Multilingualism and brain plasticity

• Abutalebi, J., Annoni, J.-M., Zimine, I., Pegna, A. J., Seghier, M. L., Lee-Jahnke, H., … Khateb, A. (2008). Language control and lexical competition in bilinguals: an event-related FMRI study. Cerebral Cortex, 18(7), 1496–505. https://doi.org/10.1093/cercor/bhm182 • Abutalebi, J., Canini, M., Della Rosa, P. A., Green, D. W., & Weekes, B. S. (2015). The neuroprotective effects of bilingualism upon the inferior parietal lobule : a structural neuroimaging study in aging Chinese bilinguals. Journal of Neurolinguistics, 33, 3–13. https://doi.org/10.1016/j.jneuroling.2014.09.008 • Abutalebi, J., Canini, M., Della Rosa, P. A., Sheung, L. P., Green, D. W., & Weekes, B. S. (2014). Bilingualism protects anterior temporal lobe integrity in aging. Neurobiology of Aging, 35(9), 2126–2133. https://doi.org/10.1016/j.neurobiolaging.2014.03.010 • Abutalebi, J., Della Rosa, P. A., Ding, G., Weekes, B. S., Costa, A., & Green, D. W. (2013). Language proficiency modulates the engagement of cognitive control areas in multilinguals. Cortex, 49(3), 905–911. https://doi.org/10.1016/j.cortex.2012.08.018 • Abutalebi, J., Della Rosa, P. A., Gonzaga, A. K. C., Keim, R., Costa, A., Perani, D., … Perani, D. (2013). The role of the left putamen in multilingual language production. Brain and Language, 125(3), 307–315. https://doi.org/10.1016/j.bandl.2012.03.009 • Abutalebi, J., Della Rosa, P. A., Green, D. W., Hernandez, M., Scifo, P., Keim, R., … Costa, A. (2012). Bilingualism tunes the anterior cingulate cortex for conflict monitoring. Cerebral Cortex, 22(9), 2076–86. https://doi.org/10.1093/cercor/bhr287 • Abutalebi, J., & Green, D. W. (2016). Neuroimaging of language control in bilinguals: neural adaptation and reserve. Bilingualism: Language and Cognition, 19(4), 689–698. https://doi.org/10.1017/S1366728916000225 • Abutalebi, J., Guidi, L., Borsa, V., Canini, M., Della Rosa, P. A., Parris, B. A., & Weekes, B. S. (2015). Bilingualism provides a neural reserve for aging populations. Neuropsychologia, 69, 201–210. https://doi.org/10.1016/j.neuropsychologia.2015.01.040 • Alladi, S., Bak, T. H., Duggirala, V., Surampudi, B., Shailaja, M., Shukla, A. K., … Kaul, S. (2013). Bilingualism delays age at onset of dementia, independent of education and immigration status. Neurology, 81(22), 1938–1944. https://doi.org/10.1212/01.wnl.0000436620.33155.a4 • Anderson, J. A. E., Grundy, J. G., De Frutos, J., Barker, R. M., Grady, C., & Bialystok, E. (2018). Effects of bilingualism on white matter integrity in older adults. NeuroImage, 167, 143–150. https://doi.org/10.1016/j.neuroimage.2017.11.038 • Becker, M., Schubert, T., Strobach, T., Gallinat, J., & Kühn, S. (2016). Simultaneous interpreters vs. professional multilingual controls: Group differences in cognitive control as well as brain structure and function. NeuroImage. https://doi.org/10.1016/j.neuroimage.2016.03.079 • Bellander, M., Berggren, R., Mårtensson, J., Brehmer, Y., Wenger, E., Li, T. Q., … Lövdén, M. (2016). Behavioral correlates of changes in hippocampal gray matter structure during acquisition of foreign vocabulary. NeuroImage, 131, 205–213. https://doi.org/10.1016/j.neuroimage.2015.10.020 • Berken, J. A., Chai, X. J., Chen, J.-K., Gracco, V. L., & Klein, D. (2016). Effects of early and late bilingualism on resting-state functional connectivity. Journal of Neuroscience, 36(4), 1165–1172. https://doi.org/10.1523/JNEUROSCI.1960-15.2016 • Berken, J. A., Gracco, V. L., Chen, J.-K., & Klein, D. (2016). The timing of language learning shapes brain structure associated with articulation. Brain Structure and Function, 221(7), 3591–3600. https://doi.org/10.1007/s00429-015-1121-9 • Bialystok, E. (2009). Bilingualism: The good, the bad, and the indifferent. Bilingualism: Language and Cognition, 12(1), 3. https://doi.org/10.1017/S1366728908003477 • Burgaleta, M., Sanjuán, A., Ventura-Campos, N., Sebastián-Gallés, N., & Ávila, C. (2016). Bilingualism at the core of the brain. Structural differences between bilinguals and monolinguals revealed by subcortical shape analysis. NeuroImage, 125, 437–445. https://doi.org/10.1016/j.neuroimage.2015.09.073 • Coggins, P. E., Kennedy, T. J., & Armstrong, T. a. (2004). Bilingual corpus callosum variability. Brain and Language, 89(1), 69–75. https://doi.org/10.1016/S0093-934X(03)00299-2 • Crinion, J. T., Turner, R., Grogan, A., Hanakawa, T., Noppeney, U., Devlin, J. T., … Price, C. J. (2006). Language control in the bilingual brain. Science, 312(5779), 1537–1540. https://doi.org/10.1126/science.1127761 • Cummine, J., & Boliek, C. A. (2013). Understanding white matter integrity stability for bilinguals on language status and reading performance. Brain Structure & Function, 218(2), 595–601. https://doi.org/10.1007/s00429-012-0466-6 • De Smet, H. J., Paquier, P. F., Verhoeven, J., & Mariën, P. (2013). The cerebellum: its role in language and related cognitive and affective functions. Brain and Language, 127(3), 334–42. https://doi.org/10.1016/j.bandl.2012.11.001 • Della Rosa, P. A., Videsott, G., Borsa, V. M., Canini, M., Weekes, B. S., Franceschini, R., & Abutalebi, J. (2013). A neural interactive location for multilingual talent. Cortex, 49(2), 605–8. https://doi.org/10.1016/j.cortex.2012.12.001 • Draganski, B., Gaser, C., Kempermann, G., Kuhn, H. G., Winkler, J., Buchel, C., & May, A. (2006). Temporal and spatial dynamics of brain structure changes during extensive learning. Journal of Neuroscience, 26(23), 6314–6317. https://doi.org/10.1523/JNEUROSCI.4628-05.2006 • Elmer, S., Hänggi, J., & Jäncke, L. (2014). Processing demands upon cognitive, linguistic, and articulatory functions promote grey matter plasticity in the adult multilingual brain: Insights from simultaneous interpreters. Cortex, 54, 179–89. https://doi.org/10.1016/j.cortex.2014.02.014 • Elmer, S., Hänggi, J., Meyer, M., & Jäncke, L. (2011). Differential language expertise related to white matter architecture in regions subserving sensory-motor coupling, articulation, and interhemispheric transfer. Human Brain Mapping, 32(12), 2064–74. https://doi.org/10.1002/hbm.21169 • Felton, A., Vazquez, D., Ramos-Nunez, A. I., Greene, M. R., Macbeth, A., Hernandez, A. E., & Chiarello, C. (2017). Bilingualism influences structural indices of interhemispheric organization. Journal of Neurolinguistics, 42, 1–11. https://doi.org/10.1016/j.jneuroling.2016.10.004 • Filippi, R., Richardson, F. M., Dick, F., Leech, R., Green, D. W., Thomas, M. S. C., & Price, C. J. (2011). The right posterior paravermis and the control of language interference. Journal of Neuroscience, 31(29), 10732–10740. https://doi.org/10.1523/JNEUROSCI.1783-11.2011 • Friederici, A. D., & Gierhan, S. M. E. (2013). The language network. Current Opinion in Neurobiology, 23(2), 250–254. https://doi.org/10.1016/j.conb.2012.10.002 • García-Pentón, L., Fernández, A. P., Iturria-Medina, Y., Gillon-Dowens, M., Carreiras, M., Pérez Fernández, A., … Carreiras, M. (2014). Anatomical connectivity changes in the bilingual brain. NeuroImage, 84, 495–504. https://doi.org/http://dx.doi.org/10.1016/j.neuroimage.2013.08.064 • Gold, B. T., Johnson, N. F., & Powell, D. K. (2013). Lifelong bilingualism contributes to cognitive reserve against white matter integrity declines in aging. Neuropsychologia, 51(13), 2841–2846. https://doi.org/10.1016/j.neuropsychologia.2013.09.037 • Golestani, N. (2014). Brain structural correlates of individual differences at low-to high-levels of the language processing hierarchy: A review of new approaches to imaging research. International Journal of Bilingualism, 18(1), 6–34. https://doi.org/10.1177/1367006912456585 • Grady, C. L., Luk, G., Craik, F. I. M., & Bialystok, E. (2015). Brain network activity in monolingual and bilingual older adults. Neuropsychologia, 66, 170–181. https://doi.org/10.1016/j.neuropsychologia.2014.10.042 • Grogan, A., Green, D. W., Ali, N., Crinion, J. T., & Price, C. J. (2009). Structural correlates of semantic and phonemic fluency ability in first and second languages. Cerebral Cortex, 19(11), 2690–8. https://doi.org/10.1093/cercor/bhp023 • Grogan, A., Parker Jones, O., Ali, N., Crinion, J. T., Orabona, S., Mechias, M. L., … Price, C. J. (2012). Structural correlates for lexical efficiency and number of languages in non-native speakers of English. Neuropsychologia, 50(7), 1347–52. https://doi.org/10.1016/j.neuropsychologia.2012.02.019 • Hämäläinen, S., Sairanen, V., Leminen, A., & Lehtonen, M. (2017). Bilingualism modulates the white matter structure of language-related pathways. NeuroImage, 152, 249–257. https://doi.org/10.1016/j.neuroimage.2017.02.081 • Hervais-Adelman, A., Moser-Mercer, B., Murray, M. M., & Golestani, N. (2017). Cortical thickness increases after simultaneous interpretation training. Neuropsychologia, 98(January), 212–219. https://doi.org/10.1016/j.neuropsychologia.2017.01.008 • Hofstetter, S., Friedmann, N., & Assaf, Y. (2017). Rapid language-related plasticity: microstructural changes in the cortex after a short session of new word learning. Brain Structure and Function, 222(3), 1231–1241. https://doi.org/10.1007/s00429-016-1273-2 • Hosoda, C., Tanaka, K., Nariai, T., Honda, M., & Hanakawa, T. (2013). Dynamic neural network reorganization associated with second language vocabulary acquisition: a multimodal imaging study. The Journal of Neuroscience, 33(34), 13663–72. https://doi.org/10.1523/JNEUROSCI.0410-13.2013 • Just, M. A., Cherkassky, V. L., Keller, T. a, Kana, R. K., & Minshew, N. J. (2007). Functional and anatomical cortical underconnectivity in autism: evidence from an FMRI study of an executive function task and corpus callosum morphometry. Cerebral Cortex, 17(4), 951–61. https://doi.org/10.1093/cercor/bhl006 • Kaiser, A., Eppenberger, L. S., Smieskova, R., Borgwardt, S., Kuenzli, E., Radue, E.-W., … Bendfeldt, K. (2015). Age of second language acquisition in multilinguals has an impact on gray matter volume in language-associated brain areas. Frontiers in Psychology, 6, 1–9. https://doi.org/10.3389/fpsyg.2015.00638 • Klein, D., Mok, K., Chen, J.-K., & Watkins, K. E. (2014). Age of language learning shapes brain structure: A cortical thickness study of bilingual and monolingual individuals. Brain and Language, 131, 20–24. https://doi.org/10.1016/j.bandl.2013.05.014 • Kousaie, S., Chai, X. J., Sander, K. M., & Klein, D. (2017). Simultaneous learning of two languages from birth positively impacts intrinsic functional connectivity and cognitive control. Brain and Cognition, 117, 49–56. https://doi.org/10.1016/j.bandc.2017.06.003 • Koziol, L. F., Budding, D., Andreasen, N., D?Arrigo, S., Bulgheroni, S., Imamizu, H., … Yamazaki, T. (2014). Consensus paper: The cerebellum’s role in movement and cognition. The Cerebellum, 13(1), 151–177. https://doi.org/10.1007/s12311-013-0511-x • Kuhl, P. K., Stevenson, J., Corrigan, N. M., van den Bosch, J. J. F., Can, D. D., & Richards, T. (2016). Neuroimaging of the bilingual brain: Structural brain correlates of listening and speaking in a second language. Brain and Language, 162, 1–9. https://doi.org/10.1016/j.bandl.2016.07.004 • Li, L., Abutalebi, J., Emmorey, K., Gong, G., Yan, X., Feng, X., … Ding, G. (2017). How bilingualism protects the brain from aging: Insights from bimodal bilinguals. Human Brain Mapping, 38(8), 4109–4124. https://doi.org/10.1002/hbm.23652 • Li, L., Abutalebi, J., Zou, L., Yan, X., Liu, L., Feng, X., … Ding, G. (2015). Bilingualism alters brain functional connectivity between “control” regions and “language” regions: Evidence from bimodal bilinguals. Neuropsychologia, 71, 236–247. https://doi.org/10.1016/j.neuropsychologia.2015.04.007 • Li, P., & Grant, A. M. (2016). Second language learning success revealed by brain networks. Bilingualism: Language and Cognition, 19(4), 657–664. https://doi.org/10.1017/S1366728915000280 • Liu, H., Hu, Z., Guo, T., & Peng, D. (2010). Speaking words in two languages with one brain: Neural overlap and dissociation. Brain Research, 1316, 75–82. https://doi.org/10.1016/j.brainres.2009.12.030 • Liu, X., Tu, L., Wang, J., Jiang, B., Gao, W., Pan, X., … Huang, R. (2017). Onset age of L2 acquisition influences language network in early and late Cantonese-Mandarin bilinguals. Brain and Language, 174, 16–28. https://doi.org/10.1016/j.bandl.2017.07.003 • Luk, G., Bialystok, E., Craik, F. I. M., & Grady, C. L. (2011). Lifelong bilingualism maintains white matter integrity in older adults. The Journal of Neuroscience, 31(46), 16808–13. https://doi.org/10.1523/JNEUROSCI.4563-11.2011 • Maguire, E., Gadian, D. G., Johnsrude, I. S., Good, C. D., Ashburner, J., Frackowiak, R. S., & Frith, C. D. (2000). Navigation-related structural change in the hippocampi of taxi drivers. Proceedings of the National Academy of Sciences of the United States of America, 97(8), 4398–403. https://doi.org/10.1073/pnas.070039597 • Mamiya, P. C., Richards, T. L., Coe, B. P., Eichler, E. E., Kuhl, P. K., Geschwind, D. H., & Paus, T. (2016). Brain white matter structure and COMT gene are linked to second-language learning in adults. Proceedings of the National Academy of Sciences, 113(26), 7249–7254. https://doi.org/10.1073/pnas.1606602113 • Mandonnet, E., Nouet, A., Gatignol, P., Capelle, L., & Duffau, H. (2007). Does the left inferior longitudinal fasciculus play a role in language? A brain stimulation study. Brain, 130(3), 623–629. https://doi.org/10.1093/brain/awl361 • Mechelli, A., Crinion, J. T., Noppeney, U., O’Doherty, J., Ashburner, J., Frackowiak, R. S., & Price, C. J. (2004). Neurolinguistics: Structural plasticity in the bilingual brain. Nature, 431(7010), 757–757. https://doi.org/10.1038/431757a • Mechelli, A., Price, C. J., Friston, K. J., & Ashburner, J. (2005). Voxel-based morphometry of the human brain: methods and applications. Current Medical Imaging Reviews, 1(2), 105–113. https://doi.org/10.2174/1573405054038726 • Mohades, S. G., Struys, E., Van Schuerbeek, P., Mondt, K., Van De Craen, P., & Luypaert, R. (2012). DTI reveals structural differences in white matter tracts between bilingual and monolingual children. Brain Research, 1435, 72–80. https://doi.org/10.1016/j.brainres.2011.12.005 • Mohades, S. G., Van Schuerbeek, P., Rosseel, Y., Van De Craen, P., Luypaert, R., & Baeken, C. (2015). White-matter development is different in bilingual and monolingual children: a longitudinal DTI study. Plos One, 10, e0117968. https://doi.org/10.1371/journal.pone.0117968 • Nichols, E. S., & Joanisse, M. F. (2016). Functional activity and white matter microstructure reveal the independent effects of age of acquisition and proficiency on second-language learning. NeuroImage, 143, 15–25. https://doi.org/10.1016/j.neuroimage.2016.08.053 • Olsen, R. K., Pangelinan, M. M., Bogulski, C., Chakravarty, M. M., Luk, G., Grady, C. L., & Bialystok, E. (2015). The effect of lifelong bilingualism on regional grey and white matter volume. Brain Research, 1612, 128–139. https://doi.org/10.1016/j.brainres.2015.02.034 • Olulade, O. A., Jamal, N. I., Koo, D. S., Perfetti, C. A., LaSasso, C., & Eden, G. F. (2016). Neuroanatomical evidence in support of the bilingual advantage theory. Cerebral Cortex, 26(7), 3196–3204. https://doi.org/10.1093/cercor/bhv152 • Osterhout, L., Poliakov, A., Inoue, K., McLaughlin, J., Valentine, G., Pitkanen, I., … Hirschensohn, J. (2008). Second-language learning and changes in the brain. Journal of Neurolinguistics, 21(6), 509–521. https://doi.org/10.1016/j.jneuroling.2008.01.001 • Park, I. S., Lee, K. J., Han, J. W., Lee, N. J., Lee, W. T., Park, K. A., & Rhyu, I. J. (2009). Experience-dependent plasticity of cerebellar vermis in basketball players. The Cerebellum, 8(3), 334–339. https://doi.org/10.1007/s12311-009-0100-1 • Patenaude, B., Smith, S. M., Kennedy, D. N., & Jenkinson, M. (2011). A Bayesian model of shape and appearance for subcortical brain segmentation. NeuroImage, 56(3), 907–922. https://doi.org/10.1016/j.neuroimage.2011.02.046 • Perani, D., & Abutalebi, J. (2015). Bilingualism, dementia, cognitive and neural reserve. Current Opinion in Neurology, 28(6), 618–625. https://doi.org/10.1097/WCO.0000000000000267 • Perani, D., Farsad, M., Ballarini, T., Lubian, F., Malpetti, M., Fracchetti, A., … Abutalebi, J. (2017). The impact of bilingualism on brain reserve and metabolic connectivity in Alzheimer’s dementia. Proceedings of the National Academy of Sciences, 114(7), 1690–1695. https://doi.org/10.1073/pnas.1610909114 • Pliatsikas, C., DeLuca, V., Moschopoulou, E., & Saddy, J. D. (2017). Immersive bilingualism reshapes the core of the brain. Brain Structure and Function, 222(4), 1785–1795. https://doi.org/10.1007/s00429-016-1307-9 • Pliatsikas, C., Johnstone, T., & Marinis, T. (2014a). fMRI evidence for the involvement of the procedural memory system in morphological processing of a second language. PLoS ONE, 9(5), e97298. https://doi.org/10.1371/journal.pone.0097298 • Pliatsikas, C., Johnstone, T., & Marinis, T. (2014b). Grey matter volume in the cerebellum is related to the processing of grammatical rules in a second language: A structural voxel-based morphometry study. The Cerebellum, 13(1), 55–63. https://doi.org/10.1007/s12311-013-0515-6 • Pliatsikas, C., & Luk, G. (2016). Executive control in bilinguals: A concise review on fMRI studies. Bilingualism: Language and Cognition, 53(9), 1689–1699. https://doi.org/10.1017/CBO9781107415324.004 • Pliatsikas, C., Moschopoulou, E., & Saddy, J. D. (2015). The effects of bilingualism on the white matter structure of the brain. Proceedings of the National Academy of Sciences, 112(5), 1334–1337. https://doi.org/10.1073/pnas.1414183112 • Prehn, K., Taud, B., Reifegerste, J., Harald, C., & Flöel, A. (2017). Neural correlates of grammatical inflection in older native and second-language speakers. Bilingualism: Language and Cognition, 1–12. https://doi.org/10.1017/S1366728916001206 • Rahmani, F., Sobhani, S., & Aarabi, M. H. (2017). Sequential language learning and language immersion in bilingualism: diffusion MRI connectometry reveals microstructural evidence. Experimental Brain Research, 235(10), 2935–2945. https://doi.org/10.1007/s00221-017-5029-x • Ressel, V., Pallier, C., Ventura-Campos, N., Díaz, B., Roessler, A., Ávila, C., & Sebastián-Gallés, N. (2012). An effect of bilingualism on the auditory cortex. The Journal of Neuroscience, 32(47), 16597–601. https://doi.org/10.1523/JNEUROSCI.1996-12.2012 • Richardson, F. M., Thomas, M. S. C., Filippi, R., Harth, H., & Price, C. J. (2010). Contrasting effects of vocabulary knowledge on temporal and parietal brain structure across lifespan. Journal of Cognitive Neuroscience, 22(5), 943–954. https://doi.org/10.1162/jocn.2009.21238 • Roberts, L., Gonzalez Alonso, J., Pliatsikas, C., & Rothman, J. (2018). Evidence from neurolinguistic methodologies: Can it actually inform linguistic/language acquisition theories and translate to evidence-based applications? Second Language Research, 34(1), 125–143. https://doi.org/10.1177/0267658316644010 • Rossi, E., Cheng, H., Kroll, J. F., Diaz, M. T., & Newman, S. D. (2017). Changes in white-matter connectivity in late second language learners: Evidence from diffusion tensor imaging. Frontiers in Psychology, 8, 1–15. https://doi.org/10.3389/fpsyg.2017.02040 • Schlegel, A. A., Rudelson, J. J., & Tse, P. U. (2012). White matter structure changes as adults learn a second language. Journal of Cognitive Neuroscience, 24(8), 1664–70. https://doi.org/10.1162/jocn_a_00240 • Schweizer, T. A., Ware, J., Fischer, C. E., Craik, F. I. M., & Bialystok, E. (2012). Bilingualism as a contributor to cognitive reserve: evidence from brain atrophy in Alzheimer’s disease. Cortex, 48(8), 991–6. https://doi.org/10.1016/j.cortex.2011.04.009 • Singh, N. C., Rajan, A., Malagi, A., Ramanujan, K., Canini, M., Della Rosa, P. A., … Abutalebi, J. (2017). Microstructural anatomical differences between bilinguals and monolinguals. Bilingualism: Language and Cognition, 1–14. https://doi.org/10.1017/S1366728917000438 • Smith, S. M., Jenkinson, M., Johansen-Berg, H., Rueckert, D., Nichols, T. E., Mackay, C. E., … Behrens, T. E. J. (2006). Tract-based spatial statistics: voxelwise analysis of multi-subject diffusion data. NeuroImage, 31(4), 1487–505. https://doi.org/10.1016/j.neuroimage.2006.02.024 • Stein, M., Federspiel, A., Koenig, T., Wirth, M., Lehmann, C., Wiest, R., … Dierks, T. (2009). Reduced frontal activation with increasing 2nd language proficiency. Neuropsychologia, 47(13), 2712–20. https://doi.org/10.1016/j.neuropsychologia.2009.05.023 • Stein, M., Federspiel, A., Koenig, T., Wirth, M., Strik, W., Wiest, R., … Dierks, T. (2012). Structural plasticity in the language system related to increased second language proficiency. Cortex, 48(4), 458–65. https://doi.org/10.1016/j.cortex.2010.10.007 • Valian, V. (2015). Bilingualism and cognition. Bilingualism: Language and Cognition, 18, 3–24. https://doi.org/10.1017/S1366728914000522 • Wei, M., Joshi, A. A., Zhang, M., Mei, L., Manis, F. R., He, Q., … Lu, Z.-L. (2015). How age of acquisition influences brain architecture in bilinguals. Journal of Neurolinguistics, 36, 35–55. https://doi.org/10.1016/j.jneuroling.2015.05.001 • Wong, P. C. M., Warrier, C. M., Penhune, V. B., Roy, A. K., Sadehh, A., Parrish, T. B., & Zatorre, R. J. (2008). Volume of left Heschl’s Gyrus and linguistic pitch learning. Cerebral Cortex, 18(4), 828–36. https://doi.org/10.1093/cercor/bhm115 • Xiang, H., van Leeuwen, T. M., Dediu, D., Roberts, L., Norris, D. G., & Hagoort, P. (2015). L2-proficiency-dependent laterality shift in structural connectivity of brain language pathways. Brain Connectivity, 5(6), 349–361. • Zou, L., Ding, G., Abutalebi, J., Shu, H., & Peng, D. (2012). Structural plasticity of the left caudate in bimodal bilinguals. Cortex, 48(9), 1197–206. https://doi.org/10.1016/j.cortex.2011.05.022