Skip to main content Accessibility help

Meta-analysis of the neural representation of first language and second language



This study reports an activation likelihood estimation meta-analysis of published functional neuroimaging studies of bilingualism. Four parallel meta-analyses were conducted by taking into account the proficiency of participants reported in the studies. The results of the meta-analyses suggest differences in the probabilities of activation patterns between high proficiency and moderate/low proficiency bilinguals. The Talairach coordinates of activation in first language processing were very similar to that of second language processing in the high proficient bilinguals. However, in the low proficient group, the activation clusters were generally smaller and distributed over wider areas in both the hemispheres than the clusters identified in the ALE maps from the high proficient group. These findings draw attention to the importance of language proficiency in bilingual neural representation.


Corresponding author

ADDRESS FOR CORRESPONDENCE Rajani Sebastian, Department of Communication Sciences & Disorders, University of Texas at Austin, Austin, TX 78712. E-mail:


Hide All
Abutalebi, J. (2008). Neural aspects of second language representation and language control. Acta Pscychologica, 128, 466478.
Abutalebi, J., & Green, D. (2007). Bilingual language production: The neurocognition of language representation and control. Journal of Neurolinguistics, 20, 242275.
Abutalebi, J., Cappa, S. F., & Perani, D. (2001). The bilingual brain as revealed by functional neuroimaging. Bilingualism: Language and Cognition, 4, 179190.
Abutalebi, J., Cappa, S. F., & Perani, D. (2005). Functional neuroimaging of the bilingual brain. In Kroll, J. F. K. & De Groot, A. M. (Eds.), Handbook of bilingualism: Psycholinguistic approaches. Oxford: Oxford University Press.
Albert, M. L., & Obler, L. K. (1978). The bilingual brain: Neuropsychological and neurolinguistic aspects of bilingualism. New York: Academic Press.
Bookheimer, S. (2002). Functional MRI of language: New approaches to understanding the cortical organization of semantic processing. Annual Review Neuroscience, 25, 151188.
Booth, J. R., Wood, L., Lu, D., Houk, J. C., & Bitan, T. (2007). The role of the basal ganglia and cerebellum in language processing. Brain Research, 1133, 136144.
Botvinick, M., Nystrom, L. E., Fissell, K., Carter, C. S., & Cohen, J. D. (1999). Conflict monitoring versus selection-for-action in anterior cingulate cortex. Nature, 402, 179181.
Braver, T. S., Barch, D. M., Gray, J. R., Molfese, D. L., & Avraham, S. (2001). Anterior cingulate cortex and response conflict: Effects of frequency, inhibition, and errors. Cerebral Cortex, 11, 825836.
Brown, S., Laird, A. R., Ingham, R. J., Ingham, J. C., & Fox, P. T. (2005). Stuttered and fluent speech production: An ALE meta-analysis of functional neuroimaging studies. Human Brain Mapping, 25, 105117.
Bush, G., Frazier, J. A., Rauch, S. L., Seidman, L. J., Whalen, P. J., Jenike, M. A., et al. (1999). Anterior cingulate cortex dysfunction in attention-deficit/hyperactivity disorder revealed by fMRI and the Counting Stroop. Biological Psychiatry, 45, 15421552.
Bush, G., Luu, P., & Posner, M. I. (2000). Cognitive and emotional influences in anterior cingulate cortex. Trends in Cognitive Sciences, 4, 215222.
Cabeza, R., & Nyberg, L. (1997). Imaging cognition: An empirical review of PET studies with normal subjects. Journal of Cognitive Neuroscience, 9, 126.
Callan, D. E., Jones, J. A., Callan, A. M., & Akahane-Yamada, R. (2004). Phonetic perceptual identification by native- and second-language speakers differentially activates brain regions involved with acoustic phonetic processing and those involved with articulatory-auditory/orosensory internal models. NeuroImage, 22, 11821194.
Carter, C. S., Braver, T. S., Barch, D. M., Botvinick, M. M., Noll, D., & Cohen, J. D. (1998). Anterior cingulate cortex, error detection, and the online monitoring of performance. Science, 280, 747749.
Chee, M. W., Tan, E. W. L., & Thiel, T. (1999). Mandarin and English single words processing studied with functional magnetic resonance imaging. Journal of Neuroscience, 15, 30503056.
Chee, M. W. L., Hon, N., Lee, H. L., & Soon, C. S. (2001). Relative language proficiency modulates BOLD signal change when bilinguals perform semantic judgments. NeuroImage, 13, 11551163.
Chee, M. W. L., Weekes, B., Lee, K. M., Soon, C. S., Schreiber, A., Hoon, J. J., et al. (2000). Overlap and dissociation of semantic processing of Chinese characters, English words, pictures: Evidence from fMRI. NeuroImage, 12, 392403.
De Bleser, R., Dupont, P., Postler, J., Bormans, G., Speelman, D., Mortelmans, L., et al. (2003). The organization of the bilingual lexicon: A PET study. Journal of Neurolinguistics, 16, 439456.
Dehaene, S., & Changeux, J. P. (1991). The Wisconsin Card Sort Test: Theoretical analysis and modeling in a neuronal network. Cerebral Cortex, 1, 6279.
Dehaene, S., Dupoux, E., Mehler, J., Cohen, L., Paulesu, E., Perani, D., et al. (1997). Anatomical variability in the cortical representation of first and second language. NeuroReport, 8, 38093815.
Derrfuss, J., Brass, M., Neumann, J., & Yves von Cramon, D. (2005). Involvement of the inferior frontal junction in cognitive control: Meta-analyses of switching and Stroop studies. Human Brain Mapping, 25, 2234.
Desimone, R., & Duncan, J. (1995). Neural mechanisms of selective attention. Annual Review of Neuroscience, 18, 193222.
Desmond, J. E., & Fiez, J. A. (1998). Neuroimaging studies of the cerebellum: Language, learning and memory. Trends in Cognitive Sciences, 2, 355361.
Ding, G., Perry, C., Peng, D., Ma, L., Li, D., Xu, S., et al. (2003). Neural mechanisms underlying semantic and orthographic processing in Chinese–English bilinguals. NeuroReport, 14, 15571562.
Edmonds, L., & Kiran, S. (2006). Effect of semantic naming treatment on cross linguistic generalization in bilingual aphasia. Journal of Speech, Language, and Hearing Research, 49, 729748.
Eickhoff, S. B., Laird, A. R., Grefkes, C., Wang, L. E., Zilles, K., & Fox, P. T. (2009). Coordinate-based activation likelihood estimation meta-analysis of neuroimaging data: A random-effects approach based on empirical estimates of spatial uncertainty. Human Brain Mapping, 30, 29072926.
Ellison-Wright, I., Glahn, D. C., Laird, A. R., Thelen, S. M., & Bullmore, E. T. (2008). The anatomy of first-episode and chronic schizophrenia: An anatomical likelihood estimation meta-analysis. American Journal of Psychology, 165, 10151023.
Fabbro, F. (1999). The neurolinguistics of bilingualism. Hove: Psychology Press.
Fernandez, B., Cardebat, D., Demonet, J. F., Joseph, P. A., Mazaux, J. M., Barat, M., et al. (2004). Functional MRI follow-up study of language processes in healthy subjects and during recovery in a case of aphasia. Stroke, 35, 21712176.
Fitzgerald, P. B., Oxley, T. J., Laird, A. R., Kulkarni, J., Egan, G. F., & Daskalakis, Z. J. (2006). An analysis of functional neuroimaging studies of dorsolateral prefrontal cortical activity in depression. Psychiatry Research, 148, 3345.
Fulbright, R. K., Jenner, A. R., Mencl, W. E., Pugh, K. R., Shaywitz, B. A., & Shaywitz, S. E. (1999). The cerebellum's role in reading: A functional MRI imaging study. American Journal of Neuroradiology, 20, 19251930.
Gandour, J., Tong, Y., Talavage, T., Wong, D., Dzemidzic, M., Xu, Y., et al. (2007). Neural basis of first and second language processing of sentence-level linguistic prosody. Human Brain Mapping, 28, 94–08.
Genovese, C. R., Laxar, N. A., & Nochols, T. (2002). Thresholding of statistical maps in functional neuroimaging using the false discovery rate. NeuroImage, 15, 870878.
Glahn, D. C., Ragland, J. D., Abramoff, A., Barrett, J., Laird, A. R., Bearden, C. E., et al. (2005). Beyond hypofrontality: A quantitative meta-analysis of functional neuroimaging studies of working memory in schizophrenia. Human Brain Mapping, 25, 6069.
Golestani, N., Alario, F., Meriaux, S., Le Bihan, D., Dehaene, S., & Pallier, C. (2006). Syntax production in bilinguals. Neuropsychologia, 44, 10291040.
Gomez-Tortosa, E., Martin, E. M., Gaviria, M., Charbel, F., & Ausman, J. I. (1995). Selective deficit of one language in a bilingual patient following surgery in the left perisylvian area. Brain and Language, 48, 320325.
Hagoort, P. (2005). On Broca, brain, and binding: A new framework. Trends in Cognitive Science, 9, 416423.
Hasegawa, M., Carpenter, P. A., & Just, M. A. (2002). An fMRI study of bilingual sentence comprehension and workload. NeuroImage, 15, 647660.
Hernandez, A. E., Dapretto, M., Mazziotta, J., & Bookheimer, S. (2001). Language switching and language representation in Spanish–English bilinguals: An fMRI study. NeuroImage, 14, 510520.
Hernandez, A. E., Martinez, A., & Kohnert, K. (2000). In search of the language switch: An fMRI study of picture naming in Spanish–English bilinguals. Brain and Language, 73, 421431.
Illes, J., Francis, W. S., Desmond, J. E., Gabrieli, J. D. E., Glover, G. H., Poldrack, R., et al. (1999). Convergent cortical representation of semantic processing in bilinguals. Brain and Language, 70, 347363.
Indefrey, P. (2006). A Meta-analysis of hemodynamic studies on first and second language processing: Which suggested differences can we trust and what do they mean? Language Learning, 56, 279304.
Indefrey, P., & Levelt, W. J. M. (2004). The spatial and temporal signatures of word production components. Cognition, 92, 101144.
Junque, C., Vendrell, P., & Vendrell, J. (1995). Differential impairments and specific phenomena in 50 Catalan–Spanish bilingual aphasic patient. In Paradis, M. (Ed.), Aspects of bilingual aphasia. Oxford: Pergamon Press.
Kerns, J. G., Cohen, J. D., MacDonald, A. W., Cho, R. Y., Stenger, V. A., & Carter, C. S. (2004). Anterior cingulate conflict monitoring and adjustments in control. Science, 303, 10231026.
Kim, K. H., Relkin, N. R., Lee, K. M., & Hirsch, J. (1997). Distinct cortical areas associated with native and second languages. Nature, 388, 171174.
Klein, D., Milner, B., Zatorre, R. J., Meyer, E., & Evans, A. C. (1995). The neural substrates underlying word generation: A bilingual functional imaging study. Proceedings of the National Academy of Science of the United States of America, 92, 28992903.
Klein, D., Milner, B., Zatorre, R. J., Zhao, V., & Nikelski, J. (1999). Cerebral organization in bilinguals: A PET study of Chinese–English verb generation. NeuroReport, 10, 28412845.
Klein, D., Zatorre, R. J., Milner, B., Meyer, E., & Evans, A. C. (1994). Left putaminal activation when speaking a second language: Evidence from PET. NeuroReport, 5, 22952297.
Krain, A. L., Wilson, A. M., Arbuckle, R., Castellanos, F. X., & Milham, M. P. (2006). Distinct neural mechanisms of risk and ambiguity: A meta-analysis of decision-making. NeuroImage, 32, 477484.
Kuchonov, P., Lancaster, J., Thompson, P., Toga, A. W., Brewer, P., & Hardies, J. (2002). An optimized individual target brain in the Talairach coordinate system. NeuroImage, 17, 922927.
Laird, A. R., Fox, M., Price, C. J., Glahn, D. C., Uecker, A. M., Lancaster, J. L., et al. (2005). ALE meta-analysis: Controlling the false discovery rate and performing statistical contrasts. Human Brain Mapping, 25, 155164.
Laird, A. R., McMillan, K. M., Lancaster, J. L., Kochunov, P., Turkeltaub, P. E., Pardo, J. V., et al. (2005). A comparison of label-based review and ALE meta-analysis in the Stroop task. Human Brain Mapping, 25, 621.
Lancaster, J. L., Tordesillas-Gutierrez, D., Martinez, M., Salinas, F., Evans, A., Zilles, K., et al. (2007). Bias between MNI and Talairach coordinates analyzed using the ICBM-152 brain template. Human Brain Mapping, 28, 11941205.
Luke, K., Liu, H., Wai, Y., Wan, Y., & Tan, L. H. (2002). Functional anatomy of syntactic and semantic processing in language comprehension. Human Brain Mapping, 16, 133145.
Ma, L., Wang, B., Narayana, S., Hazeltine, E., Chen, X., Robin, D. A., et al. (2010). Changes in regional activity are accompanied with changes in inter-regional connectivity during 4 weeks motor learning. Brain Research, 1318, 6476.
MacDonald, A. W., Cohen, J. D., Stenger, V. A., & Carter, C. S. (2000). Dissociating the role of dorsolateral prefrontal cortex and anterior cingulate cortex in cognitive control. Science, 288, 18351838.
Mahendra, N., Plante, E., Magloire, J., Milman, L., & Trouard, T. (2003). MRI variability and the localization of languages in the bilingual brain. NeuroReport, 14, 12251228.
Marian, V., Spivery, M., & Hirsch, J. (2003). Shared and separate systems in bilingual language processing: Converging evidence from eye tracking and brain imaging. Brain and Language, 86, 7082.
McMillan, K. M., Laird, A. R., Witt, S. T., & Meyerand, M. E. (2007). Self-paced working memory: Validation of verbal variations of the n-back paradigm. Brain Research, 1139, 133142.
Menzies, L. A. C., Chamberlain, S. R., Laird, A. R., Thelen, S. M., Sahakian, B. J., & Bullmore, E. T. (2008). Integrating evidence from neuroimaging and neuropsychological studies of obsessive compulsive disorder: The orbitofronto-striatal model revisited. Neuroscience Biobehavioral Review, 32, 525549.
Meschyan, G., & Hernandez, A. E. (2006). Impact of language proficiency and orthographic transparency on bilingual word reading: An fMRI investigation. NeuroImage, 29, 11351140.
Miller, E. K., & Cohen, J. D. (2001). An integrative theory of prefrontal cortex. Annual Review of Neuroscience, 24, 167202.
Moro, A., Tettamanti, M., Perani, D., Donati, C., Cappa, S. F., & Fazio, F. (2001). Syntax and the brain: Disentangling grammar by selective anomalies. NeuroImage, 13, 110118.
Nakai, T., Matsuo, K., Kato, C., Matsuzawa, M., Okada, T., Glover, G. H., et al. (1999). A functional magnetic resonance imaging study of listening comprehension of languages in human at 3-Tesla comprehension level and activation of the language areas. Neuroscience Letter, 263, 3336.
Nilipour, R., & Ashayeri, H. (1989). Alternating antagonism between two languages with successive recovery of a third in a trilingual aphasic patient. Brain and Language, 36, 2348.
Owen, A. M., McMillan, K. M., Laird, A. R., & Bullmore, E. (2005). N-back working memory paradigm: A meta-analysis of normative functional neuroimaging studies. Human Brain Mapping, 25, 4659.
Paradis, M. (1977). Bilingualism and aphasia. In Whitaker, H. & Whitaker, A. (Eds.), Studies in neurolinguistics. New York: Academic Press.
Perani, D., Dehaene, S., Grassi, F., Cohen, L., Cappa, S. F., Dupoux, F. F., et al. (1996). Brain processing of native and foreign languages. NeuroReport, 7, 24392444.
Perani, D., Paulesu, E., Galles, N. S., Dupoux, E., Dehaene, S., Bettinardi, V., et al. (1998). The bilingual brain: Proficiency and age of acquisition of the second language. Brain, 121, 18411852.
Petrides, M., Alivasatos, B., Meyer, E., & Evans, A. C. (1993). Functional activation of the human frontal cortex during the performance of verbal working memory tasks. Proceedings of the National Academy of Science of the United States of America, 90, 878882.
Pillai, J. J., Allison, J. D., Sethuraman, S., Araque, J. M., Thiruvaiyaru, D., Ison, C. B., et al. (2004). Functional MR imaging study of language-related differences in bilingual cerebellar activation. American Journal of Neuroradiology, 25, 523532.
Pillai, J. J., Araque, J. M., Allison, J. D., Sethuraman, S., Loring, D. W., & Thiruvaiyaru, D. (2003). Functional MRI study of semantic and phonological language processing in bilingual subjects: Preliminary findings. NeuroImage, 19, 565576.
Poldrack, R. A., Wagner, A. D., Prull, M. W., Desmond, J. E., Glover, G. H., & Gabrieli, J. D. E. (1999). Functional specialization for semantic and phonological processing in the left inferior prefrontal cortex. NeuroImage, 10, 1535.
Price, C. J., Devlin, J. T., Moore, C. J., Morton, C., & Laird, A. R. (2005). Meta-analyses of object naming: Effect of baseline. Human Brain Mapping, 25, 7082.
Price, C. J., Green, D. W., & von Studnitz, R. (1999). A functional imaging study of translation and language switching. Brain, 122, 22212235.
Rüschemeyer, S. A., Fiebach, C., Kempe, V., & Friederici, A. D. (2005). Processing lexical semantic and syntactic information in first and second language: fMRI evidence from Russian and German. Human Brain Mapping, 25, 266286.
Ruschemeyer, S. A., Zysset, S., & Friederici, A. D. (2006). Native and non-native reading of sentences: An fMRI experiment. NeuroImage, 31, 354365.
Siok, W. T., Perfetti, C. A., Jin, Z., & Tan, L. H. (2004). Biological abnormality of impaired reading is constrained by culture. Nature, 431, 7176.
Stowe, L., & Sabourin, L. (2005). Imaging the processing of a second language: Effects of maturation and proficiency on the neural processes involved. International Review of Applied Linguistics in Language Teaching, 43, 329354.
Suh, S., Yoon, H. W., Lee, S., Chung, J., Cho, Z., & Park, H. (2007). Effects of syntactic complexity in L1 and L2; An fMRI study of Korean–English bilinguals. Brain Research, 1136, 178189.
Talairach, J., & Tournoux, P. (1988). Co-planar stereotaxic atlas of the human brain: 3-Dimensional proportional system: An approach to cerebral imaging. New York: Thieme Medical.
Tan, L. H., Laird, A. R., Li, K., & Fox, P. T. (2005). Neuroanatomical correlates of phonological processing of Chinese characters and alphabetic words: A meta-analysis. Human Brain Mapping, 25, 8391.
Tan, L. H., Liu, H. L., Perfetti, C. A., Spinks, J. A., Fox, P. T., & Gao, J. H. (2001). The neural system underlying Chinese logograph reading. NeuroImage, 13, 836846.
Tan, L. H., Spinks, J. A., Feng, C.-M., Siok, W. T., Perfetti, C. A., Xiong, J., et al. (2003). Neural systems of second language reading are shaped by native language. Human Brain Mapping, 18, 158166.
Tham, W. W. P., Liow, S. J. R., Rajapakse, J. C., Leong, T. C., Ng, S. E. S., Lim, W. E. H., et al. (2005). Phonological processing in Chinese–English bilinguals biscriptals: An fMRI study. NeuroImage, 28, 579587.
Thulborn, K. R., Carpenter, P. A., & Just, M. A. (1999). Plasticity of language-related brain function during recovery from stroke. Stroke, 30, 749754.
Turkeltaub, P. E., Eden, G. F., Jones, K. M., & Zeffiro, T. A. (2002). Meta-analysis of the functional neuroanatomy of single-word reading: Method and validation. NeuroImage, 16, 765780.
Vandenberghe, R., Price, C., Wise, R., Josephs, O., & Frackowiak, R. S. J. (1996). Functional anatomy of a common semantic system for words and pictures. Nature, 383, 254256.
van Hell, J. G., & Dijkstra, T. (2002). Foreign language knowledge can influence native language performance in exclusively native contexts. Psychonomic Bulletin & Review, 9, 780789.
van Wijnendaele, I., & Brysbaert, M. (2002). Visual word recognition in bilinguals: Phonological priming from the second to the first language. Journal of Experimental Psychology, 28, 616627.
Wang, Y., Xue, G., Chen, C., Xue, F., & Dong, Q. (2007). Neural bases of asymmetric language switching in second-language learners: An ER-fMRI study. NeuroImage, 35, 862870.
Wartenburger, I., Heekeren, H., Abutalebi, J., Cappa, S., Villringer, A., & Perani, D. (2003). Early setting of grammatical processing in the bilingual brain. Neuron, 37, 159170.
Weiller, C., Isensee, C., Rijntjes, M., Huber, W., Muller, S., & Bier, D. (1995). Recovery from Wernicke's aphasia: A positron emission tomographic study. Annals of Neurology, 37, 723732.
Winhuisen, L., Thiel, A., Schumacher, B., Kessler, J., Rudolf, J., & Haupt, W. F. (2007). Role of the contralateral inferior frontal gyrus in recovery of language function in post stroke aphasia: A combined repetitive transcranial magnetic stimulation and positron emission tomography study. Stroke, 36, 17591763.
Xiong, J., Ma, L., Wang, B., Narayana, S., Duff, E. P., Egan, G. F., et al. (2009). Long-term motor training induced changes in regional cerebral blood flow in both task and resting states. NeuroImage, 45, 7582.
Yokoyama, S., Okamoto, H., Miyamoto, T., Yoshimoto, K., Kim, J., Iwata, K., et al. (2006). Cortical activation in the processing of passive sentences in L1 and L2: An fMRI study. NeuroImage, 30, 570579.
Zatorre, R. J., Meyer, E., Gjedde, A., & Evans, A. C. (1996). PET studies of phonetic processing of speech: Review, replication, and reanalysis. Cerebral Cortex, 6, 2130.

Related content

Powered by UNSILO

Meta-analysis of the neural representation of first language and second language



Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed.