{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T12:31:18Z","timestamp":1789475478185,"version":"build-2803163510"},"reference-count":56,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2025,8,25]],"date-time":"2025-08-25T00:00:00Z","timestamp":1756080000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Southeast University, China","award":["3218002501A2"],"award-info":[{"award-number":["3218002501A2"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Life"],"abstract":"<jats:p>Long-term physical training is known to induce brain plasticity, yet how these neural adaptations evolve across different stages of training remains underexplored. This two-year longitudinal study investigated the stage-dependent effects of endurance running on brain structure and resting-state function in healthy college students. Thirty participants were recruited into three groups based on their endurance training level: high-level runners, moderate-level runners, and sedentary controls. All participants underwent baseline and two-year follow-up MRI scans, including T1-weighted structural imaging and resting-state fMRI. The results revealed that the high-level runners exhibited a significant increase in degree centrality (DC) in the left dorsolateral prefrontal cortex (DLPFC). In the moderate-level group, more widespread changes were observed, including increased gray matter volume (GMV) in bilateral prefrontal cortices, medial frontal regions, the right insula, the right putamen, and the right temporo-parieto-occipital junction, along with decreased GMV in the posterior cerebellum. Additionally, DC decreased in the left thalamus and increased in the right temporal lobe and bilateral DLPFC; the fractional amplitude of low-frequency fluctuations (fALFF) in the right precentral gyrus was also elevated. These brain regions are involved in executive control, sensorimotor integration, and motor coordination, which may suggest potential functional implications for cognitive and motor performance; however, such interpretations should be viewed cautiously given the modest sample size and study duration. No significant changes were found in the control group. These findings demonstrate that long-term endurance training induces distinct patterns of brain plasticity at different training stages, with more prominent and widespread changes occurring during earlier phases of training.<\/jats:p>","DOI":"10.3390\/life15091342","type":"journal-article","created":{"date-parts":[[2025,8,26]],"date-time":"2025-08-26T06:25:57Z","timestamp":1756189557000},"page":"1342","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Stage-Dependent Brain Plasticity Induced by Long-Term Endurance Training: A Longitudinal Neuroimaging Study"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4868-3720","authenticated-orcid":false,"given":"Keying","family":"Zhang","sequence":"first","affiliation":[{"name":"Department of Physical Education, Southeast University, Nanjing 210096, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qing","family":"Yan","sequence":"additional","affiliation":[{"name":"Institute of Artificial Intelligence in Sports, Capital University of Physical Education and Sports, Beijing 100191, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ling","family":"Jiang","sequence":"additional","affiliation":[{"name":"Department of Physical Education, Southeast University, Nanjing 210096, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2133-2445","authenticated-orcid":false,"given":"Dongxue","family":"Liang","sequence":"additional","affiliation":[{"name":"Institute of Artificial Intelligence in Sports, Capital University of Physical Education and Sports, Beijing 100191, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3762-7651","authenticated-orcid":false,"given":"Chunmei","family":"Cao","sequence":"additional","affiliation":[{"name":"Division of Sports Science and Physical Education Tsinghua University, Tsinghua University, Beijing 100084, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7874-9092","authenticated-orcid":false,"given":"Dong","family":"Zhang","sequence":"additional","affiliation":[{"name":"Institute of Artificial Intelligence in Sports, Capital University of Physical Education and Sports, Beijing 100191, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,8,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.dcn.2013.11.003","article-title":"White matter connectivity and aerobic fitness in male adolescents","volume":"7","author":"Herting","year":"2014","journal-title":"Dev. Cogn. Neurosci."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Raichlen, D.A., Bharadwaj, P.K., Fitzhugh, M.C., Haws, K.A., Torre, G.-A., Trouard, T.P., and Alexander, G.E. (2016). Differences in resting state functional connectivity between young adult endurance athletes and healthy controls. Front. Hum. Neurosci., 10.","DOI":"10.3389\/fnhum.2016.00610"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Zhang, K., Cao, C., Wang, Y., and Zhang, D. (2024). Brain structure and function differences across varying levels of endurance training: A cross-sectional study. Front. Hum. Neurosci., 18.","DOI":"10.3389\/fnhum.2024.1503094"},{"key":"ref_4","unstructured":"Hedeker, D., and Gibbons, R.D. (2006). Longitudinal Data Analysis, John Wiley & Sons."},{"key":"ref_5","unstructured":"Ridgway, G., Ashburner, J., and Penny, W. (2013). Modelling Longitudinal Structural Change from Serial MRI, British Neuroscience Association (BNA) Festival of Neuroscience."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"898","DOI":"10.1080\/00207454.2024.2341910","article-title":"Effects on brain structural and functional in deaf children after aerobic exercise training: A pilot cluster randomized controlled study","volume":"135","author":"Qu","year":"2025","journal-title":"Int. J. Neurosci."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1093\/gerona\/61.11.1166","article-title":"Aerobic exercise training increases brain volume in aging humans","volume":"61","author":"Colcombe","year":"2006","journal-title":"J. Gerontol. A Biol. Sci. Med. Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"91","DOI":"10.3233\/JAD-201429","article-title":"Neurocognitive plasticity is associated with cardiorespiratory fitness following physical exercise in older adults with amnestic mild cognitive impairment","volume":"81","author":"Eisenstein","year":"2021","journal-title":"J. Alzheimers Dis."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1177\/1073858418771538","article-title":"Exercise-Induced Neuroplasticity: A Mechanistic Model and Prospects for Promoting Plasticity","volume":"25","author":"Harasym","year":"2019","journal-title":"Neuroscientist"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/S0166-4115(08)10015-2","article-title":"Neuroanatomical correlates of motor skill learning: Inferences from neuroimaging to behavior","volume":"139","author":"Lindquist","year":"2008","journal-title":"Adv. Psychol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1051\/sm\/2021012","article-title":"A different look at featured motor learning models: Comparison exam of Gallahue\u2019s, Fitts and Posner\u2019s and Ann Gentile\u2019s motor learning models","volume":"112","author":"Salehi","year":"2021","journal-title":"Mov. Sport Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/j.neuroimage.2015.01.039","article-title":"Decision-making conflict and the neural efficiency hypothesis of intelligence: A functional near-infrared spectroscopy investigation","volume":"109","author":"Rodrigo","year":"2015","journal-title":"Neuroimage"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/j.neulet.2006.04.006","article-title":"Using graph theoretical analysis of multi channel EEG to evaluate the neural efficiency hypothesis","volume":"402","author":"Micheloyannis","year":"2006","journal-title":"Neurosci. Lett."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Hou, H.-Y., Jia, X.-Z., Wang, P., Zhang, J.-X., Huang, S., and Li, H.-J. (2019). Intrinsic resting-state activity in older adults with video game experience. Front. Aging Neurosci., 11.","DOI":"10.3389\/fnagi.2019.00119"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1432","DOI":"10.1016\/j.neuroimage.2009.09.037","article-title":"The oscillating brain: Complex and reliable","volume":"49","author":"Zuo","year":"2010","journal-title":"Neuroimage"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1404","DOI":"10.1016\/j.neuroimage.2011.08.044","article-title":"Test-retest reliability of resting-state connectivity network characteristics using fMRI and graph theoretical measures","volume":"59","author":"Braun","year":"2012","journal-title":"Neuroimage"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1162\/089892999563698","article-title":"Set-and code-specific activation in the frontal cortex: An fMRI study of encoding and retrieval of faces and words","volume":"11","author":"McDermott","year":"1999","journal-title":"J. Cogn. Neurosci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1073\/pnas.97.1.506","article-title":"Prefrontal cortex and episodic memory retrieval mode","volume":"97","author":"Lepage","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1006\/nimg.2002.1063","article-title":"Similarities and differences in the neural correlates of episodic memory retrieval and working memory","volume":"16","author":"Cabeza","year":"2002","journal-title":"Neuroimage"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1568","DOI":"10.1126\/science.1199892","article-title":"Selective attention from voluntary control of neurons in prefrontal cortex","volume":"332","author":"Schafer","year":"2011","journal-title":"Science"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"J\u00e4ncke, L., Koeneke, S., Hoppe, A., Rominger, C., and H\u00e4nggi, J. (2009). The architecture of the golfer\u2019s brain. PLoS ONE, 4.","DOI":"10.1371\/journal.pone.0004785"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1927","DOI":"10.1016\/j.neurobiolaging.2008.10.007","article-title":"The effect of midlife physical activity on structural brain changes in the elderly","volume":"31","author":"Rovio","year":"2010","journal-title":"Neurobiol. Aging"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Wood, K.N., Nikolov, R., and Shoemaker, J.K. (2016). Impact of long-term endurance training vs. guideline-based physical activity on brain structure in healthy aging. Front. Aging Neurosci., 8.","DOI":"10.3389\/fnagi.2016.00155"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Ji, L., Zhang, H., Potter, G.G., Zang, Y.-F., Steffens, D.C., Guo, H., and Wang, L. (2017). Multiple neuroimaging measures for examining exercise-induced neuroplasticity in older adults: A quasi-experimental study. Front. Aging Neurosci., 9.","DOI":"10.3389\/fnagi.2017.00102"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1007\/s11065-006-9002-x","article-title":"Executive function and the frontal lobes: A meta-analytic review","volume":"16","author":"Alvarez","year":"2006","journal-title":"Neuropsychol. Rev."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2530","DOI":"10.1002\/hbm.20688","article-title":"At the heart of the ventral attention system: The right anterior insula","volume":"30","author":"Eckert","year":"2009","journal-title":"Hum. Brain Mapp."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"2349","DOI":"10.1523\/JNEUROSCI.5587-06.2007","article-title":"Dissociable intrinsic connectivity networks for salience processing and executive control","volume":"27","author":"Seeley","year":"2007","journal-title":"J. Neurosci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/j.acn.2007.08.010","article-title":"Assessment of executive functions: Review of instruments and identification of critical issues","volume":"23","author":"Chan","year":"2008","journal-title":"Arch. Clin. Neuropsychol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"12444","DOI":"10.1523\/JNEUROSCI.1996-11.2011","article-title":"Training-induced neural plasticity in golf novices","volume":"31","author":"Bezzola","year":"2011","journal-title":"J. Neurosci."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1016\/j.neuroimage.2007.01.003","article-title":"The mind of expert motor performance is cool and focused","volume":"35","author":"Milton","year":"2007","journal-title":"Neuroimage"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1397","DOI":"10.1016\/j.pnsc.2008.12.009","article-title":"Brain structure in diving players on MR imaging studied with voxel-based morphometry","volume":"19","author":"Wei","year":"2009","journal-title":"Prog. Nat. Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1002\/hbm.20928","article-title":"Structural neuroplasticity in the sensorimotor network of professional female ballet dancers","volume":"31","author":"Koeneke","year":"2010","journal-title":"Hum. Brain Mapp."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1177\/1073858412446202","article-title":"Emerging roles of the brain\u2019s default network","volume":"19","author":"Mantini","year":"2013","journal-title":"Neuroscientist"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1196\/annals.1440.011","article-title":"The brain\u2019s default network: Anatomy, function, and relevance to disease","volume":"1124","author":"Buckner","year":"2008","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"9673","DOI":"10.1073\/pnas.0504136102","article-title":"The human brain is intrinsically organized into dynamic, anticorrelated functional networks","volume":"102","author":"Fox","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1073\/pnas.98.2.676","article-title":"A default mode of brain function","volume":"98","author":"Raichle","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1515","DOI":"10.1002\/hbm.22008","article-title":"Neural substrates of the self-memory system: New insights from a meta-analysis","volume":"34","author":"Martinelli","year":"2013","journal-title":"Hum. Brain Mapp."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1947","DOI":"10.1111\/j.1460-9568.2009.06973.x","article-title":"Distinct brain networks in recognition memory share a defined region in the precuneus","volume":"30","author":"Werner","year":"2009","journal-title":"Eur. J. Neurosci."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1016\/j.neuroimage.2012.10.007","article-title":"Brain anatomical networks in world class gymnasts: A DTI tractography study","volume":"65","author":"Wang","year":"2013","journal-title":"Neuroimage"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"11656","DOI":"10.1073\/pnas.191374198","article-title":"Functional neuroanatomy of biological motion perception in humans","volume":"98","author":"Vaina","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"342","DOI":"10.3346\/jkms.2006.21.2.342","article-title":"Evaluation of morphological plasticity in the cerebella of basketball players with MRI","volume":"21","author":"Park","year":"2006","journal-title":"J. Korean Med. Sci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1188\/15.ONF.552-554","article-title":"Preferred reporting items for systematic reviews and meta-analyses","volume":"42","author":"Vrabel","year":"2015","journal-title":"Oncol. Nurs. Forum"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/j.euroneuro.2010.03.008","article-title":"Exploring the brain network: A review on resting-state fMRI functional connectivity","volume":"20","author":"Pol","year":"2010","journal-title":"Eur. Neuropsychopharmacol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1109\/TBDATA.2017.2734883","article-title":"Resting-state fMRI functional connectivity: Big data preprocessing pipelines and topological data analysis","volume":"3","author":"Phinyomark","year":"2017","journal-title":"IEEE Trans. Big Data"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1038\/nature10844","article-title":"Repetitive motor learning induces coordinated formation of clustered dendritic spines in vivo","volume":"483","author":"Fu","year":"2012","journal-title":"Nature"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1348","DOI":"10.1038\/nn.3470","article-title":"Multi-task connectivity reveals flexible hubs for adaptive task control","volume":"16","author":"Cole","year":"2013","journal-title":"Nat. Neurosci."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1257","DOI":"10.1016\/j.neuroimage.2006.01.033","article-title":"Building a motor simulation de novo: Observation of dance by dancers","volume":"31","author":"Cross","year":"2006","journal-title":"Neuroimage"},{"key":"ref_48","first-page":"129","article-title":"Effects of aerobic exercise training on brain structure and psychological well-being in young adults","volume":"49","author":"Gondoh","year":"2009","journal-title":"J. Sports Med. Phys. Fit."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1093\/cercor\/bhi005","article-title":"Human functional neuroimaging of brain changes associated with practice","volume":"15","author":"Kelly","year":"2005","journal-title":"Cereb. Cortex"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.neulet.2005.04.054","article-title":"The effects of regular aerobic exercise in adolescent period on hippocampal neuron density, apoptosis and spatial memory","volume":"383","author":"Uysal","year":"2005","journal-title":"Neurosci. Lett."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"230","DOI":"10.1016\/j.neuroimage.2017.11.007","article-title":"Effect of aerobic exercise on hippocampal volume in humans: A systematic review and meta-analysis","volume":"166","author":"Firth","year":"2018","journal-title":"Neuroimage"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"464","DOI":"10.1016\/j.tins.2007.06.011","article-title":"Exercise builds brain health: Key roles of growth factor cascades and inflammation","volume":"30","author":"Cotman","year":"2007","journal-title":"Trends Neurosci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1007\/s00702-008-0071-2","article-title":"Downhill training upregulates mice hippocampal and striatal brain-derived neurotrophic factor levels","volume":"115","author":"Aguiar","year":"2008","journal-title":"J. Neural Transm."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1016\/j.neuroimage.2017.08.011","article-title":"A whole brain volumetric approach in overweight\/obese children: Examining the association with different physical fitness components and academic performance. The ActiveBrains project","volume":"159","author":"Migueles","year":"2017","journal-title":"Neuroimage"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"10988","DOI":"10.1073\/pnas.1815197116","article-title":"Leptin in hippocampus mediates benefits of mild exercise by an antioxidant on neurogenesis and memory","volume":"116","author":"Yook","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"So, J.H., Huang, C., Ge, M., Cai, G., Zhang, L., Lu, Y., and Mu, Y. (2017). Intense Exercise Promotes Adult Hippocampal Neurogenesis But Not Spatial Discrimination. Front. Cell. Neurosci., 11.","DOI":"10.3389\/fncel.2017.00013"}],"container-title":["Life"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-1729\/15\/9\/1342\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:32:20Z","timestamp":1760034740000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-1729\/15\/9\/1342"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,8,25]]},"references-count":56,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2025,9]]}},"alternative-id":["life15091342"],"URL":"https:\/\/doi.org\/10.3390\/life15091342","relation":{},"ISSN":["2075-1729"],"issn-type":[{"value":"2075-1729","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,8,25]]}}}