{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,3]],"date-time":"2026-06-03T02:26:38Z","timestamp":1780453598950,"version":"3.54.1"},"reference-count":73,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2021,4,6]],"date-time":"2021-04-06T00:00:00Z","timestamp":1617667200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2021,4,6]],"date-time":"2021-04-06T00:00:00Z","timestamp":1617667200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100001691","name":"MEXT | Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["26381350"],"award-info":[{"award-number":["26381350"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"MEXT | Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["17H06380"],"award-info":[{"award-number":["17H06380"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"MEXT | Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["26560274"],"award-info":[{"award-number":["26560274"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"MEXT | Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["16KT0005"],"award-info":[{"award-number":["16KT0005"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"MEXT | Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["19H03992"],"award-info":[{"award-number":["19H03992"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Commun Biol"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Handwriting is thought to impede vocabulary learning in sighted adults because the motor execution of writing interferes with efficient audiovisual processing during encoding. However, the motor memory of writing may facilitate adult word learning when visual sensory inputs are severely restricted. Using functional MRI, we show that late-blind participants, but not sighted participants, learned novel words by recruiting the left dorsal premotor cortex known as Exner\u2019s writing area and its functional coupling with the left hippocampus. During later recall, the phonological and semantic contents of these words are represented in the activation patterns of the left hippocampus as well as in those of left frontotemporal language areas. These findings suggest that motor codes of handwriting help blind participants maintain word-form representations during learning and retrieval. We propose that such reliance on the motor system reflects a broad architecture of the cerebral language network which encompasses the limb motor system as a hardwired component.<\/jats:p>","DOI":"10.1038\/s42003-021-01971-z","type":"journal-article","created":{"date-parts":[[2021,4,6]],"date-time":"2021-04-06T10:02:48Z","timestamp":1617703368000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Graphomotor memory in Exner\u2019s area enhances word learning in the blind"],"prefix":"10.1038","volume":"4","author":[{"given":"Tomomi","family":"Mizuochi-Endo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kazuyuki","family":"Itou","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3460-641X","authenticated-orcid":false,"given":"Michiru","family":"Makuuchi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Baku","family":"Kato","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kazuhisa","family":"Ikeda","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4834-2125","authenticated-orcid":false,"given":"Kimihiro","family":"Nakamura","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2021,4,6]]},"reference":[{"key":"1971_CR1","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/0167-9457(91)90003-G","volume":"10","author":"GP van Galen","year":"1991","unstructured":"van Galen, G. P. Handwriting: issues for a psychomotor theory. Hum. Mov. Sci. 10, 165\u2013191 (1991).","journal-title":"Hum. Mov. Sci."},{"key":"1971_CR2","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1080\/14640748408402172","volume":"36","author":"DI Margolin","year":"1984","unstructured":"Margolin, D. I. The neuropsychology of writing and spelling: semantic, phonological, motor, and perceptual processes. Q. J. Exp. Psychol. Sect. A 36, 459\u2013489 (1984).","journal-title":"Q. J. Exp. Psychol. Sect. A"},{"key":"1971_CR3","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.actpsy.2004.10.019","volume":"119","author":"M Longcamp","year":"2005","unstructured":"Longcamp, M., Zerbato-Poudou, M. T. & Velay, J. L. The influence of writing practice on letter recognition in preschool children: a comparison between handwriting and typing. Acta Psychol. 119, 67\u201379 (2005).","journal-title":"Acta Psychol."},{"key":"1971_CR4","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1037\/0022-0663.82.1.159","volume":"82","author":"AE Cunningham","year":"1990","unstructured":"Cunningham, A. E. & Stanovich, K. E. Early spelling acquisition: writing beats the computer. J. Educ. Psychol. 82, 159 (1990).","journal-title":"J. Educ. Psychol."},{"key":"1971_CR5","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1037\/xge0000134","volume":"145","author":"JX Li","year":"2016","unstructured":"Li, J. X. & James, K. H. Handwriting generates variable visual output to facilitate symbol learning. J. Exp. Psychol. Gen. 145, 298\u2013313 (2016).","journal-title":"J. Exp. Psychol. Gen."},{"key":"1971_CR6","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1037\/a0023730","volume":"103","author":"CQ Guan","year":"2011","unstructured":"Guan, C. Q., Liu, Y., Chan, D. H. L., Ye, F. & Perfetti, C. A. Writing strengthens orthography and alphabetic-coding strengthens phonology in learning to read Chinese. J. Educ. Psychol. 103, 509\u2013522 (2011).","journal-title":"J. Educ. Psychol."},{"key":"1971_CR7","doi-asserted-by":"publisher","first-page":"8781","DOI":"10.1073\/pnas.0503523102","volume":"102","author":"LH Tan","year":"2005","unstructured":"Tan, L. H., Spinks, J. A., Eden, G. F., Perfetti, C. A. & Siok, W. T. Reading depends on writing, in Chinese. Proc. Natl Acad. Sci. USA 102, 8781\u20138785 (2005).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1971_CR8","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.tine.2012.08.001","volume":"1","author":"KH James","year":"2012","unstructured":"James, K. H. & Engelhardt, L. The effects of handwriting experience on functional brain development in pre-literate children. Trends Neurosci. Educ. 1, 32\u201342 (2012).","journal-title":"Trends Neurosci. Educ."},{"key":"1971_CR9","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1016\/j.tics.2008.07.006","volume":"12","author":"L Shams","year":"2008","unstructured":"Shams, L. & Seitz, A. R. Benefits of multisensory learning. Trends Cogn. Sci. 12, 411\u2013417 (2008).","journal-title":"Trends Cogn. Sci."},{"key":"1971_CR10","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1016\/j.neuroscience.2019.08.004","volume":"416","author":"F Cao","year":"2019","unstructured":"Cao, F. et al. L1 reading experience influences L2 lexical learning: Spanish learning in Chinese speakers and English speakers. Neuroscience 416, 255\u2013267 (2019).","journal-title":"Neuroscience"},{"key":"1971_CR11","doi-asserted-by":"publisher","first-page":"487","DOI":"10.1191\/0267658306sr276oa","volume":"22","author":"J Barcroft","year":"2006","unstructured":"Barcroft, J. Can writing a new word detract from learning it? More negative effects of forced output during vocabulary learning. Second Lang. Res. 22, 487\u2013497 (2006).","journal-title":"Second Lang. Res."},{"key":"1971_CR12","doi-asserted-by":"publisher","first-page":"713","DOI":"10.1111\/j.1944-9720.2007.tb02889.x","volume":"40","author":"J Barcroft","year":"2007","unstructured":"Barcroft, J. Effects of word and fragment writing during L2 vocabulary learning. Foreign Lang. Ann. 40, 713\u2013726 (2007).","journal-title":"Foreign Lang. Ann."},{"key":"1971_CR13","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1191\/0267658304sr233oa","volume":"20","author":"J Barcroft","year":"2004","unstructured":"Barcroft, J. Effects of sentence writing in second language lexical acquisition. Second Lang. Res. 20, 303\u2013334 (2004).","journal-title":"Second Lang. Res."},{"key":"1971_CR14","doi-asserted-by":"publisher","first-page":"220","DOI":"10.1037\/0033-2909.116.2.220","volume":"116","author":"H Pashler","year":"1994","unstructured":"Pashler, H. Dual-task interference in simple tasks: data and theory. Psychol. Bull. 116, 220\u2013244 (1994).","journal-title":"Psychol. Bull."},{"key":"1971_CR15","doi-asserted-by":"publisher","first-page":"609","DOI":"10.1044\/2019_LSHSS-VOIA-18-0135","volume":"50","author":"D Colenbrander","year":"2019","unstructured":"Colenbrander, D., Miles, K. P. & Ricketts, J. To see or not to see: how does seeing spellings support vocabulary learning? Lang. Speech Hearing Serv. Sch. 50, 609\u2013628 (2019).","journal-title":"Lang. Speech Hearing Serv. Sch."},{"key":"1971_CR16","doi-asserted-by":"publisher","first-page":"1359","DOI":"10.1126\/science.1194140","volume":"330","author":"S Dehaene","year":"2010","unstructured":"Dehaene, S. et al. How learning to read changes the cortical networks for vision and language. Science 330, 1359\u20131364 (2010).","journal-title":"Science"},{"key":"1971_CR17","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1016\/j.neuropsychologia.2018.01.011","volume":"111","author":"N Landi","year":"2018","unstructured":"Landi, N. et al. Neural representations for newly learned words are modulated by overnight consolidation, reading skill, and age. Neuropsychologia 111, 133\u2013144 (2018).","journal-title":"Neuropsychologia"},{"issue":"Suppl 1","key":"1971_CR18","doi-asserted-by":"publisher","first-page":"S99","DOI":"10.1016\/j.dcn.2011.06.001","volume":"2","author":"AM Clements-Stephens","year":"2012","unstructured":"Clements-Stephens, A. M. et al. Neural circuitry associated with two different approaches to novel word learning. Dev. Cogn. Neurosci. 2(Suppl 1), S99\u2013S113 (2012).","journal-title":"Dev. Cogn. Neurosci."},{"key":"1971_CR19","doi-asserted-by":"crossref","unstructured":"Ardila, A. in Foundations of Bilingual Memory (eds Heredia, R. R. & Altarriba, J.) 223\u2013234 (Springer, 2014).","DOI":"10.1007\/978-1-4614-9218-4_11"},{"key":"1971_CR20","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1177\/1745691615621279","volume":"11","author":"N Cowan","year":"2016","unstructured":"Cowan, N. Working memory maturation: can we get at the essence of cognitive growth? Perspect. Psychol. Sci. 11, 239\u2013264 (2016).","journal-title":"Perspect. Psychol. Sci."},{"key":"1971_CR21","doi-asserted-by":"publisher","first-page":"1641","DOI":"10.1111\/j.1467-8624.2010.01499.x","volume":"81","author":"JR Best","year":"2010","unstructured":"Best, J. R. & Miller, P. H. A developmental perspective on executive function. Child Dev. 81, 1641\u20131660 (2010).","journal-title":"Child Dev."},{"key":"1971_CR22","doi-asserted-by":"publisher","first-page":"258","DOI":"10.1016\/j.neuroimage.2012.02.035","volume":"61","author":"K Monzalvo","year":"2012","unstructured":"Monzalvo, K., Fluss, J., Billard, C., Dehaene, S. & Dehaene-Lambertz, G. Cortical networks for vision and language in dyslexic and normal children of variable socio-economic status. Neuroimage 61, 258\u2013274 (2012).","journal-title":"Neuroimage"},{"key":"1971_CR23","doi-asserted-by":"publisher","first-page":"537","DOI":"10.1002\/ana.21804","volume":"66","author":"FE Roux","year":"2009","unstructured":"Roux, F. E. et al. The graphemic\/motor frontal area Exner\u2019s area revisited. Ann. Neurol. 66, 537\u2013545 (2009).","journal-title":"Ann. Neurol."},{"key":"1971_CR24","doi-asserted-by":"publisher","first-page":"982","DOI":"10.1002\/hbm.21084","volume":"32","author":"M Macedonia","year":"2011","unstructured":"Macedonia, M., Muller, K. & Friederici, A. D. The impact of iconic gestures on foreign language word learning and its neural substrate. Hum. Brain Mapp. 32, 982\u2013998 (2011).","journal-title":"Hum. Brain Mapp."},{"key":"1971_CR25","doi-asserted-by":"publisher","first-page":"3631","DOI":"10.1093\/cercor\/bhy237","volume":"29","author":"E Cracco","year":"2019","unstructured":"Cracco, E., Keysers, C., Clauwaert, A. & Brass, M. Representing multiple observed actions in the motor system. Cereb. Cortex 29, 3631\u20133641 (2019).","journal-title":"Cereb. Cortex"},{"key":"1971_CR26","doi-asserted-by":"publisher","first-page":"e0177599","DOI":"10.1371\/journal.pone.0177599","volume":"12","author":"Y Kanazawa","year":"2017","unstructured":"Kanazawa, Y. et al. Phonological memory in sign language relies on the visuomotor neural system outside the left hemisphere language network. PLoS ONE 12, e0177599 (2017).","journal-title":"PLoS ONE"},{"key":"1971_CR27","doi-asserted-by":"publisher","first-page":"2043","DOI":"10.1111\/cdev.12715","volume":"88","author":"M Havy","year":"2017","unstructured":"Havy, M., Foroud, A., Fais, L. & Werker, J. F. The role of auditory and visual speech in word learning at 18 months and in adulthood. Child Dev. 88, 2043\u20132059 (2017).","journal-title":"Child Dev."},{"key":"1971_CR28","doi-asserted-by":"publisher","first-page":"15","DOI":"10.3758\/s13423-019-01647-0","volume":"27","author":"G Meade","year":"2020","unstructured":"Meade, G. The role of phonology during visual word learning in adults: an integrative review. Psychonom. Bull. Rev. 27, 15\u201323 (2020).","journal-title":"Psychonom. Bull. Rev."},{"key":"1971_CR29","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1068\/p5390","volume":"36","author":"ML Noordzij","year":"2007","unstructured":"Noordzij, M. L., Zuidhoek, S. & Postma, A. The influence of visual experience on visual and spatial imagery. Perception 36, 101\u2013112 (2007).","journal-title":"Perception"},{"key":"1971_CR30","first-page":"159","volume":"8","author":"V Occelli","year":"2014","unstructured":"Occelli, V., Lin, J. B., Lacey, S. & Sathian, K. Loss of form vision impairs spatial imagery. Front. Hum. Neurosci. 8, 159 (2014).","journal-title":"Front. Hum. Neurosci."},{"key":"1971_CR31","doi-asserted-by":"publisher","first-page":"4","DOI":"10.3389\/neuro.01.016.2008","volume":"2","author":"N Kriegeskorte","year":"2008","unstructured":"Kriegeskorte, N., Mur, M. & Bandettini, P. Representational similarity analysis - connecting the branches of systems neuroscience. Front. Syst. Neurosci. 2, 4 (2008).","journal-title":"Front. Syst. Neurosci."},{"key":"1971_CR32","doi-asserted-by":"crossref","unstructured":"Wobbrock, J. O., Findlater, L., Gergle, D. & Higgins, J. J. The aligned rank transform for nonparametric factorial analyses using only anova procedures. In Proceedings of the SIGCHI Conference on Human Factors in Computing Systems 143\u2013146 (Association for Computing Machinery, 2011).","DOI":"10.1145\/1978942.1978963"},{"key":"1971_CR33","doi-asserted-by":"publisher","first-page":"2217","DOI":"10.1081\/STA-200026599","volume":"33","author":"H Mansouri","year":"2004","unstructured":"Mansouri, H., Paige, R. L. & Surles, J. G. Aligned rank transform techniques for analysis of variance and multiple comparisons. Commun. Stat. Theory Methods 33, 2217\u20132232 (2004).","journal-title":"Commun. Stat. Theory Methods"},{"key":"1971_CR34","doi-asserted-by":"publisher","first-page":"2772","DOI":"10.1016\/j.cortex.2013.05.011","volume":"49","author":"S Planton","year":"2013","unstructured":"Planton, S., Jucla, M., Roux, F. E. & Demonet, J. F. The \u201chandwriting brain\u201d: a meta-analysis of neuroimaging studies of motor versus orthographic processes. Cortex 49, 2772\u20132787 (2013).","journal-title":"Cortex"},{"key":"1971_CR35","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1016\/j.tics.2005.05.004","volume":"9","author":"S Dehaene","year":"2005","unstructured":"Dehaene, S., Cohen, L., Sigman, M. & Vinckier, F. The neural code for written words: a proposal. Trends Cogn. Sci. 9, 335\u2013341 (2005).","journal-title":"Trends Cogn. Sci."},{"key":"1971_CR36","doi-asserted-by":"publisher","first-page":"272","DOI":"10.1016\/S1364-6613(99)01333-9","volume":"3","author":"R Jackendoff","year":"1999","unstructured":"Jackendoff, R. Possible stages in the evolution of the language capacity. Trends Cogn. Sci. 3, 272\u2013279 (1999).","journal-title":"Trends Cogn. Sci."},{"key":"1971_CR37","doi-asserted-by":"publisher","first-page":"184","DOI":"10.1080\/15475441.2010.484379","volume":"6","author":"S Carey","year":"2010","unstructured":"Carey, S. Beyond fast mapping. Lang. Learn. Dev. 6, 184\u2013205 (2010).","journal-title":"Lang. Learn. Dev."},{"key":"1971_CR38","doi-asserted-by":"publisher","first-page":"16864","DOI":"10.1523\/JNEUROSCI.1376-10.2010","volume":"30","author":"Y Shtyrov","year":"2010","unstructured":"Shtyrov, Y., Nikulin, V. V. & Pulvermuller, F. Rapid cortical plasticity underlying novel word learning. J. Neurosci. 30, 16864\u201316867 (2010).","journal-title":"J. Neurosci."},{"key":"1971_CR39","doi-asserted-by":"publisher","first-page":"304","DOI":"10.3389\/fnhum.2019.00304","volume":"13","author":"MJ Vasilyeva","year":"2019","unstructured":"Vasilyeva, M. J., Knyazeva, V. M., Aleksandrov, A. A. & Shtyrov, Y. Neurophysiological correlates of fast mapping of novel words in the adult brain. Front. Hum. Neurosci. 13, 304 (2019).","journal-title":"Front. Hum. Neurosci."},{"key":"1971_CR40","doi-asserted-by":"publisher","first-page":"857","DOI":"10.1098\/rstb.2007.2093","volume":"362","author":"S Kouider","year":"2007","unstructured":"Kouider, S. & Dehaene, S. Levels of processing during non-conscious perception: a critical review of visual masking. Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci. 362, 857\u2013875 (2007).","journal-title":"Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci."},{"key":"1971_CR41","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1353\/aad.2017.0025","volume":"162","author":"CQ Guan","year":"2017","unstructured":"Guan, C. Q. & Wang, Y. The effect of handwriting training on language learning among deaf children and their matched hearing peers in China. Am. Ann. Deaf 162, 265\u2013276 (2017).","journal-title":"Am. Ann. Deaf"},{"key":"1971_CR42","doi-asserted-by":"publisher","first-page":"20762","DOI":"10.1073\/pnas.1217749109","volume":"109","author":"K Nakamura","year":"2012","unstructured":"Nakamura, K. et al. Universal brain systems for recognizing word shapes and handwriting gestures during reading. Proc. Natl Acad. Sci. USA 109, 20762\u201320767 (2012).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1971_CR43","doi-asserted-by":"publisher","first-page":"1492","DOI":"10.1016\/S1053-8119(03)00088-0","volume":"19","author":"M Longcamp","year":"2003","unstructured":"Longcamp, M., Anton, J. L., Roth, M. & Velay, J. L. Visual presentation of single letters activates a premotor area involved in writing. Neuroimage 19, 1492\u20131500 (2003).","journal-title":"Neuroimage"},{"key":"1971_CR44","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1016\/j.cub.2014.11.068","volume":"25","author":"KM Mayer","year":"2015","unstructured":"Mayer, K. M., Yildiz, I. B., Macedonia, M. & von Kriegstein, K. Visual and motor cortices differentially support the translation of foreign language words. Curr. Biol. 25, 530\u2013535 (2015).","journal-title":"Curr. Biol."},{"key":"1971_CR45","doi-asserted-by":"publisher","first-page":"10154","DOI":"10.1073\/pnas.1803667115","volume":"115","author":"R Vos de Wael","year":"2018","unstructured":"Vos de Wael, R. et al. Anatomical and microstructural determinants of hippocampal subfield functional connectome embedding. Proc. Natl Acad. Sci. USA 115, 10154\u201310159 (2018).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1971_CR46","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-37905-9","volume":"9","author":"JJ Maller","year":"2019","unstructured":"Maller, J. J. et al. Revealing the hippocampal connectome through super-resolution 1150-direction diffusion MRI. Sci. Rep. 9, 2418 (2019).","journal-title":"Sci. Rep."},{"key":"1971_CR47","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1007\/s002210000406","volume":"133","author":"EA Murray","year":"2000","unstructured":"Murray, E. A., Bussey, T. J. & Wise, S. P. Role of prefrontal cortex in a network for arbitrary visuomotor mapping. Exp. Brain Res. 133, 114\u2013129 (2000).","journal-title":"Exp. Brain Res."},{"key":"1971_CR48","doi-asserted-by":"publisher","first-page":"442","DOI":"10.1038\/86115","volume":"4","author":"SA Small","year":"2001","unstructured":"Small, S. A. et al. Circuit mechanisms underlying memory encoding and retrieval in the long axis of the hippocampal formation. Nat. Neurosci. 4, 442\u2013449 (2001).","journal-title":"Nat. Neurosci."},{"key":"1971_CR49","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1002\/hipo.10064","volume":"13","author":"MD Greicius","year":"2003","unstructured":"Greicius, M. D. et al. Regional analysis of hippocampal activation during memory encoding and retrieval: fMRI study. Hippocampus 13, 164\u2013174 (2003).","journal-title":"Hippocampus"},{"key":"1971_CR50","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1016\/j.neuron.2009.11.031","volume":"65","author":"MS Fanselow","year":"2010","unstructured":"Fanselow, M. S. & Dong, H.-W. Are the dorsal and ventral hippocampus functionally distinct structures? Neuron 65, 7\u201319 (2010).","journal-title":"Neuron"},{"key":"1971_CR51","doi-asserted-by":"publisher","first-page":"3773","DOI":"10.1098\/rstb.2009.0111","volume":"364","author":"MH Davis","year":"2009","unstructured":"Davis, M. H. & Gaskell, M. G. A complementary systems account of word learning: neural and behavioural evidence. Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci. 364, 3773\u20133800 (2009).","journal-title":"Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci."},{"key":"1971_CR52","doi-asserted-by":"publisher","first-page":"3711","DOI":"10.1098\/rstb.2009.0130","volume":"364","author":"A Rodriguez-Fornells","year":"2009","unstructured":"Rodriguez-Fornells, A., Cunillera, T., Mestres-Misse, A. & de Diego-Balaguer, R. Neurophysiological mechanisms involved in language learning in adults. Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci. 364, 3711\u20133735 (2009).","journal-title":"Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci."},{"key":"1971_CR53","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1146\/annurev.psych.55.090902.142050","volume":"55","author":"Y Dudai","year":"2004","unstructured":"Dudai, Y. The neurobiology of consolidations, or, how stable is the engram? Annu. Rev. Psychol. 55, 51\u201386 (2004).","journal-title":"Annu. Rev. Psychol."},{"key":"1971_CR54","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1016\/j.cognition.2003.11.002","volume":"92","author":"NF Dronkers","year":"2004","unstructured":"Dronkers, N. F., Wilkins, D. P., Van Valin, R. D. Jr, Redfern, B. B. & Jaeger, J. J. Lesion analysis of the brain areas involved in language comprehension. Cognition 92, 145\u2013177 (2004).","journal-title":"Cognition"},{"key":"1971_CR55","doi-asserted-by":"publisher","first-page":"14984","DOI":"10.1073\/pnas.0403766101","volume":"101","author":"JT Devlin","year":"2004","unstructured":"Devlin, J. T., Jamison, H. L., Matthews, P. M. & Gonnerman, L. M. Morphology and the internal structure of words. Proc. Natl Acad. Sci. USA 101, 14984\u201314988 (2004).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1971_CR56","doi-asserted-by":"publisher","first-page":"1081","DOI":"10.1002\/hbm.21093","volume":"32","author":"Z Ye","year":"2011","unstructured":"Ye, Z., Mestres-Misse, A., Rodriguez-Fornells, A. & Munte, T. F. Two distinct neural networks support the mapping of meaning to a novel word. Hum. Brain Mapp. 32, 1081\u20131090 (2011).","journal-title":"Hum. Brain Mapp."},{"key":"1971_CR57","doi-asserted-by":"publisher","first-page":"1231","DOI":"10.1007\/s00429-016-1273-2","volume":"222","author":"S Hofstetter","year":"2017","unstructured":"Hofstetter, S., Friedmann, N. & Assaf, Y. Rapid language-related plasticity: microstructural changes in the cortex after a short session of new word learning. Brain Struct. Funct. 222, 1231\u20131241 (2017).","journal-title":"Brain Struct. Funct."},{"key":"1971_CR58","doi-asserted-by":"publisher","first-page":"11101","DOI":"10.1523\/JNEUROSCI.1720-17.2017","volume":"37","author":"P Ripolles","year":"2017","unstructured":"Ripolles, P. et al. Strength of temporal white matter pathways predicts semantic learning. J. Neurosci. 37, 11101\u201311113 (2017).","journal-title":"J. Neurosci."},{"key":"1971_CR59","doi-asserted-by":"publisher","first-page":"2767","DOI":"10.1093\/cercor\/bhp055","volume":"19","author":"JR Binder","year":"2009","unstructured":"Binder, J. R., Desai, R. H., Graves, W. W. & Conant, L. L. Where is the semantic system? A critical review and meta-analysis of 120 functional neuroimaging studies. Cereb. Cortex 19, 2767\u20132796 (2009).","journal-title":"Cereb. Cortex"},{"key":"1971_CR60","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1076\/vimr.1.3.131.4438","volume":"1","author":"GE Legge","year":"1999","unstructured":"Legge, G. E., Madison, C. M. & Mansfield, J. S. Measuring Braille reading speed with the MNREAD test. Vis. Impair. Res. 1, 131\u2013145 (1999).","journal-title":"Vis. Impair. Res."},{"key":"1971_CR61","doi-asserted-by":"publisher","first-page":"2153","DOI":"10.1016\/j.ridd.2012.06.012","volume":"33","author":"A Veispak","year":"2012","unstructured":"Veispak, A., Boets, B. & Ghesquiere, P. Parallel versus sequential processing in print and braille reading. Res. Dev. Disabil. 33, 2153\u20132163 (2012).","journal-title":"Res. Dev. Disabil."},{"key":"1971_CR62","doi-asserted-by":"publisher","first-page":"802","DOI":"10.1162\/jocn.2008.20504","volume":"20","author":"M Longcamp","year":"2008","unstructured":"Longcamp, M. et al. Learning through hand- or typewriting influences visual recognition of new graphic shapes: behavioral and functional imaging evidence. J. Cogn. Neurosci. 20, 802\u2013815 (2008).","journal-title":"J. Cogn. Neurosci."},{"key":"1971_CR63","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1080\/09602011.2013.809661","volume":"23","author":"R Starrfelt","year":"2013","unstructured":"Starrfelt, R., Olafsdottir, R. R. & Arendt, I. M. Rehabilitation of pure alexia: a review. Neuropsychol. Rehabil. 23, 755\u2013779 (2013).","journal-title":"Neuropsychol. Rehabil."},{"key":"1971_CR64","doi-asserted-by":"publisher","first-page":"2914","DOI":"10.1016\/j.cortex.2013.02.002","volume":"49","author":"ME Nicholls","year":"2013","unstructured":"Nicholls, M. E., Thomas, N. A., Loetscher, T. & Grimshaw, G. M. The Flinders Handedness survey (FLANDERS): a brief measure of skilled hand preference. Cortex 49, 2914\u20132926 (2013).","journal-title":"Cortex"},{"key":"1971_CR65","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1177\/1073858411420299","volume":"18","author":"Y Shtyrov","year":"2012","unstructured":"Shtyrov, Y. Neural bases of rapid word learning. Neuroscientist 18, 312\u2013319 (2012).","journal-title":"Neuroscientist"},{"key":"1971_CR66","doi-asserted-by":"publisher","first-page":"11078","DOI":"10.1073\/pnas.1507104112","volume":"112","author":"DM Nielson","year":"2015","unstructured":"Nielson, D. M., Smith, T. A., Sreekumar, V., Dennis, S. & Sederberg, P. B. Human hippocampus represents space and time during retrieval of real-world memories. Proc. Natl Acad. Sci. USA 112, 11078\u201311083 (2015).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1971_CR67","doi-asserted-by":"publisher","first-page":"5018","DOI":"10.1002\/hbm.22987","volume":"36","author":"JL Robinson","year":"2015","unstructured":"Robinson, J. L. et al. Neurofunctional topography of the human hippocampus. Hum. Brain Mapp. 36, 5018\u20135037 (2015).","journal-title":"Hum. Brain Mapp."},{"key":"1971_CR68","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1002\/hbm.20124","volume":"25","author":"DJ Bolger","year":"2005","unstructured":"Bolger, D. J., Perfetti, C. A. & Schneider, W. Cross-cultural effect on the brain revisited: universal structures plus writing system variation. Hum. Brain Mapp. 25, 92\u2013104 (2005).","journal-title":"Hum. Brain Mapp."},{"key":"1971_CR69","doi-asserted-by":"publisher","first-page":"4965","DOI":"10.1523\/JNEUROSCI.4031-14.2015","volume":"35","author":"LS Glezer","year":"2015","unstructured":"Glezer, L. S., Kim, J., Rule, J., Jiang, X. & Riesenhuber, M. Adding words to the brain\u2019s visual dictionary: novel word learning selectively sharpens orthographic representations in the VWFA. J. Neurosci. 35, 4965\u20134972 (2015).","journal-title":"J. Neurosci."},{"key":"1971_CR70","doi-asserted-by":"publisher","first-page":"424","DOI":"10.3389\/fnhum.2017.00424","volume":"11","author":"J Qu","year":"2017","unstructured":"Qu, J. et al. Neural pattern similarity in the left IFG and fusiform is associated with novel word learning. Front. Hum. Neurosci. 11, 424 (2017).","journal-title":"Front. Hum. Neurosci."},{"key":"1971_CR71","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1006\/nimg.1997.0291","volume":"6","author":"KJ Friston","year":"1997","unstructured":"Friston, K. J. et al. Psychophysiological and modulatory interactions in neuroimaging. Neuroimage 6, 218\u2013229 (1997).","journal-title":"Neuroimage"},{"key":"1971_CR72","doi-asserted-by":"publisher","first-page":"1490","DOI":"10.1523\/JNEUROSCI.2999-15.2016","volume":"36","author":"RL Jackson","year":"2016","unstructured":"Jackson, R. L., Hoffman, P., Pobric, G. & Lambon Ralph, M. A. The semantic network at work and rest: differential connectivity of anterior temporal lobe subregions. J. Neurosci. 36, 1490\u20131501 (2016).","journal-title":"J. Neurosci."},{"key":"1971_CR73","doi-asserted-by":"publisher","first-page":"3863","DOI":"10.1073\/pnas.0600244103","volume":"103","author":"N Kriegeskorte","year":"2006","unstructured":"Kriegeskorte, N., Goebel, R. & Bandettini, P. Information-based functional brain mapping. Proc. Natl Acad. Sci. USA 103, 3863\u20133868 (2006).","journal-title":"Proc. Natl Acad. Sci. USA"}],"container-title":["Communications Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-01971-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-01971-z","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-01971-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,3]],"date-time":"2022-12-03T19:25:08Z","timestamp":1670095508000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s42003-021-01971-z"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,4,6]]},"references-count":73,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2021,12]]}},"alternative-id":["1971"],"URL":"https:\/\/doi.org\/10.1038\/s42003-021-01971-z","relation":{},"ISSN":["2399-3642"],"issn-type":[{"value":"2399-3642","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,4,6]]},"assertion":[{"value":"15 October 2020","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 March 2021","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 April 2021","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"443"}}