{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T17:20:40Z","timestamp":1790184040040,"version":"4.1.0"},"reference-count":91,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T00:00:00Z","timestamp":1773273600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T00:00:00Z","timestamp":1774915200000},"content-version":"vor","delay-in-days":19,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Transl Psychiatry"],"DOI":"10.1038\/s41398-026-03851-8","type":"journal-article","created":{"date-parts":[[2026,3,12]],"date-time":"2026-03-12T17:04:02Z","timestamp":1773335042000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Consistent cortico-basal ganglia-cerebellar activation in early motor sequence learning: a systematic review and meta-analysis"],"prefix":"10.1038","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-4960-1741","authenticated-orcid":false,"given":"Weiwei","family":"Chen","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tongyue","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-8063-3148","authenticated-orcid":false,"given":"Ji","family":"Luo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1191-0697","authenticated-orcid":false,"given":"Jing","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5547-3361","authenticated-orcid":false,"given":"Yating","family":"Lv","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,3,12]]},"reference":[{"key":"3851_CR1","volume-title":"Motor learning and performance: from principles to application","author":"RA Schmidt","year":"2020","unstructured":"Schmidt RA, Lee TD Motor learning and performance: from principles to application. Sixth edition. Champaign, IL: Human Kinetics; 2020.","edition":"Sixth edition"},{"key":"3851_CR2","doi-asserted-by":"publisher","first-page":"558","DOI":"10.1037\/0033-295X.105.3.558","volume":"105","author":"DB Willingham","year":"1998","unstructured":"Willingham DB. A neuropsychological theory of motor skill learning. Psychol Rev. 1998;105:558\u201384.","journal-title":"Psychol Rev"},{"key":"3851_CR3","doi-asserted-by":"publisher","first-page":"443","DOI":"10.1016\/j.neuron.2011.10.008","volume":"72","author":"E Dayan","year":"2011","unstructured":"Dayan E, Cohen LG. Neuroplasticity Subserving Motor Skill Learning. Neuron. 2011;72:443\u201354.","journal-title":"Neuron"},{"key":"3851_CR4","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.bbr.2008.11.012","volume":"199","author":"J Doyon","year":"2009","unstructured":"Doyon J, Bellec P, Amsel R, Penhune V, Monchi O, Carrier J, et al. Contributions of the basal ganglia and functionally related brain structures to motor learning. Behav Brain Res. 2009;199:61\u201375.","journal-title":"Behav Brain Res"},{"key":"3851_CR5","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2019.116387","volume":"207","author":"K Janacsek","year":"2020","unstructured":"Janacsek K, Shattuck KF, Tagarelli KM, Lum JAG, Turkeltaub PE, Ullman MT. Sequence learning in the human brain: A functional neuroanatomical meta-analysis of serial reaction time studies. NeuroImage. 2020;207:116387.","journal-title":"NeuroImage"},{"key":"3851_CR6","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/0010-0285(87)90002-8","volume":"19","author":"MJ Nissen","year":"1987","unstructured":"Nissen MJ, Bullemer P. Attentional requirements of learning: Evidence from performance measures. Cognit Psychol. 1987;19:1\u201332.","journal-title":"Cognit Psychol"},{"key":"3851_CR7","doi-asserted-by":"publisher","first-page":"10073","DOI":"10.1523\/JNEUROSCI.2747-07.2007","volume":"27","author":"EM Robertson","year":"2007","unstructured":"Robertson EM. The serial reaction time task: Implicit motor skill learning?. J Neurosci. 2007;27:10073\u20135.","journal-title":"J Neurosci"},{"key":"3851_CR8","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1162\/jocn.1995.7.4.497","volume":"7","author":"ST Grafton","year":"1995","unstructured":"Grafton ST, Hazeltine E, Ivry R. Functional mapping of sequence learning in normal humans. J Cogn Neurosci. 1995;7:497\u2013510.","journal-title":"J Cogn Neurosci"},{"key":"3851_CR9","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1002\/(SICI)1097-0193(1997)5:2<124::AID-HBM6>3.0.CO;2-5","volume":"5","author":"SL Rauch","year":"1997","unstructured":"Rauch SL, Whalen PJ, Savage CR, Curran T, Kendrick A, Brown HD, et al. Striatal recruitment during an implicit sequence learning task as measured by functional magnetic resonance imaging. Hum Brain Mapp. 1997;5:124\u201332.","journal-title":"Hum Brain Mapp"},{"key":"3851_CR10","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.nlm.2009.02.009","volume":"92","author":"LA Boyd","year":"2009","unstructured":"Boyd LA, Edwards JD, Siengsukon CS, Vidoni ED, Wessel BD, Linsdell MA. Motor sequence chunking is impaired by basal ganglia stroke. Neurobiol Learn Mem. 2009;92:35\u201344.","journal-title":"Neurobiol Learn Mem"},{"key":"3851_CR11","doi-asserted-by":"publisher","first-page":"636","DOI":"10.1016\/j.conb.2011.06.012","volume":"21","author":"JW Krakauer","year":"2011","unstructured":"Krakauer JW, Mazzoni P. Human sensorimotor learning: adaptation, skill, and beyond. Curr Opin Neurobiol. 2011;21:636\u201344.","journal-title":"Curr Opin Neurobiol"},{"key":"3851_CR12","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/j.conb.2006.03.008","volume":"16","author":"J Ashe","year":"2006","unstructured":"Ashe J, Lungu OV, Basford AT, Lu X. Cortical control of motor sequences. Curr Opin Neurobiol. 2006;16:213\u201321.","journal-title":"Curr Opin Neurobiol"},{"key":"3851_CR13","doi-asserted-by":"publisher","first-page":"252","DOI":"10.1016\/S0028-3932(02)00158-6","volume":"41","author":"J Doyon","year":"2003","unstructured":"Doyon J, Penhune V, Ungerleider LG. Distinct contribution of the cortico-striatal and cortico-cerebellar systems to motor skill learning. Neuropsychologia. 2003;41:252\u201362.","journal-title":"Neuropsychologia"},{"key":"3851_CR14","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1016\/j.bbr.2011.09.044","volume":"226","author":"VB Penhune","year":"2012","unstructured":"Penhune VB, Steele CJ. Parallel contributions of cerebellar, striatal and M1 mechanisms to motor sequence learning. Behav Brain Res. 2012;226:579\u201391.","journal-title":"Behav Brain Res"},{"key":"3851_CR15","doi-asserted-by":"publisher","first-page":"8332","DOI":"10.1523\/JNEUROSCI.5569-09.2010","volume":"30","author":"CJ Steele","year":"2010","unstructured":"Steele CJ, Penhune VB. Specific increases within global decreases: a functional magnetic resonance imaging investigation of five days of motor sequence learning. J Neurosci Off J Soc Neurosci. 2010;30:8332\u201341.","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"3851_CR16","doi-asserted-by":"publisher","first-page":"631","DOI":"10.1146\/annurev.neuro.24.1.631","volume":"24","author":"J Tanji","year":"2001","unstructured":"Tanji J. Sequential Organization of Multiple Movements: Involvement of Cortical Motor Areas. Annu Rev Neurosci. 2001;24:631\u201351.","journal-title":"Annu Rev Neurosci"},{"key":"3851_CR17","doi-asserted-by":"publisher","first-page":"e00801","DOI":"10.7554\/eLife.00801","volume":"2","author":"T Wiestler","year":"2013","unstructured":"Wiestler T, Diedrichsen J. Skill learning strengthens cortical representations of motor sequences. Elife. 2013;2:e00801.","journal-title":"Elife"},{"key":"3851_CR18","doi-asserted-by":"publisher","first-page":"5356","DOI":"10.1523\/JNEUROSCI.3880-04.2005","volume":"25","author":"RA Poldrack","year":"2005","unstructured":"Poldrack RA, Sabb FW, Foerde K, Tom SM, Asarnow RF, Bookheimer SY, et al. The neural correlates of motor skill automaticity. J Neurosci Off J Soc Neurosci. 2005;25:5356\u201364.","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"3851_CR19","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1016\/j.neuron.2008.02.008","volume":"58","author":"G Albouy","year":"2008","unstructured":"Albouy G, Sterpenich V, Balteau E, Vandewalle G, Desseilles M, Dang-Vu T, et al. Both the hippocampus and striatum are involved in consolidation of motor sequence memory. Neuron. 2008;58:261\u201372.","journal-title":"Neuron"},{"key":"3851_CR20","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1046\/j.1528-1157.43.s.9.4.x","volume":"43","author":"O Hikosaka","year":"2002","unstructured":"Hikosaka O. A new approach to the functional systems of the brain. Epilepsia. 2002;43:9\u201315.","journal-title":"Epilepsia"},{"key":"3851_CR21","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.brainres.2009.12.073","volume":"1318","author":"L Ma","year":"2010","unstructured":"Ma L, Wang B, Narayana S, Hazeltine E, Chen X, Robin DA, et al. Changes in regional activity are accompanied with changes in inter-regional connectivity during 4 weeks motor learning. Brain Res. 2010;1318:64\u201376.","journal-title":"Brain Res"},{"key":"3851_CR22","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1146\/annurev.neuro.25.112701.142937","volume":"25","author":"MG Packard","year":"2002","unstructured":"Packard MG, Knowlton BJ. Learning and Memory Functions of the Basal Ganglia. Annu Rev Neurosci. 2002;25:563\u201393.","journal-title":"Annu Rev Neurosci"},{"key":"3851_CR23","doi-asserted-by":"crossref","unstructured":"Gamble KR, Cummings Jr. TJ, Lo SE, Ghosh PT, Howard Jr. JH, Howard DV. Implicit sequence learning in people with Parkinson\u2019s disease. Front Hum Neurosci. 2014;8.","DOI":"10.3389\/fnhum.2014.00563"},{"key":"3851_CR24","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1016\/j.brainres.2006.12.057","volume":"1137","author":"RD Seidler","year":"2007","unstructured":"Seidler RD, Tuite P, Ashe J. Selective impairments in implicit learning in Parkinson\u2019s disease. Brain Res. 2007;1137:104\u201310.","journal-title":"Brain Res"},{"key":"3851_CR25","doi-asserted-by":"publisher","first-page":"512","DOI":"10.1152\/jn.00717.2004","volume":"94","author":"A Floyer-Lea","year":"2005","unstructured":"Floyer-Lea A, Matthews PM. Distinguishable brain activation networks for short- and long-term motor skill learning. J Neurophysiol. 2005;94:512\u20138.","journal-title":"J Neurophysiol"},{"key":"3851_CR26","doi-asserted-by":"publisher","first-page":"2043","DOI":"10.1126\/science.1068524","volume":"296","author":"RD Seidler","year":"2002","unstructured":"Seidler RD, Purushotham A, Kim SG, Ugurbil K, Willingham D, Ashe J. Cerebellum activation associated with performance change but not motor learning. Science. 2002;296:2043\u20136.","journal-title":"Science"},{"key":"3851_CR27","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1016\/j.neuroimage.2012.11.020","volume":"67","author":"RM Hardwick","year":"2013","unstructured":"Hardwick RM, Rottschy C, Miall RC, Eickhoff SB. A quantitative meta-analysis and review of motor learning in the human brain. NeuroImage. 2013;67:283\u201397.","journal-title":"NeuroImage"},{"key":"3851_CR28","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1016\/j.eurpsy.2011.04.001","volume":"27","author":"J Radua","year":"2012","unstructured":"Radua J, Mataix-Cols D, Phillips ML, El-Hage W, Kronhaus DM, Cardoner N, et al. A new meta-analytic method for neuroimaging studies that combines reported peak coordinates and statistical parametric maps. Eur Psychiatry. 2012;27:605\u201311.","journal-title":"Eur Psychiatry"},{"key":"3851_CR29","doi-asserted-by":"publisher","first-page":"13","DOI":"10.3389\/fpsyt.2014.00013","volume":"5","author":"J Radua","year":"2014","unstructured":"Radua J, Rubia K, Canales-Rodr\u00edguez EJ, Pomarol-Clotet E, Fusar-Poli P, Mataix-Cols D. Anisotropic kernels for coordinate-based meta-analyses of neuroimaging studies. Front Psychiatry. 2014;5:13.","journal-title":"Front Psychiatry"},{"key":"3851_CR30","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/j.neubiorev.2017.11.012","volume":"84","author":"VI M\u00fcller","year":"2018","unstructured":"M\u00fcller VI, Cieslik EC, Laird AR, Fox PT, Radua J, Mataix-Cols D, et al. Ten simple rules for neuroimaging meta-analysis. Neurosci Biobehav Rev. 2018;84:151\u201361.","journal-title":"Neurosci Biobehav Rev"},{"key":"3851_CR31","doi-asserted-by":"publisher","first-page":"159","DOI":"10.2307\/2529310","volume":"33","author":"JR Landis","year":"1977","unstructured":"Landis JR, Koch GG. The measurement of observer agreement for categorical data. Biometrics. 1977;33:159.","journal-title":"Biometrics"},{"key":"3851_CR32","doi-asserted-by":"publisher","first-page":"1194","DOI":"10.1002\/hbm.20345","volume":"28","author":"JL Lancaster","year":"2007","unstructured":"Lancaster JL, Tordesillas-Guti\u00e9rrez D, Martinez M, Salinas F, Evans A, Zilles K, et al. Bias between MNI and Talairach coordinates analyzed using the ICBM-152 brain template. Hum Brain Mapp. 2007;28:1194\u2013205.","journal-title":"Hum Brain Mapp"},{"key":"3851_CR33","doi-asserted-by":"publisher","first-page":"677","DOI":"10.1016\/j.neuroimage.2010.02.048","volume":"51","author":"AR Laird","year":"2010","unstructured":"Laird AR, Robinson JL, McMillan KM, Tordesillas-Guti\u00e9rrez D, Moran ST, Gonzales SM, et al. Comparison of the disparity between Talairach and MNI coordinates in functional neuroimaging data: Validation of the Lancaster transform. NeuroImage. 2010;51:677\u201383.","journal-title":"NeuroImage"},{"key":"3851_CR34","doi-asserted-by":"publisher","first-page":"e68910","DOI":"10.1371\/journal.pone.0068910","volume":"8","author":"M Xia","year":"2013","unstructured":"Xia M, Wang J, He Y. BrainNet viewer: A network visualization tool for human brain connectomics. PLoS ONE. 2013;8:e68910.","journal-title":"PLoS ONE"},{"key":"3851_CR35","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1038\/nrn3475","volume":"14","author":"KS Button","year":"2013","unstructured":"Button KS, Ioannidis JPA, Mokrysz C, Nosek BA, Flint J, Robinson ESJ, et al. Power failure: why small sample size undermines the reliability of neuroscience. Nat Rev Neurosci. 2013;14:365\u201376.","journal-title":"Nat Rev Neurosci"},{"key":"3851_CR36","doi-asserted-by":"publisher","first-page":"e124","DOI":"10.1371\/journal.pmed.0020124","volume":"2","author":"JPA Ioannidis","year":"2005","unstructured":"Ioannidis JPA. Why most published research findings are false. PLoS Med. 2005;2:e124.","journal-title":"PLoS Med"},{"key":"3851_CR37","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1016\/j.neuroimage.2006.11.054","volume":"35","author":"B Thirion","year":"2007","unstructured":"Thirion B, Pinel P, M\u00e9riaux S, Roche A, Dehaene S, Poline J-B. Analysis of a large fMRI cohort: Statistical and methodological issues for group analyses. NeuroImage. 2007;35:105\u201320.","journal-title":"NeuroImage"},{"key":"3851_CR38","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1136\/bmj.315.7109.629","volume":"315","author":"M Egger","year":"1997","unstructured":"Egger M, Smith GD, Schneider M, Minder C. Bias in meta-analysis detected by a simple, graphical test. BMJ. 1997;315:629\u201334.","journal-title":"BMJ"},{"key":"3851_CR39","doi-asserted-by":"publisher","first-page":"83","DOI":"10.1016\/j.neuroimage.2008.03.061","volume":"44","author":"S Smith","year":"2009","unstructured":"Smith S, Nichols T. Threshold-free cluster enhancement: Addressing problems of smoothing, threshold dependence and localisation in cluster inference. NeuroImage. 2009;44:83\u201398.","journal-title":"NeuroImage"},{"key":"3851_CR40","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1016\/j.neuroimage.2016.05.068","volume":"141","author":"AM Winkler","year":"2016","unstructured":"Winkler AM, Ridgway GR, Douaud G, Nichols TE, Smith SM. Faster permutation inference in brain imaging. NeuroImage. 2016;141:502\u201316.","journal-title":"NeuroImage"},{"key":"3851_CR41","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1007\/s12021-016-9299-4","volume":"14","author":"CG Yan","year":"2016","unstructured":"Yan CG, Wang XD, Zuo XN, Zang YF. DPABI: Data processing & analysis for (Resting-State) brain imaging. Neuroinformatics. 2016;14:339\u201351.","journal-title":"Neuroinformatics"},{"key":"3851_CR42","doi-asserted-by":"publisher","first-page":"1013","DOI":"10.1016\/S0197-4580(03)00030-7","volume":"24","author":"SM Daselaar","year":"2003","unstructured":"Daselaar SM, Rombouts SARB, Veltman DJ, Raaijmakers JGW, Jonker C. Similar network activated by young and old adults during the acquisition of a motor sequence. Neurobiol Aging. 2003;24:1013\u20139.","journal-title":"Neurobiol Aging"},{"key":"3851_CR43","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2021.107508","volume":"185","author":"AB Fitzroy","year":"2021","unstructured":"Fitzroy AB, Kainec KA, Seo J, Spencer RMC. Encoding and consolidation of motor sequence learning in young and older adults. Neurobiol Learn Mem. 2021;185:107508.","journal-title":"Neurobiol Learn Mem"},{"key":"3851_CR44","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1016\/j.bbr.2016.11.021","volume":"319","author":"HT Karim","year":"2017","unstructured":"Karim HT, Huppert TJ, Erickson KI, Wollam ME, Sparto PJ, Sejdic E, et al. Motor sequence learning-induced neural efficiency in functional brain connectivity. Behav Brain Res. 2017;319:87\u201395.","journal-title":"Behav Brain Res"},{"key":"3851_CR45","doi-asserted-by":"publisher","first-page":"246","DOI":"10.3758\/CABN.6.3.246","volume":"6","author":"SM Landau","year":"2006","unstructured":"Landau SM, D\u2019Esposito M. Sequence learning in pianists and nonpianists: an fMRI study of motor expertise. Cogn Affect Behav Neurosci. 2006;6:246\u201359.","journal-title":"Cogn Affect Behav Neurosci"},{"key":"3851_CR46","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1176\/appi.ajp.161.1.67","volume":"161","author":"B Martis","year":"2004","unstructured":"Martis B, Wright CI, McMullin KG, Shin LM, Rauch SL. Functional magnetic resonance imaging evidence for a lack of striatal dysfunction during implicit sequence learning in individuals with animal phobia. Am J Psychiatry. 2004;161:67\u201371.","journal-title":"Am J Psychiatry"},{"key":"3851_CR47","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1016\/j.biopsych.2004.06.007","volume":"56","author":"R-A Muller","year":"2004","unstructured":"Muller R-A, Cauich C, Rubio MA, Mizuno A, Courchesne E. Abnormal activity patterns in premotor cortex during sequence learning in autistic patients. Biol Psychiatry. 2004;56:323\u201332.","journal-title":"Biol Psychiatry"},{"key":"3851_CR48","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1016\/j.jad.2010.02.114","volume":"125","author":"SL Naismith","year":"2010","unstructured":"Naismith SL, Lagopoulos J, Ward PB, Davey CG, Little C, Hickie IB. Fronto-striatal correlates of impaired implicit sequence learning in major depression: an fMRI study. J Affect Disord. 2010;125:256\u201361.","journal-title":"J Affect Disord"},{"key":"3851_CR49","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/j.neures.2005.01.008","volume":"52","author":"K Oishi","year":"2005","unstructured":"Oishi K, Toma K, Bagarinao ET, Matsuo K, Nakai T, Chihara K, et al. Activation of the precuneus is related to reduced reaction time in serial reaction time tasks. Neurosci Res. 2005;52:37\u201345.","journal-title":"Neurosci Res"},{"key":"3851_CR50","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/S0926-6410(02)00283-5","volume":"16","author":"K Werheid","year":"2003","unstructured":"Werheid K, Zysset S, Muller A, Reuter M, von Cramon DY. Rule learning in a serial reaction time task: an fMRI study on patients with early Parkinson\u2019s disease. Brain Res Cogn Brain Res. 2003;16:273\u201384.","journal-title":"Brain Res Cogn Brain Res"},{"key":"3851_CR51","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1016\/j.schres.2007.04.026","volume":"94","author":"ND Woodward","year":"2007","unstructured":"Woodward ND, Tibbo P, Purdon SE. An fMRI investigation of procedural learning in unaffected siblings of individuals with schizophrenia. Schizophr Res. 2007;94:306\u201316.","journal-title":"Schizophr Res"},{"key":"3851_CR52","doi-asserted-by":"publisher","first-page":"198","DOI":"10.1016\/j.schres.2006.06.039","volume":"88","author":"L Zedkova","year":"2006","unstructured":"Zedkova L, Woodward ND, Harding I, Tibbo PG, Purdon SE. Procedural learning in schizophrenia investigated with functional magnetic resonance imaging. Schizophr Res. 2006;88:198\u2013207.","journal-title":"Schizophr Res"},{"key":"3851_CR53","doi-asserted-by":"publisher","first-page":"492","DOI":"10.3389\/fnhum.2013.00492","volume":"7","author":"AL Jouen","year":"2013","unstructured":"Jouen AL, Verwey WB, van der Helden J, Scheiber C, Neveu R, Dominey PF, et al. Discrete sequence production with and without a pause: the role of cortex, basal ganglia, and cerebellum. Front Hum Neurosci. 2013;7:492.","journal-title":"Front Hum Neurosci"},{"key":"3851_CR54","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1016\/j.neuroimage.2005.04.025","volume":"27","author":"MG Lacourse","year":"2005","unstructured":"Lacourse MG, Orr EL, Cramer SC, Cohen MJ. Brain activation during execution and motor imagery of novel and skilled sequential hand movements. NeuroImage. 2005;27:505\u201319.","journal-title":"NeuroImage"},{"key":"3851_CR55","doi-asserted-by":"publisher","first-page":"2007","DOI":"10.1016\/j.neuroimage.2012.05.015","volume":"62","author":"CH Lin","year":"2012","unstructured":"Lin CH, Chiang MC, Wu AD, Iacoboni M, Udompholkul P, Yazdanshenas O, et al. Age related differences in the neural substrates of motor sequence learning after interleaved and repetitive practice. NeuroImage. 2012;62:2007\u201320.","journal-title":"NeuroImage"},{"key":"3851_CR56","doi-asserted-by":"publisher","first-page":"4213","DOI":"10.1093\/cercor\/bhu144","volume":"25","author":"NF Wymbs","year":"2015","unstructured":"Wymbs NF, Grafton ST. The human motor system supports sequence-specific representations over multiple training-dependent timescales. Cereb Cortex. 2015;25:4213\u201325.","journal-title":"Cereb Cortex"},{"key":"3851_CR57","doi-asserted-by":"publisher","first-page":"837","DOI":"10.1007\/s12264-025-01375-7","volume":"41","author":"J Chen","year":"2025","unstructured":"Chen J, Fan Y, Jia X, Fan F, Wang J, Zou Q, et al. The supplementary motor area as a flexible hub mediating behavioral and neuroplastic changes in motor sequence learning: A TMS and TMS-EEG study. Neurosci Bull. 2025;41:837\u201352.","journal-title":"Neurosci Bull"},{"key":"3851_CR58","doi-asserted-by":"publisher","first-page":"250","DOI":"10.1016\/j.neuroimage.2012.04.060","volume":"62","author":"S Narayana","year":"2012","unstructured":"Narayana S, Laird AR, Tandon N, Franklin C, Lancaster JL, Fox PT. Electrophysiological and functional connectivity of the human supplementary motor area. NeuroImage. 2012;62:250\u201365.","journal-title":"NeuroImage"},{"key":"3851_CR59","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1093\/cercor\/bhh198","volume":"15","author":"O Oullier","year":"2005","unstructured":"Oullier O, Jantzen KJ, Steinberg FL, Kelso JAS. Neural substrates of real and imagined sensorimotor coordination. Cereb Cortex. 2005;15:975\u201385.","journal-title":"Cereb Cortex"},{"key":"3851_CR60","doi-asserted-by":"publisher","first-page":"608","DOI":"10.1016\/j.neuroimage.2018.02.014","volume":"172","author":"D S\u00e4fstr\u00f6m","year":"2018","unstructured":"S\u00e4fstr\u00f6m D, Domell\u00f6f E. Brain activations supporting linking of action phases in a sequential manual task. NeuroImage. 2018;172:608\u201319.","journal-title":"NeuroImage"},{"key":"3851_CR61","doi-asserted-by":"publisher","first-page":"2159","DOI":"10.1093\/brain\/121.11.2159","volume":"121","author":"M Honda","year":"1998","unstructured":"Honda M. Dynamic cortical involvement in implicit and explicit motor sequence learning. A PET study. Brain. 1998;121:2159\u201373.","journal-title":"Brain"},{"key":"3851_CR62","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.cortex.2019.10.014","volume":"124","author":"M Harpaintner","year":"2020","unstructured":"Harpaintner M, Sim E-J, Trumpp NM, Ulrich M, Kiefer M. The grounding of abstract concepts in the motor and visual system: An fMRI study. Cortex. 2020;124:1\u201322.","journal-title":"Cortex"},{"key":"3851_CR63","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1007\/s00429-021-02258-z","volume":"226","author":"LC Triarhou","year":"2021","unstructured":"Triarhou LC. Cytoarchitectonics of the Rolandic operculum: morphofunctional ponderings. Brain Struct Funct. 2021;226:941\u201350.","journal-title":"Brain Struct Funct"},{"key":"3851_CR64","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1016\/j.cortex.2015.10.004","volume":"74","author":"A Steel","year":"2016","unstructured":"Steel A, Song S, Bageac D, Knutson KM, Keisler A, Saad ZS, et al. Shifts in connectivity during procedural learning after motor cortex stimulation: A combined transcranial magnetic stimulation\/functional magnetic resonance imaging study. Cortex. 2016;74:134\u201348.","journal-title":"Cortex"},{"key":"3851_CR65","doi-asserted-by":"publisher","first-page":"500","DOI":"10.1123\/mcj.8.4.500","volume":"8","author":"HI van Mier","year":"2004","unstructured":"van Mier HI, Perlmutter JS, Petersen SE. Functional changes in brain activity during acquisition and practice of movement sequences. Motor Control. 2004;8:500\u201320.","journal-title":"Motor Control"},{"key":"3851_CR66","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1016\/j.cortex.2015.06.012","volume":"71","author":"L Wilkinson","year":"2015","unstructured":"Wilkinson L, Steel A, Mooshagian E, Zimmermann T, Keisler A, Lewis JD, et al. Online feedback enhances early consolidation of motor sequence learning and reverses recall deficit from transcranial stimulation of motor cortex. Cortex. 2015;71:134\u201347.","journal-title":"Cortex"},{"key":"3851_CR67","doi-asserted-by":"publisher","first-page":"11515","DOI":"10.1523\/JNEUROSCI.6759-10.2011","volume":"31","author":"X Zhang","year":"2011","unstructured":"Zhang X, De Beukelaar TT, Possel J, Olaerts M, Swinnen SP, Woolley DG, et al. Movement observation improves early consolidation of motor memory. J Neurosci. 2011;31:11515\u201320.","journal-title":"J Neurosci"},{"key":"3851_CR68","doi-asserted-by":"crossref","unstructured":"Berlot E, Popp NJ, Diedrichsen J. A critical re-evaluation of fMRI signatures of motor sequence learning. Elife 2020;9.","DOI":"10.7554\/eLife.55241"},{"key":"3851_CR69","doi-asserted-by":"publisher","first-page":"7649","DOI":"10.1523\/JNEUROSCI.0863-21.2021","volume":"41","author":"E Berlot","year":"2021","unstructured":"Berlot E, Popp NJ, Grafton ST, Diedrichsen J. Combining repetition suppression and pattern analysis provides new insights into the role of M1 and parietal areas in skilled sequential actions. J Neurosci Off J Soc Neurosci. 2021;41:7649\u201361.","journal-title":"J Neurosci Off J Soc Neurosci"},{"key":"3851_CR70","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1038\/377155a0","volume":"377","author":"A Karni","year":"1995","unstructured":"Karni A, Meyer G, Jezzard P, Adams MM, Turner R, Ungerleider LG. Functional MRI evidence for adult motor cortex plasticity during motor skill learning. Nature. 1995;377:155\u20138.","journal-title":"Nature"},{"key":"3851_CR71","doi-asserted-by":"publisher","first-page":"414","DOI":"10.1016\/j.jphysparis.2006.03.007","volume":"99","author":"U Halsband","year":"2006","unstructured":"Halsband U, Lange RK. Motor learning in man: A review of functional and clinical studies. J Physiol-Paris. 2006;99:414\u201324.","journal-title":"J Physiol-Paris"},{"key":"3851_CR72","doi-asserted-by":"publisher","first-page":"1303","DOI":"10.1016\/j.neuroimage.2010.01.015","volume":"50","author":"A Rieckmann","year":"2010","unstructured":"Rieckmann A, Fischer H, Backman L. Activation in striatum and medial temporal lobe during sequence learning in younger and older adults: Relations to performance. NeuroImage. 2010;50:1303\u201312.","journal-title":"NeuroImage"},{"key":"3851_CR73","doi-asserted-by":"publisher","first-page":"204","DOI":"10.1037\/a0032012","volume":"127","author":"HE Schendan","year":"2013","unstructured":"Schendan HE, Tinaz S, Maher SM, Stern CE. Frontostriatal and mediotemporal lobe contributions to implicit higher-order spatial sequence learning declines in aging and Parkinson\u2019s disease. Behav Neurosci. 2013;127:204\u201321.","journal-title":"Behav Neurosci"},{"key":"3851_CR74","doi-asserted-by":"publisher","first-page":"338","DOI":"10.1038\/s41583-018-0002-7","volume":"19","author":"AC Bostan","year":"2018","unstructured":"Bostan AC, Strick PL. The basal ganglia and the cerebellum: nodes in an integrated network. Nat Rev Neurosci. 2018;19:338\u201350.","journal-title":"Nat Rev Neurosci"},{"key":"3851_CR75","doi-asserted-by":"publisher","first-page":"630","DOI":"10.1212\/WNL.58.4.630","volume":"58","author":"VS Mattay","year":"2002","unstructured":"Mattay VS, Fera F, Tessitore A, Hariri AR, Das S, Callicott JH, et al. Neurophysiological correlates of age-related changes in human motor function. Neurology. 2002;58:630\u20135.","journal-title":"Neurology"},{"key":"3851_CR76","doi-asserted-by":"publisher","first-page":"5255","DOI":"10.1002\/hbm.25191","volume":"41","author":"JA Bernard","year":"2020","unstructured":"Bernard JA, Nguyen AD, Hausman HK, Maldonado T, Ballard HK, Jackson TB, et al. Shaky scaffolding: Age differences in cerebellar activation revealed through activation likelihood estimation meta-analysis. Hum Brain Mapp. 2020;41:5255\u201381.","journal-title":"Hum Brain Mapp"},{"key":"3851_CR77","doi-asserted-by":"publisher","first-page":"270","DOI":"10.1002\/hbm.24793","volume":"41","author":"C Dahms","year":"2020","unstructured":"Dahms C, Brodoehl S, Witte O, Klingner C. The importance of different learning stages for motor sequence learning after stroke. Hum BRAIN Mapp. 2020;41:270\u201386.","journal-title":"Hum BRAIN Mapp"},{"key":"3851_CR78","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1016\/S1053-8119(03)00059-4","volume":"19","author":"RS Turner","year":"2003","unstructured":"Turner RS, Grafton ST, McIntosh AR, DeLong MR, Hoffman JM. The functional anatomy of parkinsonian bradykinesia. NeuroImage. 2003;19:163\u201379.","journal-title":"NeuroImage"},{"key":"3851_CR79","doi-asserted-by":"publisher","DOI":"10.1016\/j.nicl.2024.103601","volume":"42","author":"C Dahms","year":"2024","unstructured":"Dahms C, Noll A, Wagner F, Schmidt A, Brodoehl S, Klingner CM. Connecting the dots: Motor and default mode network crossroads in post-stroke motor learning deficits. Neuroimage Clin. 2024;42:103601.","journal-title":"Neuroimage Clin"},{"key":"3851_CR80","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1162\/jocn_a_00890","volume":"28","author":"A Dovern","year":"2016","unstructured":"Dovern A, Fink GR, Timpert DC, Saliger J, Karbe H, Weiss PH, et al. Timing matters? learning of complex spatiotemporal sequences in left-hemisphere stroke patients. J Cogn Neurosci. 2016;28:223\u201336.","journal-title":"J Cogn Neurosci"},{"key":"3851_CR81","doi-asserted-by":"publisher","first-page":"174","DOI":"10.1038\/s41593-017-0054-4","volume":"21","author":"R Polan\u00eda","year":"2018","unstructured":"Polan\u00eda R, Nitsche MA, Ruff CC. Studying and modifying brain function with non-invasive brain stimulation. Nat Neurosci. 2018;21:174\u201387.","journal-title":"Nat Neurosci"},{"key":"3851_CR82","doi-asserted-by":"publisher","first-page":"1071","DOI":"10.1016\/j.clinph.2015.02.001","volume":"126","author":"PM Rossini","year":"2015","unstructured":"Rossini PM, Burke D, Chen R, Cohen LG, Daskalakis Z, Di Iorio R, et al. Non-invasive electrical and magnetic stimulation of the brain, spinal cord, roots and peripheral nerves: Basic principles and procedures for routine clinical and research application. An updated report from an I.F.C.N. Committee. Clin Neurophysiol. 2015;126:1071\u2013107.","journal-title":"Clin Neurophysiol"},{"key":"3851_CR83","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2019.116424","volume":"208","author":"G Koch","year":"2020","unstructured":"Koch G, Esposito R, Motta C, Casula EP, Di Lorenzo F, Bonn\u00ec S, et al. Improving visuo-motor learning with cerebellar theta burst stimulation: Behavioral and neurophysiological evidence. NeuroImage. 2020;208:116424.","journal-title":"NeuroImage"},{"key":"3851_CR84","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1016\/j.brs.2019.08.001","volume":"13","author":"T Shimizu","year":"2020","unstructured":"Shimizu T, Hanajima R, Shirota Y, Tsutsumi R, Tanaka N, Terao Y, et al. Plasticity induction in the pre-supplementary motor area (pre-SMA) and SMA-proper differentially affects visuomotor sequence learning. Brain Stimulat. 2020;13:229\u201338.","journal-title":"Brain Stimulat"},{"key":"3851_CR85","doi-asserted-by":"crossref","unstructured":"Thong S, Doery E, Biabani M, Rogasch N, Chong T, Hendrikse J, et al. Disinhibition across secondary motor cortical regions during motor sequence learning: A TMS-EEG STUDY. J Neurosci. 2025;45.","DOI":"10.1101\/2024.02.26.581077"},{"key":"3851_CR86","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2020.117449","volume":"224","author":"G Hartwigsen","year":"2021","unstructured":"Hartwigsen G, Volz LJ. Probing rapid network reorganization of motor and language functions via neuromodulation and neuroimaging. NeuroImage. 2021;224:117449.","journal-title":"NeuroImage"},{"key":"3851_CR87","doi-asserted-by":"crossref","unstructured":"Kim YK, Shin SH. Comparison of effects of transcranial magnetic stimulation on primary motor cortex and supplementary motor area in motor skill learning (randomized, cross over study). Front Hum Neurosci 2014;8.","DOI":"10.3389\/fnhum.2014.00937"},{"key":"3851_CR88","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.brs.2012.02.005","volume":"6","author":"ZD Deng","year":"2013","unstructured":"Deng ZD, Lisanby SH, Peterchev AV. Electric field depth\u2013focality tradeoff in transcranial magnetic stimulation: Simulation comparison of 50 coil designs. Brain Stimulat. 2013;6:1\u201313.","journal-title":"Brain Stimulat"},{"key":"3851_CR89","doi-asserted-by":"publisher","first-page":"689","DOI":"10.1016\/j.biopsych.2020.05.033","volume":"90","author":"RFH Cash","year":"2021","unstructured":"Cash RFH, Weigand A, Zalesky A, Siddiqi SH, Downar J, Fitzgerald PB, et al. Using brain imaging to improve spatial targeting of transcranial magnetic stimulation for depression. Biol Psychiatry. 2021;90:689\u2013700.","journal-title":"Biol Psychiatry"},{"key":"3851_CR90","doi-asserted-by":"publisher","first-page":"1054","DOI":"10.1126\/science.1252900","volume":"345","author":"JX Wang","year":"2014","unstructured":"Wang JX, Rogers LM, Gross EZ, Ryals AJ, Dokucu ME, Brandstatt KL, et al. Targeted enhancement of cortical-hippocampal brain networks and associative memory. Science. 2014;345:1054\u20137.","journal-title":"Science"},{"key":"3851_CR91","doi-asserted-by":"publisher","first-page":"2191","DOI":"10.1001\/jama.2013.281053","volume":"310","author":"World Medical Association.","year":"2013","unstructured":"World Medical Association. World Medical Association Declaration of Helsinki: Ethical principles for medical research involving human subjects. JAMA. 2013;310:2191.","journal-title":"JAMA"}],"container-title":["Translational Psychiatry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-03851-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-03851-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-03851-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,31]],"date-time":"2026-03-31T15:02:55Z","timestamp":1774969375000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41398-026-03851-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,12]]},"references-count":91,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["3851"],"URL":"https:\/\/doi.org\/10.1038\/s41398-026-03851-8","relation":{},"ISSN":["2158-3188"],"issn-type":[{"value":"2158-3188","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,12]]},"assertion":[{"value":"19 June 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 December 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"20 January 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 March 2026","order":4,"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"}},{"value":"The validation study was approved by the Ethics Committee of the Affiliated Hospital of Hangzhou Normal University (No. 2023 (E2) - HS - 089) and was conducted in accordance with the Declaration of Helsinki [\n                      \n                      ]. All participants provided written informed consent prior to participation.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Compliance with ethical standards"}}],"article-number":"175"}}