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Journal of Neurolinguistics 33 (2015) 14e28

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Journal of Neurolinguistics
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White-matter structure in the right hemisphere
predicts Mandarin Chinese learning success
Zhenghan Qi a, b, *, Michelle Han a, b, Keri Garel a, b,
Ee San Chen a, b, John D.E. Gabrieli a, b, c

Department of Brain and Cognitive Sciences, Massachusetts Institute of Technology, Cambridge, MA 02139,
McGovern Institute for Brain Research, Massachusetts Institute of Technology, Cambridge, MA 02139, USA
Institute for Medical Engineering & Science, Massachusetts Institute of Technology, Cambridge, MA 02139,

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 18 March 2014
Received in revised form 11 August 2014
Accepted 13 August 2014
Available online 2 September 2014

Second language learning becomes increasingly difficult with age,
but some adults learn more successfully than others. We examined
whether inter-subject variability in the microstructure of white
matter pathways, as measured by diffusion tensor imaging (DTI),
would predict native English speakers' outcomes in learning
Mandarin Chinese. Twenty-one adults were scanned before
participating in an intensive 4-week Mandarin course. At the end
of the Mandarin course, participants completed a final exam that
assessed their skills in both spoken and written Mandarin. Individual participants' white-matter tracts were reconstructed from
their native DTI data and related to final-exam performance. Superior language learning was correlated with DTI measures in the
right hemisphere, but not in the left hemisphere. In particular,
greater initial fractional anisotropy (FA) in both the right superior
longitudinal fasciculus (parietal bundle) and the right inferior
longitudinal fasciculus was associated with more successful Mandarin learning. The relation between white-matter structure in the
right hemisphere of native English speakers and successful initial
language learning may reflect the tonal and visuo-spatial properties, respectively, of spoken and written Mandarin Chinese.
© 2014 The Authors. Published by Elsevier Ltd. This is an open
access article under the CC BY-NC-ND license (http://

Second-language acquisition
Diffusion tensor imaging
White matter
Right hemisphere

* Corresponding author. Massachusetts Institute of Technology, 43 Vassar St., 46-4033, Cambridge, MA 02139, USA.
Tel.: þ1 617 324 4959.
E-mail address: (Z. Qi).
0911-6044/© 2014 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://

2005). we asked whether structural differences in whitematter pathways are related to differences among English-speaking adults in learning spoken and written Mandarin. the left AF is most prominently implicated in language function. 1841e1852.. Rauschecker et al. & Cabanis. Conversely. Johnson & Newport. a white matter tract in the dorsal pathway (VenturaCampos et al. 2013). 2009. 2011). and reading (Dronkers. 2013). 2007. Instead.. but neuroimaging studies have started to explore the structural and functional neural differences underlying individual variation in adult language learning. 2011). & Wandell. with few learners achieving native-like fluency in either the spoken or written form of the new language. two white-matter tracts have been most strongly associated with language: the left arcuate fasciculus (AF) component of the superior longitudinal fasciculus (SLF) and the left inferior longitudinal fasciculus (ILF). Yeatman. / Journal of Neurolinguistics 33 (2015) 14e28 15 1. Because Mandarin differs from English and other alphabetic languages in both spoken and written forms. second language acquisition during adulthood is exceptionally challenging (Hull & Vaid. and even these studies have reported mixed findings. Li. Honda.. & Schlaug. The left ILF has also been implicated more specifically in visual (orthographic) aspects of reading. 2007. Adults vary substantially in their second-language learning efficacy. Cramon. & Ffytche. 2004.. 2012. Sakai. 2008. diffusion measures of the left AF strongly associated with superior learning of English pseu pez-Barroso et al. Perani et al. Introduction In contrast to native language acquisition. perhaps by transmitting visual information about words between the fusiform gyrus at the temporaloccipital junction and anterior/interior temporal regions (Epelbaum et al.Z. Tittgemeyer. Yeatman et al. auditory comprehension. 1989. Here.. & Hanakawa. 2004. 2007). Iba-Zizen. 2013). Mandarin has four lexical tones (high- . Ben-Shachar. an arguably indistinguishable tract from the left ILF (Wakana et al.. 2012). only a few studies have explored relations between structural or functional connectivity in the right hemisphere and language learning. is highest in early bilinguals and lowest in monolinguals with late bilinguals being in between (Mohades et al. Nariai. 2007). In one study. 2005).. one of the diffusivity measures indexing white matter microstructure. such that fractional anisotropy (FA). 1998.. In studies of English and other alphabetic languages. Jones. The SLF connects a dorsal language network (Hickok & Poeppel. disruption of the left AF leads to impairment in speech production. Tanaka. learning proficiency correlated with the training-induced increases in the strength of the structural connectivity between the right inferior frontal gyrus and caudate (Hosoda. Garibaldi. One study found diffusion measures in the right AF correlated with individual's performance in pitch-related grammar learning (Loui. 2011).. Neuroimaging in relation to language learning has shown an association between individual learning performance and both structural and functional connectivity in the left hemisphere. Kuhl. pp. In contrast to the attention given to the left hemisphere. which are two regions connected by the left SLFp. Yeatman et al. Friederici. & Wong. Qi et al. Bilingualism also affects the left inferior frontal-occipital fasciculus. the neural bases that support learning Mandarin may be different from those that support learning European (alphabetic) languages. which seems to be effortless during the first years of childhood. Another study found a positive correlation between diffusion dowords (Lo measures of the left ILF and the learning of associations between Mandarin tones and English pseudowords (Wong. via another pathway. Plaisant. Little is known about how adult language learning capacities are represented in the brain. The ILF connects a ventral language network that includes Broca's area and posterior occipitotemporal regions and. a study of Japanese speakers learning English found no association between the learning performance and the white-matter structure in either the left or right hemisphere before learning. Dougherty. & Kno Among these language-related white-matter tracts. 2012). A resting-state functional magnetic resonance imaging (rsfMRI) study found that successful learning of foreign sound contrasts was associated with greater functional connectivity between Broca's area and the left superior parietal area. von €sche.. Vandermosten et al. the anterior temporal lobe with the uncinate fasciculus (Anwander. 2012. For spoken language. 2007) and is thought to consist of a direct AF pathway connecting posterior (superior temporal gyrus/Wernicke's area) and anterior (inferior frontal gyrus/ Broca's area) language cortices and an indirect pathway including the SLFp connecting the inferior parietal cortex and anterior language cortices (Catani. Chandrasekaran.

Xu et al. 2005). lexical tone is a crucial phonetic component signaling the semantics of the syllable. Zhang. processing Chinese characters with increased spacing between strokes elicited bilateral activation in the cuneus. Liu. 2011).. 2001. Zatorre & Gandour. there are two alternative possibilities in regards to the lateralization of early learning for naïve Mandarin learners whose native language is English. Further. Tan et al. 2009. we hypothesized that the visuo-spatial demands of learning Chinese characters may invoke right-hemisphere networks specialized for visuo-spatial processing (Pisella et al. rising. Hoosain. and there is evidence that the right hemisphere is dominant in processing pitch-related acoustic information (see review: Wong. Using diffusion tensor imaging (DTI). Learners were students enrolled in an intensive onemonth Mandarin training program in the classroom. Thus. & Schneider. a traditional learning environment similar to reallife learning experiences. Desroches. the processing of pitch information in speech has been associated with increased activation in both the right superior temporal gyrus and the right inferior frontal gyrus (Meyer. there is evidence that phonological processing of lexical tones is subserved by the left hemisphere in native Mandarin speakers (Gandour et al.. Unlike alphabetic languages. Tong et al. & Peng. may also depend upon left-hemisphere networks.. Wang. 2002). Pugh et al. Alsop. 2004. In Chinese-English bilingual readers. 2013. 2004.. & Chen. & von Cramon. Qi et al. in whom speech and reading is highly left-lateralized. Tan. Therefore. 2005). 2002. Pell. Mandarin learners cannot apply the same grapheme-to-phoneme scheme of alphabetic systems while reading Chinese.16 Z. 2012). 2005. especially for pitch contour information. a number of cross-linguistic studies have reported languagespecific effects on the recruited neural network. 2006). 2002. / Journal of Neurolinguistics 33 (2015) 14e28 level. each Chinese character is a monosyllabic word.. which map visual symbols onto phonemes. & Grodd. Wang. Gu.. Chinese characters usually consist of two or more spatially arranged radicles with a large variety of spatial relationships. Electrophysiological and fMRI evidence has shown that processing lexical tones is left-lateralized in native Mandarin speakers. The development of stronger inter-hemispheric connectivity suggested that Mandarin training in native English speakers. Hu. including production and perception of speech prosody (Baum & Pell. Saygin et al. Rudelson.. & Fox. One DTI study directly examined Mandarin-learning induced change in white matter structure. Warrier & Zatorre. Loui. 2012). 2013. the role of pitch perception in spoken Mandarin may invoke greater right-hemisphere participation than that occurring for non-tonal languages. Thus. This study did not examine whether initial variability in white-matter pathways was associated with better or worse learning. might lead to increased right-hemisphere involvement. but instead access the phonological information through visuo-orthographic information of the character (Tan. & Schlaug. Together with the consonantevowel combination. Mandarin employs a logographic system. Despite widespread consensus on left-hemisphere dominance in reading (Cohen & Dehaene. Alternatively. Ackermann. we examined whether the characteristics of language-relevant white-matter pathways (left and right AF. Thus. Wildgruber. & Tan. while processing acoustic tones remains right-lateralized regardless of listeners' language background (Gandour et al. 2005). Liu. Li. 1998. Tan et al. Perfetti. 2011. learning Mandarin lexical tones. 2008). Laird. 2005). Yeatman et al.. Friederici. SLFp. Early learning of spoken Mandarin could depend upon right-hemisphere networks involved in pitch processing. Alter. Erb. Yang. In the present study. 2013. The pronunciation of characters is usually opaque with only 28% of the phonetic radicles carrying the same phoneme information as the whole character (Perfetti. 2004. This longitudinal study measured white-matter microstructure over the course of nine months of Mandarin instruction and found increased fractional anisotropy (FA) in frontal-lobe tracts crossing the genu of the corpus callosum in the language-training group but not in the control group (Schlegel. 1999. 2004).. we extracted diffusion measures of . 2011. which is dominated by the left hemisphere in native Mandarin speakers. dipping and falling) that confer prosodic features to speech at the syllable level. The four tones can be distinguished by the shapes of the internal pitch contour. Chinese readers showed greater activation of the right middle occipital and right fusiform regions than English readers when reading in their native languages (Bolger. while processing English words with increased spacing did not (Sun. Despite the role of the right-hemisphere in acoustic processing of lexical tones that is important for spoken Mandarin. Lohmann.. and ILF) at baseline were correlated with variation in proficiency of learning spoken and written Mandarin in adults. Damage to the right auditory cortex or disruptions to its connectivity pattern leads to impairment in pitch-related processing. & Zhao.. 2011). & Tse. 1995. & Peng. Tracy et al. For written language. Pihan.

Z.5 vs. we hypothesized that higher FA values would predict greater Mandarin proficiency in the learners. 10 quizzes. The final exam was composed of speech production. mostly written in Chinese characters. All volunteers gave written consent that was approved by the institutional review board at Massachusetts Institute of Technology prior to their participation. Because outlier values can have strong influences on correlational analyses. Orthographic skill was measured in a computer-based task at the laboratory administered within one week after the Mandarin course. All the Chinese characters were taught in the class and were paired based on their close resemblance in visual forms (e. if individual sensitivity to pitch information and visuo-spatial information is important for early Mandarin learning. Spanish. Qi et al. scores were percentage of correct answers.. one participant was excluded based on diffusion measures that deviated more than 2. Because previous research has reported positive correlations between FA values and superior language learning. Participants Twenty-two participants were recruited for the study. For both tests. Kaufman & Kaufman. Among these. Importantly. Daily homework was assigned from the companion workbook and was supplemented with additional material designed by the instructor. participants saw an English word and were asked to choose the correct translation from the two given Chinese characters. on average. participants spent. three participants spoke the other language fluently and one participant spoke English exclusively during childhood. Four participants reported growing up in a bilingual environment (Polish. 2010) as the main textbook.6. An anonymous mid-term survey (in which 14 of the 21 students responded) showed participants also spent. and syntactic information. / Journal of Neurolinguistics 33 (2015) 14e28 17 fractional anisotropy (FA) for each tract within each individual. The course used The New Practical Chinese Reader Volume 1 (Liu. 13 males and 8 females.2.74. listening. The participants had an average standard IQ of 117 (SD ¼ 12. 62.8.g. on average. which tapped into the students' ability to effectively integrate orthographic information with phonologic. All analyses proceeded with the remaining twenty-one participants (mean age ¼ 23. the right-hemisphere pathways may be more associated with gaining early Mandarin proficiency than the left-hemisphere pathways. All participants were exposed to foreign . were native American-English speakers. we included two more tests that more specifically measured individuals' phonological skill and orthographic skill.3 h in the classroom and completed 11 assignments.5 standard deviations from the mean in 5 of the 6 tracts of interest. 18 right-handed and 3 left-handed). 2. which took place on the MIT campus. In particular. semantic. 星or果?). In each of the 30 trials. and 1 final exam. STAR. SD ¼ 3. as indicated by the Language History Questionnaire (Linck et al. Phonological skill was measured by participants' accuracy in the pinyin (an alphabetic phonetic system for Mandarin pronunciation) dictation part of the final exam. out of 100). In addition to measuring the participants' holistic usage of the language with the final exam total scores. The final exam was graded by the instructor on a scale of 0e100 and was used as the overall measure for the students' Mandarin learning performance.84) measured by Kaufman Brief Intelligence Test (KBIT-2. the four bilingual participants did not differ from the monolinguals in respect to their average final exam scores (77. and reading sections. 2. of which 17 were monolinguals with no extensive exposure to languages other than English as a child. 2004). Methods 2. Hindi and Hebrew). Language training procedure and proficiency tests Participants signed up for one of two course sections taught by the same instructor.5 h per day for 5 days per week over 4 weeks.1. All 21 participants. 2013). Behavioral accuracy reflected how well one could match the slight difference in the spatial arrangement of strokes with different semantic meanings. Participants were screened for MRI compatibility and no history of neurological diseases. The students wrote down the pinyin after listening to each of 10 Mandarin phrases/sentences twice. 76. The course. 3. 2.7 h every day outside of class studying Mandarin. provided students with intensive training in introductory Modern Standard Mandarin. Throughout the course. 1 mid-term exam.

. which automatically excluded extensively distorted volumes. radial diffusivity (RD). with further manual editing for accuracy. 176 slices).64 ms. FSL's bedpostx tool was used to fit a ball-and-stick model of diffusion to each subject's DWIs. Japanese Kanji). none of the participants had prior exposure to Chinese. matrix ¼ 128 mm  128 mm) and 64 slices per volume.0 (limited) and 5. we examined three bilateral tracts of interest. 2014) automated artifact detection software. 2011). The selfreport fluency of the second language ranged between 1. The b0 volumes were collected in the first 10 volumes of the entire scan and were later averaged. Of these.mgh. 2. Imaging acquisition and processing Diffusion tensor images were acquired before Mandarin training began. The diffusion-weighted scan included 10 non-diffusion-weighted reference volumes (b ¼ 0) and 60 diffusion-weighted volumes (b ¼ 700 s/mm2) with voxel resolution of 2.0 (excellent). The diffusion tensor eigenvalues extracted from TRACULA outputs were used to compute axial diffusivity (AD). red). AD describes diffusion along the longest principal axis of a fiber. purple). crucially. Data were collected on a Siemens Trio 3-Tesla MRI scanner with a standard 32-channel phased array head coil (TE ¼ 84 ms.harvard. defined a priori: the ILF. FA describes the ratio of normalized standard deviation and root mean square of the three diffusion directions. Qi et al. RD describes diffusion perpendicular to the principle diffusion direction. Thai).. / Journal of Neurolinguistics 33 (2015) 14e28 languages. with FA values of 0. Illustration of tracts of interest in axial (left) and sagittal (right) views. A high-resolution T1weighted magnetization prepared rapid gradient echo (MPRAGE) scan was also acquired (TE ¼ 1. 1. .2e1 indicating white matter and greater values connoting greater white matter integrity (Kunimatsu et al. FOV ¼ 220 mm. The data were then processed using TRACULA (Yendiki et al. TR ¼ 2530 ms.and inter-registration of each subject's b0 diffusion images to their T1-weighted images and the MNI template respectively. 2004). TR ¼ 8040 ms). The DTI data were pre-processed using DTIPrep's (Oguz et al.g.15).0 mm isotropic (field of view ¼ 256 mm. FA was used as the main measure because it summarizes white matter microstructure. other tonal languages (e. ranging from 1 to 5 different languages. The FA values range from 0 (equal diffusion in all directions) to 1. and fractional anisotropy (FA) for the three bilateral tracts of interest. In our analysis. with large variability in the level of Tracts were extracted from an example subject and were registered to Montreal Neurological Imaging (MNI) template space. However. TRACULA's pathway reconstruction software generated probability distributions of the trajectory of the 18 major white matter pathways. 1. the SLFp. Cortical and white matter surfaces were generated according to individual landmarks using the FreeSurfer image analysis suite (http://surfer.18 Z. calculated the tensor fit at each voxel to generate maps of diffusion tensor eigenvalues. 1). performed both intra. which corrected for eddy currents and head motion. or any other language with a logographic writing system (e.0 (standard deviation.3.nmr. and the AF (Fig. The tracts of interest include: Inferior Longitudinal Fasciculus (ILF. Superior Longitudinal Fasciculus parietal bundle (SLFp. and estimated the endpoints and waypoints for each major white matter pathway using subject-specific anatomical information. blue) and Arcuate Fasciculus (AF. matrix ¼ 220 mm  220 mm. with an average of 3. AD and RD values were used in a Fig..

which generated a confidence interval based on 105 random resamplings in order to test whether they were statistically different Fig. one-tailed statistical tests were used. All reported p values have been false detection rate (FDR) corrected for multiple comparisons (FDR < 0. i. Results 3. Statistical methods We examined the relations between the participants' final exam scores and three measures derived from the FA values: 1) average FA across all three tracts of interest within each hemisphere. one-tailed p ¼ . 2) average FA within each tract of interest. Therefore. 2007). two-tailed p ¼ . data normality was not assumed. The average FA value of (a) the left tracts did not correlate with subsequent Mandarin learning success. Lee. More positive laterality scores indicate higher average FA values in the left tract. Spearman's rank correlation. Because FA values are bounded between 0 and 1 with considerable skew and kurtosis (Alexander. We directly compared the strengths of the leftand right-hemisphere correlations using a bootstrap procedure. we adopted a non-parametric method. Fig. Lazar.Z. one-tailed p ¼ . but there was a significant correlation between the mean baseline FA of the three right-hemisphere tracts and final exam scores (Spearman's r ¼ . . while the average FA value of (b) the right tracts did correlate significantly and positively with subsequent Mandarin learning success. Right but not left hemisphere tracts predicted Mandarin learning ability The mean baseline FA of the three left-hemisphere tracts did not significantly correlate with final exam scores (Spearman's r ¼ . Fig. / Journal of Neurolinguistics 33 (2015) 14e28 19 post hoc analysis after we located tracts that were important for Mandarin learning. Black lines represent the linear trend of the correlation and grey shades represent the 95% confidence interval of the data. The selective correlation with right-hemisphere pathways occurred despite strong overall correlations between the means of the left and right pathways across individuals (Spearman's r ¼ . i. in order to better characterize the underlying biological factors contributing to the inter-subject variability of the FA values. For analyses where both directions of hypothesis were possible (correlation between tract lateralization and learning performance).1. 2b). right-lateralized. 2.4.24. For analyses with unidirectional hypotheses (greater FA in the tracts for analysis are associated with better learning performance). and 3) laterality scores of each pair of tracts calculated as Laterality Score ¼ (FAleft  FAright)/(FAleft þ FAright).55. 2. twotailed tests were used. More negative laterality scores indicate higher average FA values in the right tract. SLFp and AF) and final exam scores. left-lateralized.e.14. 2a).001).005. 3. Qi et al.e. Correlation between the average FA across three tracts of interest (ILF.05).65. & Field. for all the following analyses.

3b). In both (a) and (b). 3a) and the right SLFp (Spearman's r ¼ .02). 1996). right AF. The tracts are colored according to the FA values. adding either right SLFp or right ILF FA values as a second predictor increased the variance of data explained by the model from 15.06). onetailed FDR-corrected. black lines represent the linear trend of the correlation and grey shades represent the 95% confidence interval of the data. The correlation between final exam scores and FA of both tracts remained significant (p's < .4%. we analyzed two additional diffusion parameters. In order to further investigate the possible anatomical structure differences that contributed to the inter-subject variability of FA values. in the right ILF and right SLFp. one-tailed p ¼ . 3.20 Z.32.28). 3. and left ILF and participants' final exam scores (uncorrected p's < . Table 1). Fig. The average AD did not correlate with individual final exam scores in either tract (Spearman's r's < . AD and RD. p ¼ .10. (c) Right ILF and (d) right SLFp were rendered in two example subjects (circled points) to illustrate the FA difference between a less successful (Subj. Qi et al. we conducted a partial correlation analysis to control for individual IQ. Compared with the model using IQ as the only predictor.06. Fig. . Because final exam scores were also marginally correlated with IQ (Spearman's r ¼ .1% and 22. There was a significantly stronger correlation between the final exam scores and FA values in the right hemisphere than those in the left hemisphere (one-tailed p < .02).60. one-tailed FDR-corrected. p ¼ .01. one-tailed p's > .2. Right ILF and right SLFp predicted Mandarin learning ability Individual final exam scores correlated positively with FA of the right ILF (Spearman's r ¼ . 1) and a more successful (Subj.50. the average RD significantly correlated with Fig. Individual IQ scores and tract FA values (arcsin transformed in order to meet the normality assumption of the linear regression analyses) were fit into linear regression models for the prediction of the final exam scores.03.08).04. There were also marginally significant correlations between the FA values in the left AF. / Journal of Neurolinguistics 33 (2015) 14e28 from each other (DiCiccio & Efron. Greater initial FA values of (a) the right ILF and (b) the right SLFp correlate with subsequent Mandarin learning success. However. respectively (model comparison p ¼ . 2) learner. p ¼ .1% to 25.

one-tailed p ¼ 0. we investigated the relations between initial white-matter microstructure and subsequent Mandarin learning success. Previous research. Greater right-lateralization of ILF FA correlated positively with superior final exam scores (Spearman's r ¼ .10 .56. higher FA in the right ILF and right SLFp predicted greater success in learning Mandarin. we calculated laterality scores for each tract in each participant.3. has shown that FA in homologous white matter structures are highly correlated in individuals (Wahl et al. Correlation p (Uncorrected) p (FDR) Left ILF Left SLFp Left AF Right ILF Right SLFp Right AF . Fig.12 FDR ¼ false-discovery rate. 4). We found a significantly stronger positive correlation between students' final exam scores and initial FA in righthemisphere tracts than in left-hemisphere tracts. two-tailed FDR-corrected p ¼ .12 .45.57..05 .01 .16.06. we further examined the relation between hemispheric asymmetry of white-matter microstructure and Mandarin learning performance.51 . one-tailed p ¼ 0. Participants displayed great variability in lateralization.41 . correlated significantly only with the average FA values of the right ILF (Spearman's r ¼ 0.49). There was no relation between final exam scores and AF lateralization (Spearman's r ¼ .54. More than 50% of the participants showed right-lateralized ILF. Participants' performance on the final orthography measure. two-tailed FDR-corrected p ¼ .29 .005) and the right SLFp (Spearman's r ¼ 0. . As revealed by separate behavioral subtests.50 .006) and not the right SLFp (Spearman's r ¼ 0. whereas more than 50% showed left-lateralized SLFp (Fig. 4b). Participants' performance on the final phonology measure correlated significantly with baseline average FA values of both the right ILF (Spearman's r ¼ 0. Our laterality analysis of tract FA values showed left-lateralized SLF tracts Table 1 Spearman correlations between white matter anisotropy and the final exam score. one-tailed p ¼ 0. two-tailed FDR-corrected p ¼ . / Journal of Neurolinguistics 33 (2015) 14e28 21 individual final exam score in both the right ILF (Spearman's r ¼ 0.53.18).002 .004). Tract laterality and handedness The FA characterization of white-matter tracts is consistent with other studies. 4a) and greater right-lateralization of SLFp FA tended to correlate with superior final exam scores (Spearman's r ¼ . FA in both right-hemisphere tracts was associated with phonology learning success.42.Z. 4. while FA in the right ILF was associated with only orthography learning success.35 .04.45. Discussion In this study. 4. Analysis of hemispheric asymmetries of FA further revealed that more rightward ILF and less leftward SLFp were associated with higher final exam scores. however. one-tailed p ¼ 0.60 .01 .03) and the right SLFp (Spearman's r ¼ 0. 3. By comparing the FA values between the two hemispheres for each pair of tracts of interest.1.12 . one-tailed p ¼ 0.32 . 2010). Fig. We also examined the specific relations of the right ILF and SLFp tracts to Mandarin phonology learning (as measured by the pinyin dictation task in the final exam) and orthography learning (as measured by the two-alternative forced-choice task).21. one-tailed p ¼ 0. like our study. Qi et al. In particular. Laterality of ILF and SLFp predicted Mandarin learning ability Given that right-hemisphere tracts played a greater role in predicting Mandarin learning efficacy than left-hemisphere tracts.02). Native English speakers enrolled in a one-month introductory Mandarin course and reached a wide range of proficiency at the end of the training. These results provide converging evidence for the hypothesis that the right-hemisphere tracts play a more important role than left hemisphere tracts in preparing adults for initial learning of Mandarin as a second language.08 .06 .03 . Numbers in bold fonts indicate statistical significance.

2008). First..22 Z. and myelination (Beaulieu.. Saur et al. 4. TRACULA does not. 2012). This result is consistent with previous findings of an overall leftward asymmetry of AF regardless of handedness or functional language lateralization.2. fiber density. Therefore. Takahashi et al. 2010.. however that the ILF includes fibers from other language-related ventral pathways. Kno Turner. membrane permeability. & diameter. Jones. based on the anatomical regions of interest defined in Makris et al. This finding suggests that increased axonal myelination or denser axonal tracts (lower RD). in most participants.. that carried the correlation between the white matter microstructure and Mandarin learning success.. (2005) (Yendiki et al. 2008. The SLF is divided into AF and SLFp. The AF in this study corresponds best to the classically defined “direct” pathway of the arcuate fasciculus. 2002. both of which connect the posterior parietal area with the posterior temporal area. Lorenz et al. 2010). It is possible. 2004. 2002). as determined by the hemispheric difference in response to a verb production task (Vernooij et al. while the SLFp is closest to the anterior segment of the “indirect” pathway (Catani et al. Qi et al. / Journal of Neurolinguistics 33 (2015) 14e28 Fig. rather than axonal integrity measured by AD.. similar to the majority of the right-handers. the tract laterality scores of all three left-handers showed left lateralized SLFp. there are limitations to the accurate delineation of tracts by TRACULA. TRACULA reconstructs white matter into 18 tracts. TRACULA identifies ILF as one of the ventral stream pathways. We also reanalyzed the data excluding the three left-handers and found the same significant correlations between the right-hemisphere tracts and Mandarin learning. differentiate the posterior segment of the indirect pathway from the posterior part of the AF. 4.. Glasser & Rilling. Likewise. 2005). supported superior Mandarin learning. but not AD. consistent with findings of volume asymmetry of the tracts (Ellmore et al. 2002. which can be associated with higher AD and/or lower RD (Song et al. such as the middle . the precise neurobiological basis of the FA correlations cannot be determined. Second. DTI analyses of white-matter microstructure Higher FA values likely reflect a number of different tract-specific properties. 2007). Initial hemispheric asymmetry of tract FA values correlates with subsequent Mandarin learning success. There are limitations to DTI analyses of white-matter tracts.. however. diffusion tensor modeling is not able to solve the problem of intra-voxel crossing fibers.. 2011). one possible concern regarding the current data set is that our sample included three left-handers. Because the incidence of right functional laterality in language increases with the degree of left-handedness (Basic et al. (a) More rightlateralized ILF and (b) less left-lateralized SLFp were associated with higher final exam scores. including fiber € sche. Black lines represent the linear trend of the correlation and grey shades represent the 95% confidence interval of the data. Our findings showed it was RD. Reduced RD and increased FA might result from fewer crossing fibers in successful learners. However.

Qi et al. Prior studies found associations between diffusivity measures in the left hemisphere.3. 2011).. Higher FA in the right superior temporal and the right lingual gyrus white-matter regions were positively correlated with faster reading time of English words in a Chinese-English bilingual group.. white-matter diffusivity measures in the right hemisphere predicted Mandarin learning. Further. 2005). native Mandarin speakers recruited the right inferior frontal gyrus more strongly in the production of characters with different tones than the production of characters with different vowels (Liu et al. during a tone discrimination task predicted whether an individual was successful in learning new pseudowords with Mandarin lexical tones (Wong. which is associated with visual processes. The other challenge Mandarin learners usually face is learning to read Chinese characters. 4. but not in European multilinguals who did not speak Mandarin (Crinion et al. The most relevant evidence comes from a training study in native English speakers who were naïve to Mandarin. Moreover. Similarly. Right-hemisphere contribution to Mandarin learning In the present study. Wong et al. It is noteworthy that only the properties of the right ILF.. Saur et al. which is more associated with auditory processes. Our finding that learners' performance in the orthography task positively associated with their FA values of the right ILF tract before training is consistent with these findings. hemisphere tracts in learning new phonemes. Wong et al. Increased white-matter volume in the tract arising from the right Heschl's gyrus was found in multilingual Mandarin speakers and native Mandarin speakers. Moreover. Previous studies showed greater activation of the right middle occipital and right fusiform regions in Chinese readers than English readers when reading in their native languages (Bolger et al. A DTI-identified pathway between superior/middle temporal gyrus and the frontal lobe is located in right-hemisphere regions associated with prosodic processing in European languages (Glasser & Rilling. 2009. & Parrish. The level of activation in the right temporal gyrus. those studies focused on European languages without the tonal properties of Mandarin or the non-alphabetic visuo-spatial properties of Chinese characters. which have great visuo-spatial complexity. 2013. Fiez. the engagement of the right hemisphere in early attention to pitch information in Mandarin appears to be a shared mechanism irrespective of language experience. and not the right SLFp. crosssectional studies discovered right-hemisphere structural characteristics specific to Chinese speakers.. Neuroimaging studies have found evidence suggesting that the brain areas connected by the ILF and the SLFp in the right hemisphere participate in processing pitch information of Mandarin speech.. 2009). but not in an English monolingual group (Cummine & Boliek... there is greater right-hemisphere involvement for Mandarin speech than for nontonal European languages. 2007). White-matter studies provide evidence for a structural counterpart to the right-hemisphere functional specialization for pitch processing. & Perfetti. / Journal of Neurolinguistics 33 (2015) 14e28 23 longitudinal fasciculus and inferior fronto-occipital fasciculus (Dick & Tremblay.. 2009). these findings may seem to contradict the dominant involvement of left € el. 2005. On the surface. there was a positive correlation between FA of the right temporal-parietal conjunction and individual's ability to learn an artificial pitch-related grammar (Loui et al. However. In a Chinese character naming task. The reduction of right SLF volume was found to be associated with the behavioral deficiencies of tone-deafness (Loui et al. 2008).. 2009). 2004). 2005). A rightward asymmetry of activation in the superior temporal sulcus and the middle frontal gyrus was found in both Mandarin and English native speakers performing same-different judgment on Mandarin intonation across syllables and on tones within syllables (Gandour et al. 2011). 2006). Lo Friederici.. & Johansen-Berg. a rightward asymmetry in the middle frontal gyrus was reported in both Mandarin and English native speakers processing sentence-level prosody of Mandarin (Tong et al. but not in the right hemisphere.. and grammar in adults (Flo  pez-Barroso et al. as well as the left temporal gyrus. English speakers learning Mandarin increased recruitment of the right fusiform regions when viewing Chinese characters (Nelson. Tan et al. 2012.Z. Liu.. 2013). Overall. words. 2009). Scholz. de Vries. 2008. Perrachione. 2011. these studies support the idea that due to the tonal nature of Mandarin speech. see review: Breitenstein. and language learning performance. had a significant association with .

Similar findings from a MEG study suggested greater right visual word form area activation and therefore more bilateral recruitment in L2 (English) word processing compared to L1 (Spanish. & Hutchins. especially in early learning. Legault & Litcofsky. and there are many homophones per single-syllable word. Li. 1998. with superior orthographic learning is consistent with the idea that early learning of Chinese characters is more related to visual and less to phonological processes than that occurs in alphabetic languages. and therefore may show a different relationship with brain structure. Marshall. 1994. Therefore. 2001. / Journal of Neurolinguistics 33 (2015) 14e28 orthography learning in Mandarin Chinese. 2007). 1999.. One longitudinal study revealed Ferna learning-induced change of right-hemisphere structure within individuals. 2003. whose native language is less similar to English compared to German speakers (D'Anselmo. but not the right SLFp. direct mapping of orthographic information to meaning may be efficient. Japanese speakers showed enhancement of grey matter and white matter density in the right IFG after learning English (Hosoda. Honda. Craik. & Hernandez. & Carreiras. & Brancucci. & Grady. Iturria-Medina. switching. Leonard et al. Pisella et al. Bialystok. these studies suggest right-hemisphere involvement in second-language learning may not be limited to Mandarin learning. Loui et al. Language learning at a more advanced stage may involve distinct underlying processes from earlier stages. early bilingual speakers of two alphabetical languages showed stronger anterior-posterior structural and functional connectivity within both the left and right hemispheres (Luk. An fMRI study revealed more bilateral activation elicited by children who processed English speech as a second language compared to those who processed English speech as a native language (Archila-Suerte. Hsieh. 2014). Gillon-Dowens. 2011). Together. Tommasi. as well as selecting. it is noteworthy that the association between white matter structure and Mandarin learning success revealed in the current study is based on the proficiency tests after only one month of training. & Ferreira. 2013). 2004). A behavioral meta-analyses study of bilingual laterality found that second language proficiency in late bilinguals was positively correlated with right-hemisphere involvement (Hull & Vaid... such as bilateral prefrontal and parietal areas (Abutalebi & Green. Warrier & Zatorre. Zevin. The extent of right-hemisphere involvement appears to be modulated by the similarity between L1 and L2. Neuropsychological evidence also supports the possibility that learning tonal and visuo-spatial aspects of Mandarin may invoke right-hemisphere brain regions. Both the right dorsal and ventral visual streams have been implicated to play important roles in the cognitive control of visual search (Lane... 2004. Luk & Green. Only an experiment with another control group of participants learning an alphabetic language could test the specific role of the right hemisphere in Mandarin learning. 2011). The present study is not able to differentiate between these two alternative explanations. 2007. Gandour. & Vallar. Pell. 2013). Groisman. . 2001. semantic analysis of Chinese characters is difficult via phonology because there is a lack of transparent mapping between phonemes and graphemes.. The correlation of the right ILF. including production and perception of speech prosody (Baum & Pell. & Zhao. In Mandarin. and translating between native and foreign languages might lead to recruiting areas outside of the network for one's native language. & Makashvili. 2006). Early bilingual speakers also showed increased structural connectivity in a subnetwork involving the right  n. Wong. Pe rez superior frontal gyrus and the left reading network measured by graph efficiency (García-Pento ndez.24 Z. Fink. An alternative interpretation of our finding is that the microstructure of white matter in the right hemisphere is involved not only in Mandarin learning. Ramos. Tong et al. The importance of lefthemisphere engagement in fluent Mandarin has been implicated by the findings that native Mandarin speakers show left-lateralized functional representation of language (Gandour et al. Mkheidze.. Zatorre. Mosidze. Compared to monolingual English speakers. 2011). 2014. 2012). Lesion at the right posterior parietal regions and occipital regions leads to disruption of visuo-spatial attention network (Halligan. Damage to the right auditory cortex leads to impairment in pitch-related calculation. Milner. Zuccarini. Nariai. Klein. 2010). but also second-language learning in general. Finally. Xu et al. A dichotic behavioral test with English words as L2 showed reduced leftward asymmetry in Italian speakers. The contribution of right hemisphere in bilingualism is further supported by a number of neuroimaging studies. Tanaka. The cognitive control demands of adapting to new linguistic rules. & Hanakawa. Qi et al. Structural MRI studies found that increased exposure to second language is associated with greater grey matter density in the right hippocampus (Mårtensson et al. 2013). Reiterer. 2009. 2005.

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