Beruflich Dokumente
Kultur Dokumente
School of Psychology, University of Kent, Canterbury, Kent CT2 7NP, United Kingdom; and Departments of cPsychology and dClinical Language Sciences and
Centre for Integrative Neuroscience and Neurodynamics, School of Psychology and Clinical Language Sciences, University of Reading, Reading RG6 6AL,
United Kingdom
bilingualism
TBSS
www.pnas.org/cgi/doi/10.1073/pnas.1414183112
NEUROSCIENCE
PSYCHOLOGICAL AND
COGNITIVE SCIENCES
Edited by Patricia K. Kuhl, University of Washington, Seattle, WA, and approved December 12, 2014 (received for review July 29, 2014)
PSYCHOLOGICAL AND
COGNITIVE SCIENCES
values in the right IFOF and Mohades and colleagues (11) in the
left IFOF for bilinguals vs. monolinguals. Because this bundle
has been implicated in semantic processing (12), the observed
effect may signify more efficient semantic processing in bilinguals, which can be related to the fact that bilinguals need to
constantly select between L1 and L2 naming alternatives (19).
Another region that was found to be affected by bilingualism is
the genu of the CC, also reported in Luk and colleagues (5).
Schlegel and colleagues (22) also reported increased FA values
in the genu of the CC for their participants in intensive language
training courses, which they attributed to increased myelination
of the CC fibers as a result of increased language switching
demands. Although the role of the CC in language processing is
not fully understood (22), it has been heavily implicated in effective interhemispheric communication and in executive functioning (25, 26). The explanation provided by Schlegel and
colleagues also applies to the bilingual participants in our study,
as well as in Luk and colleagues, with the difference being that
our participants face similar switching demands as a result of
linguistic immersion, and not language training.
The final two tracts that appeared to be affected by bilingualism
in our study, as well as in Luk and colleagues (5), were the superior longitudinal fasciculus and uncinate fasciculus, bilaterally.
According to Friederici (27), these tracts constitute, respectively,
a dorsal and a ventral pathway connecting Brocas area to temporal areas, notably the superior temporal gyrus and the middle
temporal gyrus, which have been linked to phonological, semantic,
and syntactic processing in various, and sometimes competing,
theoretical models (see ref. 27 for a review). Differentiating the
exact involvement of these WM pathways in language acquisition
and in language processing is beyond the scope of this article and
cannot be achieved with our current dataset; however, effective
Pliatsikas et al.
connectivity between the same areas has also been recently proposed in early bilinguals (15).
Despite the proposed links between WM structure and cognitive benefits in bilingualism, these hypotheses have not been
tested systematically. Cummine and Boliek (24), for example,
reported a negative correlation between FA values and reaction
times in a task involving reading aloud words with regular and
irregular phonology. Similarly, Hosoda and colleagues (23)
showed a positive correlation between FA values and L2 vocabulary size in several WM tracts. Given the significant effects
we found for our L2 group in the language-related WM tracts
described here, we would also expect significant correlations
between the FA values in the reported tracts and the participants performance in tasks tapping on semantics and syntax,
signifying more efficient processing. In addition, the significant
effects on the CC should predict enhanced executive functioning
by the same group, as has also been proposed in ref. 1, highlighting further the effects of bilingualism on general cognition.
However, as the appropriate behavioral tasks were not administered in our groups, we can only speculate about these effects.
It is important to highlight that our young late bilingual group
demonstrated increased FA values in a widespread network of
language-related WM tracts. This network resembles the networks identified in older bilinguals (5), as well as those in early
bilinguals (15). In addition, and in contrast to language-training
studies (22, 23), the WM effects persevere as a function of everyday use of L2, a suggestion that can account for the effects
reported in Luk and colleagues, and can also explain the absence
of significant effects in the linguistically trained but not immersed late learners (23, 24). The close mapping of the present
results to those from Luk and colleagues suggests that everyday
handling of more than one language functions as an intensive
cognitive stimulation that benefits specific language-related
PNAS Early Edition | 3 of 4
NEUROSCIENCE
Fig. 1. Significant L2 > NS differences in FA values (red/yellow), expressed in 1-P values (P < 0.05, corrected) and overlaid onto a standard space WM skeleton (green).
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Curr Dir Psychol Sci 19:1923.
2. Schweizer TA, Ware J, Fischer CE, Craik FIM, Bialystok E (2012) Bilingualism as a contributor to cognitive reserve: Evidence from brain atrophy in Alzheimers disease.
Cortex 48(8):991996.
3. Li P, Legault J, Litcofsky KA (2014) Neuroplasticity as a function of second language
learning: Anatomical changes in the human brain. Cortex 58:301324.
4. Stein M, Winkler C, Kaiser A, Dierks T (2014) Structural brain changes related to bilingualism: Does immersion make a difference? Front Psychol 5:1116.
5. Luk G, Bialystok E, Craik FIM, Grady CL (2011) Lifelong bilingualism maintains white
matter integrity in older adults. J Neurosci 31(46):1680816813.
6. Smith SM, et al. (2006) Tract-based spatial statistics: Voxelwise analysis of multisubject diffusion data. Neuroimage 31(4):14871505.
7. Gold BT, Johnson NF, Powell DK (2013) Lifelong bilingualism contributes to cognitive
reserve against white matter integrity declines in aging. Neuropsychologia 51(13):
28412846.
8. Gold BT, Kim C, Johnson NF, Kryscio RJ, Smith CD (2013) Lifelong bilingualism
maintains neural efficiency for cognitive control in aging. J Neurosci 33(2):387396.
9. Pliatsikas C, Marinis T (2013) Processing of regular and irregular past tense morphology in highly proficient L2 learners of English: A self-paced reading study. Appl
Psycholinguist 34:943970.
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between bilingual and monolingual children. Brain Res 1435:7280.
12. Leclercq D, et al. (2010) Comparison of diffusion tensor imaging tractography of language tracts and intraoperative subcortical stimulations. J Neurosurg 112(3):503511.
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(2014) Anatomical connectivity changes in the bilingual brain. Neuroimage 84:495504.
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4 of 4 | www.pnas.org/cgi/doi/10.1073/pnas.1414183112
Twenty-five native speakers of English (female, 14; mean age, 28.16 y; SD,
5.33 y) formed the NS group of this study. No one from the NS group reported
speaking an L2. The demographics of the NS group are illustrated in Table S2.
The NS group was also tested in the QPT test (mean score, 98.56; SD, 4),
where they scored significantly higher than the L2 group [F(1,44) = 37.324;
P < 0.001], and in the Backward Digit Span test (mean score, 8.12; SD, 2.14),
where their performance did not differ from that of the L2 group [F(1,44) =
1.307; P = 0.259].
Data Acquisition, Preparation, and Analysis. We used a 3.0-Tesla Siemens
MAGNETOM Trio MRI scanner with Syngo software and 36-channel Head
Matrix coil to acquire a whole-brain diffusion-weighted Echo-Plannar Imaging image (two averages, 30 directions, 60 axial slices; slice thickness, 2 mm,
no interslice gap; field of view, 256 256 mm; acquisition matrix, 128 128;
voxel size, 2 mm isotropic; echo time, 93 ms; repetition time, 8,200 ms;
b-value, 1,000 s/mm2).
The images were preprocessed with FMRIBs Diffusion Toolbox, part of FSL
(31), and were initially corrected for eddy-current distortion and head motion, and subsequently a diffusion tensor model was fit at each voxel of the
corrected data with DTIFIT. This resulted in one FA image per participant.
Preprocessing was followed by a TBSS (6) analysis in FSL as follows: The FA
images for all participants were nonlinearly registered to a standard space
FA target image and subsequently affine-transformed into the 1 1 1
Montreal Neurological Institute 152 space in a combined process avoiding
sampling the images twice. This resulted in standard-space FA images for
each participant that were merged and averaged to create a mean 4D image. The mean image was fed into the FA skeletonization program to create
an FA skeleton that included voxels identified as WM across all participants.
The mean skeleton image was thresholded at 0.2. We then analyzed the
skeletonized data with a between-groups voxel-wise analysis with permutation-based nonparametric testing, corrected for multiple comparisons
with TFCE (32). Sex was included in the model as a covariate of no interest,
and two contrasts were examined: L2 > NS, and NS > L2. This resulted in
corrected whole-brain t statistical images of the significant differences,
which were thresholded at P < 0.05.
ACKNOWLEDGMENTS. We thank Francesca Allerton for assisting in the
recruitment of participants for this project. This study was internally funded
by the Centre for Integrative Neuroscience and Neurodynamics, University
of Reading.
Pliatsikas et al.
Supporting Information
Pliatsikas et al. 10.1073/pnas.1414183112
Table S1. Demographics of the L2 group
ID
Age, y
Sex
Country of
origin
Native
language
Highest obtained
educational level
101
103
105
106
107
108
109
110
111
112
113
114
115
116
119
120
121
123
124
126
Mean
SD
31
31
30
24
23
34
27
47
20
35
34
31
37
20
27
23
46
41
43
33
31.85
8.07
Male
Male
Female
Female
Female
Female
Female
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
Female
Male
Male
Greece
Italy
China
Poland
Greece
Serbia
France
Hong Kong
Bulgaria
Netherlands
Poland
France
Poland
Romania
Romania
France
Germany
Spain
Uganda
Italy
Greek
Italian
Mandarin
Polish
Greek
Serbian
French
Cantonese
Bulgarian
Dutch
Polish
French
Polish
Romanian
Romanian
French
German
Spanish
German
Italian
PhD
PhD
PhD
MSc/MA
BSc/BA
PhD
MSc/MA
BSc/BA
Undergraduate
PhD
MSc/MA
Undergraduate
BSc/BA
Undergraduate
MSc/MA
BSc/BA
MSc/MA
BSc/BA
PhD
PhD
U.K. residency, mo
Age of acquisition
of L2 English, y
99
86
88
18
40
44
34
374
13
102
114
144
92
16
42
44
175
196
81
18
91.00
84.87
8
15
12
6
8
6
12
6
10
11
4
15
18
13
5
13
11
4
10
16
10.15
4.17
Place of L2 learning
School
School
School
School
Home and school
Home and school
School
School
School
School
School
School
School
Home and school
School
School
School
School
School
School
Age, y
Sex
Highest obtained
educational level
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
Mean
SD
23
24
23
28
30
24
26
32
25
24
23
22
22
31
39
33
37
34
31
32
23
34
33
31
20
28.16
5.33
Male
Male
Male
Female
Female
Female
Male
Male
Male
Male
Female
Female
Female
Male
Female
Male
Male
Female
Female
Female
Male
Female
Female
Female
Female
BSc/BA
MSc/MA
BSc/BA
MSc/MA
PhD
MSc/MA
PhD
PhD
MSc/MA
BSc/BA
MSc/MA
MSc/MA
MSc/MA
PhD
BSc/BA
MSc/MA
PhD
MSc/MA
BSc/BA
PhD
BSc/BA
MSc/MA
PhD
Undergraduate
BSc/BA
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