulphate solution-filled
spherical phantom (General Electric standard) and found to be less than
0.0005. The centroid of a cluster of language-activated voxels was determined as
the two-dimensional centre of mass, and the centre-to-centre distance between
centroids was taken as the separation (mm) between language-specific activity.
Task. The sentence-generation task was performed silently (internal speech) to
minimize head movement and was similar to tasks previously employed in
neuroimaging language studies
23
. The subject was instructed to ‘‘describe’
events that occurred during a specified period of the previous day (morning,
afternoon, night); this task was practised before the imaging sessions. Imme-
diately before each run, the subject was instructed which language he/she was to
imagine speaking, and graphical cues signalling morning, afternoon, and night
were displayed in various orders for 10 s during the 30-s task period. These
graphics provided common non-linguistic cues for the task and the unpre-
dictable order of presentation presumably reduced the tendency to rehearse
mentally before the cue. The languages were alternated during the imaging
session to prevent habituation and a potentially time-dependent bias.
Subjects. Twelve healthy multilingual volunteers, 9 males and 3 females, were
recruited according to institutional informed consent procedures. Subjects
were either right-handed or ambidextrous, as assessed by the Edinburgh
handedness inventory
24
(Table 1). The mean age of subjects was 29.3 (64.2)
years. Six subjects (‘early’ bilinguals) were exposed to two languages during
infancy, and six subjects (‘late’ bilinguals) were exposed to a second language in
early adulthood. The mean age of initial exposure to the second language was
11.2 (61.5) years and the mean age that conversational fluency was achieved
was 19.2 (64.1) years. Each of the ‘late’ bilingual subjects had lived in the
country of the second language, which assured a high standard for fluency. Each
of the early bilinguals was raised in a home where either the parents spoke one
language and siblings and friends spoke another, or the parents spoke two
languages. Ten languages were represented as indicated on Table 1, and all
subjects reported approximately equal fluency and frequent usage in each
language at the time of testing.
Received 20 February; accepted 30 April 1997.
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Acknowledgements. We thank K. Zakian and D. Ballon for use of the 5 3 5 mesh dome resonator,
J. Victor, G. Krol, J. Posner, R. Cappiello, M. Ruge, D. Correa, S. Harris, J. Salvagno, P. Kuhl, F. Nottebohm,
G. E. Vates and R. DeLePaz for technical assistance and helpful comments, and Y. Popowich, N. Rubin,
T. Ozaki, D. R. Moreno, B. Aghazadeh, D. Barbut-Heinemann, D. Orbach, R. Valencia, J. Carton, E. Go
¨
tte,
R. Ha
¨
rtl, O. Torres and M. Li for volunteering as subjects. Supported by the William T. Morris Foundation
fellowship, the Tri-Institutional MD/Ph.D Program (KHSK); the Charles A. Dana Foundation, Johnson &
Johnson Focused Giving Foundation, Cancer Center Support Grant NCI (J.H.); the C. V. Starr
Foundation and the Lookout Fund (N.R.R.).
Correspondence and requests for materials should be addressed to J. Hirsch (e-mail: hirsch@vision.
mskcc.org).
Figure 5 A representative axial slice from an ‘early’ bilingual subject (G) who
learned English and Turkish simultaneously during early childhood shows all
voxels that pass the multistage statistical criteria at P # 0:0005. Red indicates the
Turkish language task and yellow indicates the English language task. An
expanded view of the region of interest (Brocas area
1–3
) indicates multiple
common voxels between the two language areas. The geometric centres-of-
mass indicate that the centroids are within 1.5 voxels. R indicates the right side of
the brain.
... Previous task-based fMRI studies have extensively investigated how the brain system supports L1 and L2 processing in bilinguals. These studies have found that the fronto-tempo-parietal functional network was largely shared for L1 and L2 (Abutalebi, 2008;Chee et al., 1999;Sulpizio et al., 2020), but differences in neural responses between languages were also observed, as indicated with either cortical activation (Abutalebi, 2008;Kim et al., 1997) or multivoxel patterns (Nichols et al., 2021;Ou et al., 2020;Xu et al., 2017). Moreover, researchers have investigated inter-regional functional connectivity and topological properties, and they revealed that a more globally integrated network emerged during L1 processing, while a more segregated network supported L2 processing (Cao et al., 2014;Dodel et al., 2005;Feng et al., 2015). ...
... On the one hand, distinct brain state dynamics in late bilinguals may result from differences in the acquisition and proficiency of L1 compared to L2, as well as their long-term effects on tuning functional networks in late bilinguals. Previous studies showed that the activation or multivoxel pattern tended to be more different between L1 and L2 conditions in late bilinguals, while they tended to be more similar in early bilinguals (Kim et al., 1997;Nichols et al., 2021;Nichols and Joanisse, 2016;Wartenburger et al., 2003). In addition, proficiency in L2 could also predict dissimilarity between language representations in several brain areas within the language network (Nichols et al., 2021;Nichols and Joanisse, 2016;Wartenburger et al., 2003). ...
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