Content uploaded by Augusto Buchweitz
Author content
All content in this area was uploaded by Augusto Buchweitz on Sep 06, 2017
Content may be subject to copyright.
Japanese and English sentence reading comprehension and
writing systems: An fMRI study of first and second language
effects on brain activation
Augusto Buchweitz,
Carnegie Mellon University
Robert A. Mason,
Carnegie Mellon University
Mihoko Hasegawa, and
RIKEN Brain Science Institute, Japan
Marcel A. Just
Carnegie Mellon University
Abstract
Functional magnetic resonance imaging (fMRI) was used to compare brain activation from Japanese
readers reading hiragana (syllabic) and kanji (logographic) sentences, and English as a second
language (L2). Kanji showed more activation than hiragana in right-hemisphere occipito-temporal
lobe areas associated with visuospatial processing; hiragana, in turn, showed more activation than
kanji in areas of the brain associated with phonological processing. L1 results underscore the
difference in visuospatial and phonological processing demands between the systems. Reading in
English as compared to either of the Japanese systems showed more activation in inferior frontal
gyrus, medial frontal gyrus, and angular gyrus. The additional activation in English in these areas
may have been associated with an increased cognitive demand for phonological processing and verbal
working memory. More generally, L2 results suggest more effortful reading comprehension
processes. The study contributes to the understanding of differential brain responses to different
writing systems and to reading comprehension in a second language.
Introduction
The objective of the study was to investigate the brain activation associated with reading
comprehension and different orthographies in two Japanese writing systems, and in English,
a second language. A remarkable characteristic of the Japanese writing systems (kanji,
katakana, and hiragana) is that they present readers with the cognitive challenge of decoding
different types of mappings of words to sound and to meaning. In the study, participants read
kanji sentences and hiragana sentences, their native language (L1), as well as English sentences,
their second language (L2). The investigation of Japanese readers of English as a second
language allows for a unique three-faceted comparison of reading different writing systems,
with the L1 being written in a logographic system and in a non-alphabetic, syllabary system,
and the L2 in an alphabetic, syllabic system.
Address for correspondence: Center for Cognitive Brain Imaging, Carnegie Mellon University, Department of Psychology, 5000 Forbes
Avenue, Pittsburgh, PA 15213 United States.
NIH Public Access
Author Manuscript
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
Published in final edited form as:
Biling (Camb Engl). 2009 January 28; 12: 141–151. doi:10.1017/S1366728908003970.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Two hypotheses motivated this investigation. First, there should be differences in brain
activation associated with the processing of Japanese kanji and Japanese hiragana writing
systems. Even though kanji and hiragana are orthographies of the same language, the different
mapping of print to phonology and to meaning requires readers to rely on different cognitive
processes. Second, there should be differences in brain activation associated with the
processing of English compared with Japanese. The L2-L1 differences are expected to reflect
additional cognitive processes of reading in a foreign language, and in a different writing
system in which the readers are relatively less proficient.
Logographic and syllabic writing systems and their brain representations
The brain activation associated with reading syllabic and logographic systems has both
universal and orthography-specific characteristics. Syllabic systems have print mapped to
sound at the level of a single letter and of letter combinations (graphemes) with varying degrees
of regularity between languages (assembled phonology (Tan, Laird, Li, & Fox, 2005a).
Logographic systems have print mapped to sound at the level of meaning (morphemes)
(morpho-syllabic systems (Perfetti et al., 2007)). Reading logographic characters requires
processing of visual and spatial information to access the mental lexicon and to access the
pertinent lexical and phonological information to sound-out a word (addressed phonology
(Tan et al., 2005a)).
The dual-coding model (Yamadori, 2000; Thuy et al., 2004) proposes a representation of the
different cognitive processes of processing ideographic and syllabic systems in Japanese, i.e.,
kanji and katakana (kana), respectively. According to the model, the processing of kanji
requires that readers access word semantics first (semantics-first system) during reading. Kana,
in its turn, requires that readers access the phonology of the words first (phonology-first system)
(Yamadori, 2000). Meaning in kanji is intrinsic, or irregular, as determined by sound. Kana
has definite phonetic values, and its syllabograms are unique (an extreme form of regular
words) (Thuy et al., 2004).
Two companion meta-analyses by Bolger, Perfetti, and Schneider (2005) and Tan et al.
(2005a) reviewed neuroimaging studies of word-reading in Western European languages, and
in Chinese logographic, Japanese katakana (kana), and Japanese kanji. There were three
regions of robust convergence of brain activation from different orthographies: (1) mid-anterior
aspect of the left superior temporal lobe; (2) left inferior frontal gyrus (LIFG); and (3) left
occipito-temporal region (Bolger et al., 2005). The left occipito-temporal network (including
the left visual word form area (Cohen & Dehaene, 2004)) is postulated as an area that is
universally responsible for feedback between phonology and print.
The diverging loci of activation accommodate some of the variances between the printed forms.
Right hemisphere occipital and temporal lobe areas were associated with visuospatial
processing of logographic systems (Matsuo et al., 2003; Tan et al., 2005a). Left hemisphere
temporal and parietal areas were associated with phonological processing of alphabets. Tan et
al. (2005a) found differences in: (a) posterior aspect of the superior temporal lobe activated
only in syllabic systems (alphabetic and non-alphabetic); this activation is possibly associated
with the retrieval of phonological information (Tan et al., 2005a); (b) left anterior-dorsal frontal
region recruited in Chinese character reading; this activation may be associated with the
coordination of the processing of the unique combination of graphic, phonological, and
semantic information in logographic characters; (c) right occipito-temporal region (Brodmann
Area 37) activated in reading Chinese logographs compared with syllabic languages; this
activation is possibly associated with spatial processing of logographs (Bolger et al., 2005). In
a brain imaging study of European languages, Paulesu et al. (2000) found differences in brain
activation associated with the regularity of Italian, and the irregularity of English orthographies.
Buchweitz et al. Page 2
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Bolger et al.’s (2005) meta-analysis reported common right hemisphere occipito-temporal
areas for reading Chinese logographs. This activation was not found in the meta-image
generated for the studies of reading kanji logographs. The absence of right occipito-temporal
activation for reading kanji may have been an effect of the averaging over results in the meta-
analysis. One of the studies included in the meta-analysis did report right medial occipital
activation for Japanese kanji (Nakamura et al., 2000). The authors of the study argued that the
right occipital activation was a result of the visual processing differences between reading kanji
words and reading single kana characters. Other studies have also reported more activation of
bilateral fusiform areas (BA 37) for kanji (Thuy et al., 2004; Nakamura, Dehaene, Jobert,
Bihan, & Kouider, 2005).
The unique right occipito-temporal activation in reading kanji is evidence that there are
additional visual processes associated with reading this type of orthography. The distinctive
visual processing demands of reading logographs are also mirrored developmentally. Learning
to read the Chinese logographic system depends on developing good handwriting skills (Tan,
Spinks, Eden, Perfetti, & Siok, 2005b), and on the ability to integrate visuospatial information
(Tan et al., 2001).
Clinical studies have also shown that visual and spatial processes are fundamental for reading
kanji characters. Sakurai et al. (2000), in an investigation of alexia and agraphia in Japanese,
reported the case of a fusiform lesion patient who had an equally impaired ability to understand
written kanji and kana (alexia), but who made more paragraphic errors for kanji, had minor
agraphia for kanji, and could write kanji that he could not read. The patient used kinesthetic
reading (or facilitation) as a strategy to overcome kanji reading difficulties (Sakurai et al.,
2000). The importance of kinesthetic reading for processing logographic characters was also
shown by Matsuo et al. (2003), who reported that finger movements lightened the cognitive
load of reading logographic characters.
Evidently, Chinese and Japanese logographs are systems of two different languages. But
Chinese and Japanese logographs do have a similarity that is of interest for the study, that is,
they are morpho-syllabic writing systems. The similarity between Japanese kanji and Chinese
logographs has historical origins: kanji characters are originally Chinese characters. The review
of brain imaging studies of reading Chinese logographs is, in our understanding, pertinent to
the discussion of brain representations for reading kanji logographs.
In the present study, it was expected that there would be differences in brain activation
associated with the unique characteristics of Japanese kanji and hiragana writing systems. The
different mapping of print to phonology and meaning elicits different cognitive processes for
reading in the two systems. The different processes for reading hiragana and kanji were
expected to be associated with differences in brain activation. To our knowledge, there are no
previous studies that contrasted the brain activation from reading kanji and hiragana sentences.
Brain representation of language in bilinguals
Brain activation in bilinguals varies with the specific processing demands of each language,
as well as with proficiency, age of acquisition (Perani & Abutalebi, 2005) and writing systems
(Meschyan & Hernandez, 2006). Brain imaging studies of bilinguals have been carried out on
the cortical representation of different language-pairs (Kim, Relkin, Lee, & Hirsch, 1997); on
bilingual auditory sentence comprehension (Hasegawa, Carpenter, & Just, 2002); bilingual
reading comprehension (Meschyan & Hernandez, 2006), and on bilingual phonetic, lexical,
and phonological processing (Marian, Spivey, & Hirsch, 2003; Tham et al., 2005). In a
precursor to this paper, Hasegawa et al. (2002) showed that the workload of listening
comprehension in the less-proficient L2 resulted in activation of additional cortical areas in
comparison with L1. Meschyan and Hernandez (2006) reported increased cortical activity
Buchweitz et al. Page 3
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
associated with differences in the orthographic transparency of Spanish (shallow orthography)
and English (deep orthography).
Participants were late bilinguals and self-rated their reading skill in L2 at an intermediate level
of proficiency (see Methods section). Few studies (Chee et al., 1999; Shieh et al., 2001;
Yokoyama et al., 2006) have investigated bilinguals reading logographic and alphabetic
systems at a level above that of words. Yokoyama et al. (2006) contrasted the brain activation
associated with reading in Japanese and English. The study focused on the differences between
processing structurally complex sentences (active and passive sentences) in the two languages.
Others have investigated listening comprehension effects in the English-Japanese language
pair (Hasegawa et al., 2002), and the Chinese-English and Korean-English language pairs
(Jeong et al., 2007). It was expected that the results of the present study would provide evidence
of an increased workload in L2 sentence reading comprehension in a different writing system
(English) in which participants are less proficient than in the L1.
Method
Participants
Ten right-handed (Edinburgh handedness inventory (Oldfield, 1971)) bilingual speakers of
Japanese (first language) and English participated in the study. The results are reported for nine
participants (age M=27.4 years, SD=4.27, range 24–38 years). One participant was excluded
due to excessive head movement greater than 3.0 mm. All gave signed, informed consent
approved by the University of Pittsburgh and Carnegie Mellon Institutional Review boards.
Participants in this study were the same as those from a previous study (Hasegawa et al.,
2002).
Participants were enrolled as university undergraduate or graduate students in the United
States. At the time of the study, they had been in the US for 0.5 to 3 years (M=1.13; SD=0.8).
Age of acquisition of the second language showed that participants were late bilinguals, as
defined by Paradis (2003) (second language learned after the age of seven years) (M=26.9,
SD=4.8, range 21–38). Proficiency in the L2 was assessed using a language background
questionnaire (Hasegawa et al., 2002), in which bilinguals were asked to self-rate their reading
ability in English. The rating was on a scale of 1.0–5.0 with 0.5 intervals, where 5.0 was
excellent and 1.0 poor. Overall, participants rated their proficiency at an intermediate level
(M=3.0; SD=0.9).
Materials
Two-clause target sentences were presented in English, and Japanese hiragana and kanji scripts
(see Figure 1 for examples). The kanji sentences had mixed in katakana and hiragana characters,
but still were mostly kanji. This mixed kanji text is the typical manner in which modern
Japanese logograph is written. Kana characters are used together with kanji to, for example,
transcribe neologisms. Japanese sentences written only with hiragana, however, are unusual.
Sentences in hiragana were included to focus on the brain activation associated with reading
of L1 syllabograms. Sentences had either a negative statement, for example: The uncle didn’t
take out a notebook and lent an umbrella to the doctor, or an affirmative statement: The worker
read a magazine and showed some pictures to the brother. Half the sentences in the experiment
stimuli were negative, half positive.
To assess L1 and L2 comprehension accuracy, and to ensure that participants were performing
the comprehension task (rather than just superficially reading the sentences), a single-clause
probe sentence followed the target sentence, to which participants had to respond true or false.
The probe sentence always referred to the agent-patient relationship in the sentence. For
Buchweitz et al. Page 4
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
example, the negative sentence above was followed by the probe The uncle lent an umbrella.
Forty percent of the probes were false. The nouns and verbs for English sentences were chosen
to be at beginner or lower intermediate levels (junior high school level), and therefore would
be familiar to the participants. In the precursor to this study, English sentences were found to
be quite comprehensible to native speakers (Hasegawa et al., 2002). Translation equivalents
were used to make the lexical level of sentences comparable.
Procedure
There were three separate fMRI scans, one for each writing system. The presentation order of
the conditions was balanced across participants. In all cases, sentences were displayed in full
on a screen, one at a time, using Coglab Experimental Control software (www.ccbi.cmu.edu).
Participants were familiarized with the scanner before the actual study. The experiment was
self-paced; the display of a sentence was terminated by the participant clicking a mouse button.
The first sentence was then followed by a probe sentence. Display of the probe sentence was
terminated by the participant making a true or false response, also by clicking mouse buttons.
The durations of each of the two instances of sentence presentation were labeled target-sentence
reading time and probe-sentence reading time. There was no feedback to the participant
following response to the probe sentence. Target and probe sentences were separated by 500
ms, and an additional 1500-ms interval followed the probe sentence before display of the next
target-probe pair.
Each of the three conditions consisted of 8 epochs; each epoch had a series of five successively
presented trials of either affirmative or negative target sentences. Participants were given one
epoch of affirmative sentences, then one epoch of negative sentences. Two additional epochs
were presented to four participants who showed considerably fast reading times in both kanji
and hiragana during the practice session. The additional epochs were presented to ensure the
acquisition of at least 40 images per participant for each Japanese condition. Sentence epochs
were alternated with fixation periods of either six or 30 seconds. The 30-second fixation periods
were inserted after every two epochs. The shorter six-second fixation was included to allow
the hemodynamic response to decrease between the epochs that were not followed by a 30-
second fixation, before the next epoch began. Fixation consisted of the display of a crosshair
in the middle of the screen.
Image Acquisition
Images for all three scans were acquired on a 1.5-T scanner (with quadrature birdcage head
coil) at the MR Research Center at the University of Pittsburgh Medical Center. The acquisition
parameters for gradient-echo EPI with 14 oblique axial slices were TR=3000 ms, TE=50 ms,
flip angle 90°, 128 × 64 acquisition matrix, FOV 40 × 20 cm, 5-mm slice thickness, 1-mm gap,
in-plane voxel resolution of 3.125 × 3.125 mm, and RF whole-head coil. Four disabled
acquisitions preceded each scan.
fMRI data analysis
To compare the distribution of activation, data were analyzed using SPM99 (Wellcome Dept.
of Cognitive Neurology, University College London). Images were corrected for slice
acquisition timing, motion-corrected, normalized to the Montreal Neurological Institute (MNI)
template, resampled to 2 × 2 × 2 mm voxels, and smoothed with an 8-mm Gaussian kernel to
decrease spatial noise. The model for each participant included regressors for each of the six
conditions scanned (kanji negative and positive, hiragana negative and positive, English
negative and positive) and for the fixation condition. The regressors were convolved with the
canonical SPM99 hemodynamic response function. The length of the regressors was the
individual box car function convolved with each subject’s self-paced reading time for each
sentence in the eight or ten-epoch blocks.
Buchweitz et al. Page 5
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Statistical analysis was performed on individual and group data by using the general linear
model and Gaussian random field theory as implemented in SPM99 (Friston et al., 1995). The
extent threshold was 20 voxels. Group analyses were performed using a random-effects model.
Statistical maps were superimposed on normalized T1-weighted images. Group images were
generated for each condition relative to fixation (the common baseline condition). After group
analyses were carried out, we collapsed across the images for negative and affirmative
sentences relative to fixation in each writing system, Japanese hiragana, Japanese kanji, and
English. Images were collapsed across the two types of sentences because there were no
systematic effects of negative versus affirmative sentences. The collapsed images for each
individual subject were used to carry out the condition subtractions in SPM99: kanji > hiragana;
hiragana > kanji; English > hiragana; and English > kanji (paired t-test; p=.005, uncorrected).
An additional direct contrast was carried out between English and the average of the two
Japanese conditions. We further collapsed the images for each individual subject across the
two Japanese writing system conditions by generating an average contrast file for each
individual subject, and the collapsed images were subsequently used to carry out the L2 > L1
subtraction in SPM99: English > Japanese (average of hiragana and kanji) (paired t-test; p=.
005, uncorrected). Due to the more modest number of participants, a more liberal height
threshold of p=.005, uncorrected, was applied in the analysis of fMRI results. The activation
clusters that remained significant (p<.05) after cluster-level correction for multiple
representations are marked with an asterisk (*) in the tables of results.
Results and Discussion
Behavioral results
Comparison of target and probe-sentence reading times across languages showed that L2
reading comprehension was significantly more time-consuming than L1 reading
comprehension. Total target sentence reading time comparison: English versus kanji, t(8)
=4.74, p<.05; English versus hiragana, t(8)=3.75, p<.05. Probe-sentence reading time: English
versus kanji, t(8)=3.98, p<.05; English versus hiragana, t(8)=2.80, p<.05. Comprehension
accuracy was not affected in L2 (there were no significant differences in the accuracy of the
responses in the comparison between Japanese writing systems and English). The bilinguals
were slower to read in English, but were still able to understand the sentences (hiragana error
M=0.11; SD=0.08; kanji error M=0.10; SD=0.07; English error M=0.10; SD=0.07). There
were no differences in reading times or accuracy between the two Japanese writing systems
(kanji and hiragana). English and Japanese target- and probe-sentence reading times are shown
in Table 1.
fMRI results
Kanji and hiragana—The direct contrasts between the Japanese writing systems indicate
that the brain activation reflects the differences in the mapping of print to sound and to meaning
in the two orthographies. The contrast between kanji and hiragana shows more activation for
kanji in right inferior (BA 37) and mid temporal gyri (BA 39). The cortical response to kanji
sentences may be associated with the visuospatial processing associated with logographic
characters. More activation of right BA 37 for kanji corroborates previous studies that reported
activation in this area for reading logographs (Nakamura et al., 2005; Tan et al. 2005a). The
activation of right hemisphere areas in kanji logograph reading can be associated with the
specialization of the right side of the brain for pictorial and spatial processing. This
interpretation of the results is supported by the findings of imaging, developmental, and clinical
studies of reading logographs (Bolger et al., 2005; Tan et al., 2005a; 2005b; Matsuo et al.,
2003; Tan et al., 2001; Sakurai et al., 2000). Figure 2 and Table 2 report the results for the
comparison of kanji with hiragana.
Buchweitz et al. Page 6
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
The contrast between hiragana and kanji shows more activation for hiragana in left inferior
parietal lobe (LIPL), right supramarginal gyrus, and right supplementary motor area (SMA).
The LIPL activation indicates that there is an additional demand on phonological coding of
words in hiragana, relative to kanji. The activation of right-hemisphere supramarginal gyrus
may be a spill-over of activation from the left-hemisphere supramarginal area, which is
associated with phonological processing. In language tasks, the additional, less-specialized
right-hemisphere area may be recruited because it is capable of performing similar types of
cognitive functions to the first, well-specialized area (Just, Carpenter, Keller, Eddy, &
Thulborn, 1996). Figure 3 and Table 2 report the activation of hiragana compared with kanji.
Reading a sentence requires comprehension processes that are not in effect at the word level,
such as integration of sentence information and inference-making. Thus, the brain activation
differences associated with single word reading in different writing systems could have been
less evident at the level of sentence reading. But direct comparison of brain activation from
comprehending sentences in the two Japanese writing systems shows activation that mirrors
differences at the level of processing logographic and syllabic words. The results for the
contrast between kanji and hiragana corroborate the dual-coding model of different routes to
meaning in processing logographs and syllables (Yamadori, 2000; Thuy et al., 2004).
English > hiragana and English > kanji—The L2-L1 comparisons show recruitment of
cortical areas that may be associated with the greater workload imposed by reading
comprehension processes in the less-proficient L2. The results show systematic brain activation
in the L2 in relation to each of the L1 writing systems. English activated more left angular,
bilateral superior frontal, and left inferior frontal gyri in comparison with kanji, and in
comparison with hiragana. The differences between L2 and L1 reading may be associated with
the additional demand on internal phonological rehearsal processes and verbal working
memory in reading a second-language, alphabetic system. English orthography is
characteristically irregular (deep orthography) in its mapping of print to sound; it possibly
places additional cognitive demand on phonological processes in comparison with either first-
language system. The longer reading times in English might have been expected to produce
more activation in the visual cortex (longer duration of visual input on the screen), but no such
differences emerged.
English showed more activation than hiragana in a cluster surrounding the left angular gyrus
and extending anteriorly to the LIPL, which was mirrored laterally in the right supramarginal
gyrus. In the comparison with kanji, not only did the extra activation of left angular gyrus in
English extend anteriorly to the left supramarginal gyrus, but it was also mirrored laterally in
the right hemisphere (right angular and supramarginal gyri). A similar activation in the left
supramarginal gyrus (BA 40) has been reported in association with phonological processing
in English versus Chinese logographs (Tham et al., 2005). Figure 4 shows the activation from
reading English compared with reading Japanese hiragana and kanji (see also Tables 3 and 4).
Studies of reading syllabic systems in a native language show that the angular gyrus is one of
the areas associated with phonological processes and graphophoneme conversion (Bolger et
al., 2005). In the present study, activation of the angular gyrus was found in both contrasts of
English with Japanese. Activation of the angular gyrus was also found in the analysis of English
> Japanese (average of kanji + hiragana) (Table 5 and bottom rendering in Figure 4). The
consistent activation of the angular gyrus may be associated with the cognitive load of
phonological processes of reading an alphabetic system in a second language.
The peaks for the two clusters of angular gyrus activation in the English > kanji and the English
> hiragana contrasts are relatively close (approximately 18 mm in both cases) to the left inferior
parietal region where Mechelli et al. (2004) reported an increase in the density of grey matter
Buchweitz et al. Page 7
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
for both early and late bilinguals. It is possible that the cortical region that includes the left
inferior parietal lobe and the angular gyrus plays a role in the access to L2 phonology.
The activations of the left inferior frontal, of the superior frontal, and of the angular gyri form
a network of areas that were consistently more activated in L2. The activation of this network
suggests that for the Japanese bilinguals in the study there was an increase in cognitive demand
stemming from phonological rehearsal processes. The additional phonological processes might
have made the comprehension of the sentences slower and more demanding in L2, as the
significant differences between English and Japanese reading times also indicate. The network
of frontal lobe and angular gyrus activation indicates that the readers may be resorting to a
phonological rehearsal process in reading in the L2 (a phonological rehearsal loop). Activation
in areas associated with a phonological rehearsal loop, just prior to the probe processing,
indicates that in these intermediately proficient bilinguals there may be a cognitive mechanism
that assists comprehension processes in the less-proficient L2 by keeping sound-to-meaning
representations active.
Conclusion
Sentence reading in the Japanese logographic and syllabic systems produces orthography-
specific activation. The brain differentially responds to the two orthographies, activating right-
hemisphere areas associated with the pictorial and visuospatial characteristic of kanji
characters, and activating left and right-hemisphere areas associated with the syllabic nature
of hiragana.
Direct comparison across languages and different writing systems shows that there is activation
that indicates a brain response to reading in the less-proficient L2. For the Japanese bilinguals
in question, reading in English poses a cognitive challenge of processing in a language in which
they are less proficient, and a language which uses a different orthography. What the brain
activation indicates is that there is activation in cortical areas that may be associated with the
recognition and processing of words and letter-to-sound mappings, and with the additional
working memory load of keeping these mappings active. Ultimately, the data indicate that
within the same language, sentence-level processing can elicit differential cognitive responses
associated with word-level comprehension processes, and across languages, with more
cognitively demanding phonological rehearsal processes.
Acknowledgments
This research was supported by the National Institute of Mental Health Grant MH029617. We would like to thank
Natasha Tokowicz and the current members of the CCBI reading group for helpful comments on a previous draft of
this paper.
References
Bolger DJ, Perfetti CA, Schneider W. A cross-cultural effect on the brain revisited. Human Brain Mapping
2005;25:91–104.
Chee MW, Caplan D, Soon CS, Sriram N, Tan EW, Thiel T, Weekes B. Processing of visually presented
sentences in Mandarin and English studied with fMRI. Neuron 1999;23:127–137. [PubMed:
10402199]
Cohen L, Dehaene S. Specialization within the ventral stream: the case for the visual word form area.
NeuroImage 2004;22:466–476. [PubMed: 15110040]
Friston K, Ashburner J, Frith C, Poline JB, Heather J, Frackowiak R. Spatial registration and
normalization of images. Human Brain Mapping 1995;2:165–189.
Hasegawa M, Carpenter PA, Just MA. An fMRI study of bilingual sentence comprehension and workload.
NeuroImage 2002;15:647–660. [PubMed: 11848708]
Buchweitz et al. Page 8
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Jeong H, Sugiura M, Sassa Y, Yokoyama S, Horie K, Sato S, Taira M, Kawashima R. Cross-linguistic
influence on brain activation during second language processing: An fMRI study. Bilingualism:
Language and Cognition 2007;10:175–187.
Just MA, Carpenter PA, Keller TA, Eddy WF, Thulborn KR. Brain activation modulated by sentence
comprehension. Science 1996;274:114–116. [PubMed: 8810246]
Kim KHS, Relkin NR, Lee K, Hirsch J. Distinct cortical areas associated with native and second
languages. Nature 1997;388:171–174. [PubMed: 9217156]
Marian V, Spivey M, Hirsch J. Shared and separate systems in bilingual language processing: Converging
evidence from eye-tracking and brain imaging. Brain and Language 2003;86:70–82. [PubMed:
12821416]
Matsuo K, Kato C, Okada T, Moriya T, Glover GH, Nakai T. Finger movements lighten the neural loads
in the recognition of ideographic characters. Cognitive Brain Research 2003;17:263–272. [PubMed:
12880898]
Mechelli A, Crinion JT, Noppeney U, O’Doherty J, Ashburner J, Frackowiak RS, Price CJ. Structural
plasticity in the bilingual brain. Nature 2004;431:757. [PubMed: 15483594]
Meschyan G, Hernandez AE. Impact of language proficiency and orthographic transparency on bilingual
word reading: An fMRI investigation. NeuroImage 2006;29:1134–1140.
Nakamura K, Honda M, Okada T, Hanakawa T, Toma K, Fukuyama H, Konishi J, Shibasaki H.
Participation of the left posterior inferior temporal cortex in writing and mental recall of kanji
orthography: a functional MRI study. Brain 2000;123:954–967. [PubMed: 10775540]
Nakamura K, Dehaene S, Jobert A, Bihan DL, Kouider S. Subliminal convergence of kanji and kana
words: Further evidence for functional parcellation of the posterior temporal cortex in visual word
perception. Journal of Cognitive Neuroscience 2005;17:954–968. [PubMed: 15969912]
Oldfield RC. The assessment and analysis of handedness: The Edinburgh inventory. Neuropsychologia
1971;9:97–113. [PubMed: 5146491]
Paradis, M. Differential use of cerebral mechanisms in bilinguals. In: Banich, MT.; Mack, M., editors.
Mind, Brain, and Language: Multidisciplinary perspectives. Mahwah, NJ: Erlbaum Associates; 2003.
p. 351-370.
Paulesu E, McCrory F, Fazio F, Menoncello L, Brunswick N, Cappa SF, Cotelli M, Cossu G, Corte F,
Lorusso M, Pesenti S, Gallagher A, Price C, Frith CD, Frith U. A cultural effect on brain activation.
Nature Neuroscience 2000;3:91–96.
Perani D, Abutalebi J. The neural basis of first and second language processing. Current opinion in
Neurobiology 2005;15:202–206. [PubMed: 15831403]
Perfetti CA, Liu Y, Fiez J, Nelson J, Bolger D, Tan LH. Reading in two writing systems: Accommodation
and assimilation of the brain’s reading network. Bilingualism: Language and Cognition
2007;10:131–146.
Sakurai Y, Takeuchi S, Takada T, Horiuchi E, Nakase H, Sakuta M. Alexia caused by a fusiform or
posterior inferior temporal lesion. Journal of Neurological Sciences 2000;178:42–51.
Shieh CC, Luke KK, Tan LH, Wai YY, Wan YL, Liu HL. Brain mechanisms for syntactic and semantic
processing by Chinese and English bilinguals. NeuroImage 2001;13:602.
Tan LH, Liu H, Perfetti CA, Spinks JA, Fox PT, Gao J. The neural system underlying Chinese logograph
reading. NeuroImage 2001;13:836–846. [PubMed: 11304080]
Tan LH, Laird AR, Li K, Fox PT. Neuroanatomical correlates of phonological processing of Chinese
characters and alphabetic words: A meta-analysis. Human Brain Mapping 2005a;25:83–91.
[PubMed: 15846817]
Tan LH, Spinks JA, Eden GF, Perfetti CA, Siok WT. Reading depends on writing, in Chinese. Proceedings
of the National Academy of Sciences 2005b;102:8781–8785.
Tham WWP, Rickard Liow SJ, Rajapakse JC, Leong TC, Ng SES, Lim WEH, Ho LG. Phonological
processing in Chinese-English bilingual biscriptals: an fMRI study. NeuroImage 2005;28:579–587.
[PubMed: 16126414]
Thuy DHD, Matsuo K, Nakamura K, Toma K, Oga T, Nakai T, Shibasaki H, Fukuyama H. Implicit and
explicit processing of kanji and kana words and non-words studied with fMRI. NeuroImage
2004;23:878–889. [PubMed: 15528088]
Buchweitz et al. Page 9
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Tzourio-Mazoyer N, Landeau B, Papathanassiou D, Crivello F, Etard O, Delcroix N, Mazoyer B, Joliot
M. Automated anatomical labeling of activations in SPM using a macroscopic anatomical
parcellation of the MNI MRI single subject brain. NeuroImage 2002;15:273–289. [PubMed:
11771995]
Yamadori A. Neuropsychological model of reading based on Japanese experiences. Psychologia
2000;43:1–14.
Yokoyama S, Okamoto H, Miyamoto T, Yoshimoto K, Kim J, Iwata K, Jeong H, Uchida S, Ikuta N,
Sassa Y, Nakamura W, Horie K, Sato S, Kawashima R. Cortical activation in the processing of passive
sentences in L1 and L2: An fMRI study. NeuroImage 2006;30:570–579. [PubMed: 16300965]
Buchweitz et al. Page 10
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Figure 1.
Examples of the kanji and hiragana sentences.
Buchweitz et al. Page 11
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Figure 2.
Brain areas showing activation for kanji > hiragana (p < .005 uncorrected; T = 3.36; extent
threshold voxels = 20; right, a view of the inferior surface of the brain without the cerebellum;
ellipses highlight the activation of right-hemisphere inferior and mid temporal gyri).
Buchweitz et al. Page 12
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Figure 3.
Brain areas showing activation for hiragana > kanji (p < .005 uncorrected; T = 3.36; extent
threshold voxels = 20; top, surface renderings; bottom, coronal view showing SMA activation
(MNI coordinate: x = 10; y = –16; z = –56)).
Buchweitz et al. Page 13
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Figure 4.
Brain areas showing activation for English > hiragana, English > kanji, and English > Japanese
(average hiragana + kanji) (p < .005 uncorrected; T = 3.36; extent threshold voxels = 20; ellipses
highlight the loop of activation in the left frontal lobe and left angular gyrus).
Buchweitz et al. Page 14
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Buchweitz et al. Page 15
Table 1
Target-Sentence and Probe-Sentence Reading Times for English and Japanese
Target-Sentence Reading T (SD) (ms) Probe-Sentence Reading T (SD) (ms)
English 7215.51 (1452.33) 2390.48 (605.54)
kanji 4755.63 (1347.96) 1695.01 (370.52)
hiragana 4879.61 (1640.71) 1817.09 (407.31)
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Buchweitz et al. Page 16
Table 2
Clusters of activation for the Japanese writing system contrasts
kanji > hiragana BA Cluster size Z T-value MNI coordinate
x y z
R Mid Temporal gyrus* 39 127 4.42 9.81 56 −64 18
L Amygdala/Putamen 34 23 3.58 5.93 −22 −2−10
L Lingual gyrus 19 23 3.39 5.34 −22 −50 −4
R Inf. Temporal gyrus 37 31 3.28 5.02 58 −54 −12
hiragana > kanji BA Cluster size Z T-value MNI coordinate
x y z
L Inf. Parietal lobe 40 25 4.42 9.78 −48 −52 46
R Mid Cingulate gyrus* 5/7 162 3.79 6.71 16 −42 48
R Precuneus/Cingulate gyrus 31 112 3.62 6.10 24 −52 26
R Cuneus 18 22 3.58 5.95 16 −86 24
R Supp. Motor Area 6 99 3.37 5.28 10 −16 56
Left Precuneus 31 48 3.22 4.83 −20 −46 32
Left Ant. Cingulate gyrus 32 74 3.07 4.46 −10 26 28
R Supramarginal gyrus 40 22 3.02 4.32 50 −44 32
L Sup. Temporal gyrus 22/ 30 2.92 4.08 −46 −42 6
21
(P<.005 uncorrected; T3.36; extent threshold voxels=20; T-value, Z, and MNI coordinate are for the peak activated voxel in each cluster only. AAL labeling (Tzourio-Mazoyer, et al., 2002) clusters
marked with (*) were also significant at p<.05 after cluster-level correction for multiple representations)
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Buchweitz et al. Page 17
Table 3
Clusters for of activation for English > hiragana
BA Cluster size Z T- value MNI coordinate
x y z
L Angular gyrus* 39/40 134 3.88 7.06 −54 −62 34
L Inf. Parietal lobe
L Sup. Frontal gyrus 10 34 3.62 6.09 −18 60 18
R Sup. Frontal gyrus 9 69 3.52 5.75 16 38 36
R Sup. Frontal gyrus 9/1 58 3.30 5.08 26 44 16
0
R Mid Cingulate gyrus 31 58 3.26 4.94 0 −30 44
R Fusiform gyrus 20 34 3.25 4.91 32 −34 −20
L Inf. Frontal gyrus 47 75 3.22 4.86 −26 28 −10
L Lingual gyrus 19 21 3.17 4.70 −22 −50 −6
L Mid Cingulate gyrus 31 28 3.15 4.66 0 −46 36
R Sup. Medial Frontal gyrus 10 79 3.12 4.57 0 54 0
L Sup. Medial Frontal gyrus
L Cerebellum 4, 5 47 3.11 4.55 −8−56 −20
L Sup./Mid Temporal gyrus 21 48 3.09 4.50 −42 −12 −10
R Sup. Temporal gyrus 41 22 3.02 4.31 38 −32 16
R Supramarginal gyrus 40 25 3.01 4.29 60 −12 24
R Rolandic Operculum 13 21 2.91 4.06 42 −16 18
L Medial Frontal gyrus 10 23 2.85 3.94 −14 44 18
L Sup. Medial Frontal gyrus
(P<.005 uncorrected; T3.36; extent threshold voxels=20; T-value, Z, and MNI coordinate are for the peak activated voxel in each cluster only. AAL labeling (Tzourio-Mazoyer, et al., 2002) clusters
marked with (*) were also significant at p<.05 after cluster-level correction for multiple representations)
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Buchweitz et al. Page 18
Table 4
Clusters of activation for English > kanji
BA Cluster size Z T-value MNI coordinate
x y z
L Sup. Medial Frontal gyrus* 9 23 4.47 10.13 −4 44 30
L Precuneus 31 310 4.31 9.15 −4−50 32
R Supramarginal gyrus 40/43 21 3.98 7.53 58 −18 20
R Angular gyrus 39 26 3.69 6.32 48 −60 26
L Mid Frontal gyrus 8 33 3.65 6.21 −28 20 50
L Angular gyrus 40 37 3.51 5.71 −56 −58 30
L Supramarginal gyrus
R Mid Cingulate gyrus 31 57 3.35 5.22 14 −38 42
R Mid Temporal gyrus 22 46 3.32 5.14 68 −36 2
R Mid Cingulate gyrus 24 49 3.27 4.99 4 −20 44
L Mid Cingulate gyrus 24 28 3.22 4.84 −12 6 36
R Sup. Temporal gyrus 41 20 3.22 4.83 42 −32 10
R Sup. Frontal gyrus 9 48 3.21 4.82 20 38 36
L Inf. Frontal gyrus 47 39 3.06 4.42 −34 32 −18
L Mid Frontal gyrus
R Rolandic Operculum 13 26 3.00 4.27 44 −24 18
L Sup. Frontal gyrus 10 54 2.92 4.08 −12 54 12
(P<.005 uncorrected; T3.36; extent threshold voxels=20; T-value, Z, and MNI coordinate are for the peak activated voxel in each cluster only. AAL labeling (Tzourio-Mazoyer, et al., 2002) clusters
marked with (*) were also significant at p<.05 after cluster-level correction for multiple representations)
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
NIH-PA Author Manuscript NIH-PA Author Manuscript NIH-PA Author Manuscript
Buchweitz et al. Page 19
Table 5
Clusters of activation for English > (hiragana + kanji)
BA Cluster size Z T-value MNI coordinate
x y z
R Mid Cingulate gyrus* 24 320 4.09 8.03 2 −6 44
L Post. Cingulate gyrus 23 23 3.86 7.00 −4−36 30
L Precuneus 29 41 3.79 6.70 0 −54 6
R Mid Temporal gyrus 22 51 3.55 5.84 64 −40 8
L Mid Cingulate gyrus 31 30 3.41 5.40 −10 −42 44
R Sup. Frontal gyrus 9 80 3.31 5.08 16 42 36
R Supramarginal gyrus 43 53 3.31 5.08 58 −18 22
L Angular gyrus 39 63 3.18 4.74 −56 −58 28
L Inf. Parietal lobe
L Inf. Frontal gyrus 47 85 3.02 4.32 −42 40 −14
L Sup. Frontal gyrus 10 40 3.02 4.31 −18 60 18
L Sup. Medial Frontal gyrus 9/10 20 2.77 3.76 −8 50 22
R Sup. Medial Frontal gyrus 10 25 2.75 3.71 16 52 16
(P<.005 uncorrected; T3.36; extent threshold voxels=20; T-value, Z, and MNI coordinate are for the peak activated voxel in each cluster only. AAL labeling (Tzourio-Mazoyer, et al., 2002) clusters
marked with (*) were also significant at p<.05 after cluster-level correction for multiple representations)
Biling (Camb Engl). Author manuscript; available in PMC 2009 November 25.
