{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,1]],"date-time":"2026-05-01T05:40:45Z","timestamp":1777614045454,"version":"3.51.4"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1013093","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,9,10]],"date-time":"2025-09-10T00:00:00Z","timestamp":1757462400000}}],"reference-count":109,"publisher":"Public Library of Science (PLoS)","issue":"9","license":[{"start":{"date-parts":[[2025,9,2]],"date-time":"2025-09-02T00:00:00Z","timestamp":1756771200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN\/05221-2020"],"award-info":[{"award-number":["RGPIN\/05221-2020"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100013200","name":"Government of Alberta","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100013200","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100009192","name":"Alberta Innovates","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100009192","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Although the brain\u2019s structural and functional alterations with age are individually well documented, how differences in cognitive abilities emerge from variations in the underlying spatio-temporal patterns of regional brain activity is largely unknown. Patterns of increased synchronization between brain regions are taken as enhanced cognitive integration, while decreased synchronization is indicative of cognitive segregation. The ability to dynamically switch between different levels of integration and segregation across different cognitive systems is believed to be crucial for overall cognitive performance. Building on a recently proposed cognitively informed, synchronization-based framework, we study here age-related variations in dynamical flexibility between segregation and integration, as captured by changes in the variable patterns of partial synchronization or chimera states. Leveraging personalized brain network models based on large-scale, multisite datasets of cross-sectional healthy cohorts, we systematically show how regional brain stimulation produces distinct patterns of synchronization. We find that chimera states play a crucial role in regulating the balance between cognitive integration and segregation as the brain ages, providing new insights into the mechanisms underlying cognitive decline and preservation in aging. Whereas the emergent synchronization behavior of brain regions belonging to the same cognitive system often show the same aging trends, different cognitive systems can demonstrate distinct trends. This supports the idea that aging affects cognitive systems differently and that understanding this variability is essential for a more comprehensive view of neuro-cognitive aging. At the same time, dynamical flexibility increases in the oldest age groups across most cognitive systems. This may reflect compensatory mechanisms to counteract age-related cognitive declines and points towards a phenomenon of dedifferentiation. Yet, the multiplicity of behaviors highlights that whereas dedifferentiation emerges in certain cognitive systems, differentiation can also occur in others. This illustrates that these processes, though seemingly oppositional, can coexist and unfold in parallel across different cognitive systems.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1013093","type":"journal-article","created":{"date-parts":[[2025,9,2]],"date-time":"2025-09-02T18:12:25Z","timestamp":1756836745000},"page":"e1013093","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":3,"title":["Changing cognitive chimera states in human brain networks with age: Variations in cognitive integration and segregation"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4566-0016","authenticated-orcid":true,"given":"Andrew","family":"Patton","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8103-4442","authenticated-orcid":true,"given":"J\u00f6rn","family":"Davidsen","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2025,9,2]]},"reference":[{"key":"pcbi.1013093.ref001","unstructured":"United Nations Department of Economy and Social Affairs. Population Division 2022 : Summary of Results. 2022."},{"issue":"4","key":"pcbi.1013093.ref002","doi-asserted-by":"crossref","first-page":"1439","DOI":"10.3390\/ijerph17041439","article-title":"The impact of population aging and public health support on EU labor markets","volume":"17","author":"M Cristea","year":"2020","journal-title":"Int J Environ Res Public Health."},{"issue":"1","key":"pcbi.1013093.ref003","doi-asserted-by":"crossref","DOI":"10.1038\/s42949-021-00023-z","article-title":"Ageing and population shrinking: implications for sustainability in the urban century","volume":"1","author":"MP Jarzebski","year":"2021","journal-title":"npj Urban Sustain."},{"issue":"4","key":"pcbi.1013093.ref004","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1093\/geront\/37.4.433","article-title":"Successful aging","volume":"37","author":"JW Rowe","year":"1997","journal-title":"Gerontologist."},{"issue":"2","key":"pcbi.1013093.ref005","first-page":"359","article-title":"Definitions of successful ageing: a brief review of a multidimensional concept","volume":"90","author":"A Urtamo","year":"2019","journal-title":"Acta Biomed."},{"issue":"1","key":"pcbi.1013093.ref006","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1080\/0361073X.2017.1398849","article-title":"Successful aging: the role of cognitive gerontology","volume":"44","author":"A Hartley","year":"2018","journal-title":"Exp Aging Res."},{"issue":"3","key":"pcbi.1013093.ref007","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1016\/j.cger.2014.04.001","article-title":"Dementia and cognitive impairment: epidemiology, diagnosis, and treatment","volume":"30","author":"J Hugo","year":"2014","journal-title":"Clin Geriatr Med."},{"issue":"9","key":"pcbi.1013093.ref008","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1016\/S1474-4422(11)70072-2","article-title":"The projected effect of risk factor reduction on Alzheimer\u2019s disease prevalence","volume":"10","author":"DE Barnes","year":"2011","journal-title":"Lancet Neurol."},{"key":"pcbi.1013093.ref009","doi-asserted-by":"crossref","first-page":"31","DOI":"10.3389\/neuro.09.031.2009","article-title":"The human brain in numbers: a linearly scaled-up primate brain","volume":"3","author":"S Herculano-Houzel","year":"2009","journal-title":"Front Hum Neurosci."},{"issue":"1","key":"pcbi.1013093.ref010","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1162\/jocn.1990.2.1.58","article-title":"The bounded brain: toward quantitative neuroanatomy","volume":"2","author":"C Cherniak","year":"1990","journal-title":"J Cogn Neurosci."},{"issue":"5","key":"pcbi.1013093.ref011","doi-asserted-by":"crossref","first-page":"336","DOI":"10.1038\/nrn3214","article-title":"The economy of brain network organization","volume":"13","author":"E Bullmore","year":"2012","journal-title":"Nat Rev Neurosci."},{"issue":"5","key":"pcbi.1013093.ref012","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1177\/1073858416667720","article-title":"Small-World brain networks revisited","volume":"23","author":"DS Bassett","year":"2017","journal-title":"Neuroscientist."},{"key":"pcbi.1013093.ref013","doi-asserted-by":"crossref","first-page":"200","DOI":"10.3389\/fnins.2010.00200","article-title":"Modular and hierarchically modular organization of brain networks","volume":"4","author":"D Meunier","year":"2010","journal-title":"Front Neurosci."},{"issue":"44","key":"pcbi.1013093.ref014","doi-asserted-by":"crossref","first-page":"15775","DOI":"10.1523\/JNEUROSCI.3539-11.2011","article-title":"Rich-club organization of the human connectome","volume":"31","author":"MP van den Heuvel","year":"2011","journal-title":"J Neurosci."},{"issue":"2","key":"pcbi.1013093.ref015","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.conb.2012.11.015","article-title":"Network attributes for segregation and integration in the human brain","volume":"23","author":"O Sporns","year":"2013","journal-title":"Curr Opin Neurobiol."},{"issue":"18","key":"pcbi.1013093.ref016","doi-asserted-by":"crossref","first-page":"6349","DOI":"10.1002\/hbm.26511","article-title":"Integration and segregation manifolds in the brain ensure cognitive flexibility during tasks and rest","volume":"44","author":"K Capouskova","year":"2023","journal-title":"Hum Brain Mapp."},{"issue":"2","key":"pcbi.1013093.ref017","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1016\/j.neuron.2016.09.018","article-title":"The dynamics of functional brain networks: integrated network states during cognitive task performance","volume":"92","author":"JM Shine","year":"2016","journal-title":"Neuron."},{"issue":"48","key":"pcbi.1013093.ref018","doi-asserted-by":"crossref","first-page":"12083","DOI":"10.1523\/JNEUROSCI.2965-15.2016","article-title":"The segregation and integration of distinct brain networks and their relationship to cognition","volume":"36","author":"JR Cohen","year":"2016","journal-title":"J Neurosci."},{"issue":"37","key":"pcbi.1013093.ref019","doi-asserted-by":"crossref","first-page":"11678","DOI":"10.1073\/pnas.1422487112","article-title":"Dynamic reconfiguration of frontal brain networks during executive cognition in humans","volume":"112","author":"U Braun","year":"2015","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"23","key":"pcbi.1013093.ref020","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2022288118","article-title":"Segregation, integration, and balance of large-scale resting brain networks configure different cognitive abilities","volume":"118","author":"R Wang","year":"2021","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"1","key":"pcbi.1013093.ref021","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1146\/annurev-neuro-071013-014030","article-title":"The Brain\u2019s default mode network","volume":"38","author":"ME Raichle","year":"2015","journal-title":"Annu Rev Neurosci."},{"issue":"2","key":"pcbi.1013093.ref022","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1038\/s41593-018-0312-0","article-title":"Human cognition involves the dynamic integration of neural activity and neuromodulatory systems","volume":"22","author":"JM Shine","year":"2019","journal-title":"Nat Neurosci."},{"issue":"4","key":"pcbi.1013093.ref023","doi-asserted-by":"crossref","first-page":"1835","DOI":"10.1109\/JBHI.2021.3119940","article-title":"Flexible brain transitions between hierarchical network segregation and integration associated with cognitive performance during a multisource interference task","volume":"26","author":"R Wang","year":"2022","journal-title":"IEEE J Biomed Health Inform."},{"key":"pcbi.1013093.ref024","doi-asserted-by":"crossref","first-page":"13217","DOI":"10.1038\/ncomms13217","article-title":"Integration and segregation of large-scale brain networks during short-term task automatization","volume":"7","author":"H Mohr","year":"2016","journal-title":"Nat Commun."},{"key":"pcbi.1013093.ref025","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.neuroimage.2016.11.006","article-title":"Multi-scale brain networks","volume":"160","author":"RF Betzel","year":"2017","journal-title":"Neuroimage."},{"issue":"1","key":"pcbi.1013093.ref026","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1038\/s42003-025-07470-9","article-title":"Dynamic switching between brain networks predicts creative ability","volume":"8","author":"Q Chen","year":"2025","journal-title":"Commun Biol."},{"issue":"1","key":"pcbi.1013093.ref027","doi-asserted-by":"crossref","first-page":"5135","DOI":"10.1038\/s41598-017-05425-7","article-title":"Cognitive performance in healthy older adults relates to spontaneous switching between states of functional connectivity during rest","volume":"7","author":"J Cabral","year":"2017","journal-title":"Sci Rep."},{"issue":"11","key":"pcbi.1013093.ref028","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1038\/s41583-018-0068-2","article-title":"Maintenance, reserve and compensation: the cognitive neuroscience of healthy ageing","volume":"19","author":"R Cabeza","year":"2018","journal-title":"Nat Rev Neurosci."},{"issue":"10","key":"pcbi.1013093.ref029","doi-asserted-by":"crossref","first-page":"502","DOI":"10.1016\/j.tics.2013.08.012","article-title":"Efficiency, capacity, compensation, maintenance, plasticity: emerging concepts in cognitive reserve","volume":"17","author":"D Barulli","year":"2013","journal-title":"Trends Cogn Sci."},{"issue":"10","key":"pcbi.1013093.ref030","doi-asserted-by":"crossref","first-page":"2015","DOI":"10.1016\/j.neuropsychologia.2009.03.004","article-title":"Cognitive reserve","volume":"47","author":"Y Stern","year":"2009","journal-title":"Neuropsychologia."},{"issue":"5","key":"pcbi.1013093.ref031","doi-asserted-by":"crossref","first-page":"292","DOI":"10.1016\/j.tics.2012.04.005","article-title":"Memory aging and brain maintenance","volume":"16","author":"L Nyberg","year":"2012","journal-title":"Trends Cogn Sci."},{"key":"pcbi.1013093.ref032","doi-asserted-by":"crossref","unstructured":"Nyberg L, Lindenberger U. Brain maintenance and cognition in old age. In: Poeppel D, Mangun GR, Gazzaniga MS, editors. The cognitive neurosciences. 6th ed. Cambridge, MA: MIT Press; 2020. p. 81\u20139.","DOI":"10.7551\/mitpress\/11442.003.0011"},{"issue":"3","key":"pcbi.1013093.ref033","doi-asserted-by":"crossref","first-page":"1394","DOI":"10.1006\/nimg.2002.1280","article-title":"Aging gracefully: compensatory brain activity in high-performing older adults","volume":"17","author":"R Cabeza","year":"2002","journal-title":"Neuroimage."},{"issue":"5","key":"pcbi.1013093.ref034","doi-asserted-by":"crossref","first-page":"1201","DOI":"10.1093\/cercor\/bhm155","article-title":"Que PASA? The posterior-anterior shift in aging","volume":"18","author":"SW Davis","year":"2008","journal-title":"Cereb Cortex."},{"issue":"3","key":"pcbi.1013093.ref035","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1007\/s11065-014-9270-9","article-title":"How does it STAC up? Revisiting the scaffolding theory of aging and cognition","volume":"24","author":"PA Reuter-Lorenz","year":"2014","journal-title":"Neuropsychol Rev."},{"issue":"9","key":"pcbi.1013093.ref036","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1008144","article-title":"Relations between large-scale brain connectivity and effects of regional stimulation depend on collective dynamical state","volume":"16","author":"L Papadopoulos","year":"2020","journal-title":"PLoS Comput Biol."},{"issue":"4","key":"pcbi.1013093.ref037","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1038\/35067550","article-title":"The brainweb: phase synchronization and large-scale integration","volume":"2","author":"F Varela","year":"2001","journal-title":"Nat Rev Neurosci."},{"issue":"1","key":"pcbi.1013093.ref038","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/j.neuron.2013.12.022","article-title":"The metastable brain","volume":"81","author":"E Tognoli","year":"2014","journal-title":"Neuron."},{"issue":"4","key":"pcbi.1013093.ref039","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1113\/jphysiol.2012.239590","article-title":"Synchronization and desynchronization in epilepsy: controversies and hypotheses","volume":"591","author":"P Jiruska","year":"2013","journal-title":"J Physiol."},{"key":"pcbi.1013093.ref040","unstructured":"Kuramoto Y, Battogtokh D. Coexistence of coherence and incoherence in nonlocally coupled phase oscillators. arXiv preprint 2002."},{"issue":"17","key":"pcbi.1013093.ref041","doi-asserted-by":"crossref","first-page":"174102","DOI":"10.1103\/PhysRevLett.93.174102","article-title":"Chimera states for coupled oscillators","volume":"93","author":"DM Abrams","year":"2004","journal-title":"Phys Rev Lett."},{"issue":"1","key":"pcbi.1013093.ref042","doi-asserted-by":"crossref","first-page":"012205","DOI":"10.1103\/PhysRevE.93.012205","article-title":"Chimera states in bursting neurons","volume":"93","author":"BK Bera","year":"2016","journal-title":"Phys Rev E."},{"key":"pcbi.1013093.ref043","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.plrev.2018.09.003","article-title":"Chimera states in neuronal networks: a review","volume":"28","author":"S Majhi","year":"2019","journal-title":"Phys Life Rev."},{"key":"pcbi.1013093.ref044","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.physrep.2020.10.003","article-title":"Chimeras","volume":"898","author":"F Parastesh","year":"2021","journal-title":"Physics Reports."},{"issue":"12","key":"pcbi.1013093.ref045","doi-asserted-by":"crossref","first-page":"120402","DOI":"10.1063\/5.0249682","article-title":"Introduction to focus issue: chimera states: from theory and experiments to technology and living systems","volume":"34","author":"J Davidsen","year":"2024","journal-title":"Chaos."},{"issue":"1","key":"pcbi.1013093.ref046","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1038\/s42005-024-01728-0","article-title":"An exact mathematical description of computation with transient spatiotemporal dynamics in a complex-valued neural network","volume":"7","author":"RC Budzinski","year":"2024","journal-title":"Commun Phys."},{"issue":"1","key":"pcbi.1013093.ref047","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1038\/s42005-022-00984-2","article-title":"Embedded chimera states in recurrent neural networks","volume":"5","author":"M Masoliver","year":"2022","journal-title":"Commun Phys."},{"key":"pcbi.1013093.ref048","doi-asserted-by":"crossref","DOI":"10.1101\/2023.07.28.550946","volume-title":"The hippocampal theta oscillation may be generated by chimera dynamics","author":"M Masoliver","year":"2023"},{"key":"pcbi.1013093.ref049","doi-asserted-by":"crossref","first-page":"12468","DOI":"10.1038\/ncomms12468","article-title":"Evidence that birds sleep in mid-flight","volume":"7","author":"NC Rattenborg","year":"2016","journal-title":"Nat Commun."},{"key":"pcbi.1013093.ref050","doi-asserted-by":"crossref","first-page":"221","DOI":"10.2147\/NSS.S71970","article-title":"Unihemispheric sleep and asymmetrical sleep: behavioral, neurophysiological, and functional perspectives","volume":"8","author":"GG Mascetti","year":"2016","journal-title":"Nat Sci Sleep."},{"issue":"9","key":"pcbi.1013093.ref051","doi-asserted-by":"crossref","first-page":"1190","DOI":"10.1016\/j.cub.2016.02.063","article-title":"Night watch in one brain hemisphere during sleep associated with the first-night effect in humans","volume":"26","author":"M Tamaki","year":"2016","journal-title":"Curr Biol."},{"issue":"21","key":"pcbi.1013093.ref052","doi-asserted-by":"crossref","first-page":"218403","DOI":"10.1103\/PhysRevLett.132.218403","article-title":"Spatial-temporal analysis of neural desynchronization in sleeplike states reveals critical dynamics","volume":"132","author":"D Curic","year":"2024","journal-title":"Phys Rev Lett."},{"issue":"4","key":"pcbi.1013093.ref053","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.aau8535","article-title":"Cognitive chimera states in human brain networks","volume":"5","author":"K Bansal","year":"2019","journal-title":"Sci Adv."},{"key":"pcbi.1013093.ref054","doi-asserted-by":"crossref","first-page":"425","DOI":"10.3389\/fneur.2019.00425","article-title":"10Kin1day: a bottom-up neuroimaging initiative","volume":"10","author":"MP van den Heuvel","year":"2019","journal-title":"Front Neurol."},{"issue":"3","key":"pcbi.1013093.ref055","doi-asserted-by":"crossref","first-page":"968","DOI":"10.1016\/j.neuroimage.2006.01.021","article-title":"An automated labeling system for subdividing the human cerebral cortex on MRI scans into gyral based regions of interest","volume":"31","author":"RS Desikan","year":"2006","journal-title":"Neuroimage."},{"issue":"2","key":"pcbi.1013093.ref056","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1016\/j.jneumeth.2011.09.031","article-title":"Mapping the human connectome at multiple scales with diffusion spectrum MRI","volume":"203","author":"L Cammoun","year":"2012","journal-title":"J Neurosci Methods."},{"issue":"7028","key":"pcbi.1013093.ref057","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1038\/nature03288","article-title":"Functional cartography of complex metabolic networks","volume":"433","author":"R Guimer\u00e0","year":"2005","journal-title":"Nature."},{"issue":"12","key":"pcbi.1013093.ref058","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1016\/j.tics.2013.09.012","article-title":"Network hubs in the human brain","volume":"17","author":"MP van den Heuvel","year":"2013","journal-title":"Trends Cogn Sci."},{"key":"pcbi.1013093.ref059","doi-asserted-by":"crossref","first-page":"561","DOI":"10.1016\/j.neuroimage.2016.10.044","article-title":"Cortical rich club regions can organize state-dependent functional network formation by engaging in oscillatory behavior","volume":"146","author":"M Senden","year":"2017","journal-title":"Neuroimage."},{"issue":"2","key":"pcbi.1013093.ref060","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1038\/nphys209","article-title":"Detecting rich-club ordering in complex networks","volume":"2","author":"V Colizza","year":"2006","journal-title":"Nature Phys."},{"issue":"14","key":"pcbi.1013093.ref061","doi-asserted-by":"crossref","first-page":"4239","DOI":"10.1002\/hbm.25950","article-title":"Trajectory of rich club properties in structural brain networks","volume":"43","author":"L Riedel","year":"2022","journal-title":"Hum Brain Mapp."},{"key":"pcbi.1013093.ref062","doi-asserted-by":"crossref","DOI":"10.1016\/S1053-8119(09)71822-1","article-title":"Brain connectivity toolbox: a collection of complex network measurements and brain connectivity datasets.","volume":"47","author":"M Rubinov","year":"2009","journal-title":"NeuroImage."},{"issue":"7","key":"pcbi.1013093.ref063","doi-asserted-by":"crossref","first-page":"430","DOI":"10.1038\/nrn3963","article-title":"Rethinking segregation and integration: contributions of whole-brain modelling","volume":"16","author":"G Deco","year":"2015","journal-title":"Nat Rev Neurosci."},{"issue":"1","key":"pcbi.1013093.ref064","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0006-3495(72)86068-5","article-title":"Excitatory and inhibitory interactions in localized populations of model neurons","volume":"12","author":"HR Wilson","year":"1972","journal-title":"Biophys J."},{"key":"pcbi.1013093.ref065","volume-title":"Neuroscience","author":"D Purves","year":"2001"},{"issue":"12","key":"pcbi.1013093.ref066","doi-asserted-by":"crossref","first-page":"2952","DOI":"10.3390\/ijms20122952","article-title":"Diversity and function of somatostatin-expressing interneurons in the cerebral cortex","volume":"20","author":"T Riedemann","year":"2019","journal-title":"Int J Mol Sci."},{"issue":"9","key":"pcbi.1013093.ref067","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1005076","article-title":"Stimulation-based control of dynamic brain networks","volume":"12","author":"SF Muldoon","year":"2016","journal-title":"PLoS Comput Biol."},{"issue":"4","key":"pcbi.1013093.ref068","doi-asserted-by":"crossref","first-page":"665","DOI":"10.1016\/j.neuron.2011.09.006","article-title":"Functional network organization of the human brain","volume":"72","author":"JD Power","year":"2011","journal-title":"Neuron."},{"issue":"3","key":"pcbi.1013093.ref069","doi-asserted-by":"crossref","first-page":"1125","DOI":"10.1152\/jn.00338.2011","article-title":"The organization of the human cerebral cortex estimated by intrinsic functional connectivity","volume":"106","author":"BTT Yeo","year":"2011","journal-title":"J Neurophysiol."},{"key":"pcbi.1013093.ref070","volume-title":"Changing cognitive chimera states in human brain networks with age: variations in cognitive integration and segregation","author":"D Patton","year":"2025"},{"issue":"10","key":"pcbi.1013093.ref071","doi-asserted-by":"crossref","DOI":"10.1088\/1742-5468\/2008\/10\/P10008","article-title":"Fast unfolding of communities in large networks","volume":"2008","author":"VD Blondel","year":"2008","journal-title":"J Stat Mech."},{"issue":"5980","key":"pcbi.1013093.ref072","doi-asserted-by":"crossref","first-page":"876","DOI":"10.1126\/science.1184819","article-title":"Community structure in time-dependent, multiscale, and multiplex networks","volume":"328","author":"PJ Mucha","year":"2010","journal-title":"Science."},{"issue":"3","key":"pcbi.1013093.ref073","doi-asserted-by":"crossref","first-page":"297","DOI":"10.2307\/1932409","article-title":"Measures of the amount of ecologic association between species","volume":"26","author":"LR Dice","year":"1945","journal-title":"Ecology."},{"key":"pcbi.1013093.ref074","doi-asserted-by":"crossref","first-page":"6","DOI":"10.1038\/s44172-023-00140-w","article-title":"Learning multi-site harmonization of magnetic resonance images without traveling human phantoms","volume":"3","author":"S Liu","year":"2024","journal-title":"Commun Eng."},{"issue":"2","key":"pcbi.1013093.ref075","doi-asserted-by":"crossref","first-page":"172","DOI":"10.2967\/jnumed.121.262464","article-title":"A guide to ComBat harmonization of imaging biomarkers in multicenter studies","volume":"63","author":"F Orlhac","year":"2022","journal-title":"J Nucl Med."},{"issue":"9","key":"pcbi.1013093.ref076","doi-asserted-by":"crossref","first-page":"1970","DOI":"10.1111\/jgs.16022","article-title":"Investigating and remediating selection bias in geriatrics research: the selection bias toolkit","volume":"67","author":"HR Banack","year":"2019","journal-title":"J Am Geriatr Soc."},{"key":"pcbi.1013093.ref077","unstructured":"Mowinckel A, Pineiro D. Ggseg package for R. [cited 2019 December]. https:\/\/github.com\/LCBC-UiO\/ggseg"},{"key":"pcbi.1013093.ref078","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1016\/j.neuroimage.2014.07.067","article-title":"Changes in structural and functional connectivity among resting-state networks across the human lifespan","author":"RF Betzel","year":"2014","journal-title":"Neuroimage."},{"key":"pcbi.1013093.ref079","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.neuroimage.2017.01.077","article-title":"Effects of aging on functional and structural brain connectivity","volume":"160","author":"JS Damoiseaux","year":"2017","journal-title":"Neuroimage."},{"issue":"1","key":"pcbi.1013093.ref080","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1002\/jnr.24669","article-title":"Functional brain connectivity changes across the human life span: from fetal development to old age","volume":"99","author":"M Edde","year":"2021","journal-title":"J Neurosci Res."},{"key":"pcbi.1013093.ref081","doi-asserted-by":"crossref","first-page":"116756","DOI":"10.1016\/j.neuroimage.2020.116756","article-title":"Functional network reorganization in older adults: Graph-theoretical analyses of age, cognition and sex","volume":"214","author":"J Stumme","year":"2020","journal-title":"Neuroimage."},{"key":"pcbi.1013093.ref082","doi-asserted-by":"crossref","first-page":"116974","DOI":"10.1016\/j.neuroimage.2020.116974","article-title":"The modular organization of brain cortical connectivity across the human lifespan","volume":"218","author":"MG Puxeddu","year":"2020","journal-title":"Neuroimage."},{"issue":"5","key":"pcbi.1013093.ref083","doi-asserted-by":"crossref","first-page":"1354","DOI":"10.1002\/jnr.24795","article-title":"Reorganization of brain structural networks in aging: a longitudinal study","volume":"99","author":"A Coelho","year":"2021","journal-title":"J Neurosci Res."},{"issue":"46","key":"pcbi.1013093.ref084","doi-asserted-by":"crossref","first-page":"15254","DOI":"10.1523\/JNEUROSCI.2135-15.2015","article-title":"Default mode dynamics for global functional integration","volume":"35","author":"D Vatansever","year":"2015","journal-title":"J Neurosci."},{"issue":"1","key":"pcbi.1013093.ref085","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1037\/0882-7974.17.1.85","article-title":"Hemispheric asymmetry reduction in older adults: the HAROLD model","volume":"17","author":"R Cabeza","year":"2002","journal-title":"Psychol Aging."},{"issue":"33","key":"pcbi.1013093.ref086","doi-asserted-by":"crossref","first-page":"7303","DOI":"10.1523\/JNEUROSCI.1701-17.2018","article-title":"Increased prefrontal activity with aging reflects nonspecific neural responses rather than compensation","volume":"38","author":"AM Morcom","year":"2018","journal-title":"J Neurosci."},{"issue":"3","key":"pcbi.1013093.ref087","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1007\/s00221-012-3319-x","article-title":"Reassessing the HAROLD model: is the hemispheric asymmetry reduction in older adults a special case of compensatory-related utilisation of neural circuits?","volume":"224","author":"M Berlingeri","year":"2013","journal-title":"Exp Brain Res."},{"issue":"45","key":"pcbi.1013093.ref088","doi-asserted-by":"crossref","first-page":"9361","DOI":"10.1523\/JNEUROSCI.1111-21.2021","article-title":"Does hemispheric asymmetry reduction in older adults in motor cortex reflect compensation?","volume":"41","author":"E Knights","year":"2021","journal-title":"J Neurosci."},{"issue":"2","key":"pcbi.1013093.ref089","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pbio.3002494","article-title":"Older adults preserve audiovisual integration through enhanced cortical activations, not by recruiting new regions","volume":"22","author":"SA Jones","year":"2024","journal-title":"PLoS Biol."},{"issue":"14","key":"pcbi.1013093.ref090","doi-asserted-by":"crossref","first-page":"2527","DOI":"10.1523\/JNEUROSCI.1331-22.2023","article-title":"Widespread fMRI BOLD signal overactivations during cognitive control in older adults are not matched by corresponding increases in fPET glucose metabolism","volume":"43","author":"L Stiernman","year":"2023","journal-title":"J Neurosci."},{"issue":"10","key":"pcbi.1013093.ref091","doi-asserted-by":"crossref","first-page":"3777","DOI":"10.1002\/hbm.22877","article-title":"Age-related changes in the topological organization of the white matter structural connectome across the human lifespan","volume":"36","author":"T Zhao","year":"2015","journal-title":"Hum Brain Mapp."},{"issue":"50","key":"pcbi.1013093.ref092","doi-asserted-by":"crossref","first-page":"15684","DOI":"10.1523\/JNEUROSCI.2308-09.2009","article-title":"Age- and gender-related differences in the cortical anatomical network","volume":"29","author":"G Gong","year":"2009","journal-title":"J Neurosci."},{"issue":"2","key":"pcbi.1013093.ref093","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1002\/gps.2087","article-title":"Aging of cerebral white matter: a review of MRI findings","volume":"24","author":"FM Gunning-Dixon","year":"2009","journal-title":"Int J Geriatr Psychiatry."},{"issue":"8","key":"pcbi.1013093.ref094","doi-asserted-by":"crossref","first-page":"4651","DOI":"10.1093\/cercor\/bhaa066","article-title":"Age-related decline in the topological efficiency of the brain structural connectome and cognitive aging","volume":"30","author":"X Li","year":"2020","journal-title":"Cereb Cortex."},{"issue":"3","key":"pcbi.1013093.ref095","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.1093\/cercor\/bhz193","article-title":"Contributions of white matter connectivity and BOLD modulation to cognitive aging: a lifespan structure-function association study","volume":"30","author":"CE Webb","year":"2020","journal-title":"Cereb Cortex."},{"key":"pcbi.1013093.ref096","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.neuroimage.2015.04.009","article-title":"The organisation of the elderly connectome","volume":"114","author":"A Perry","year":"2015","journal-title":"Neuroimage."},{"key":"pcbi.1013093.ref097","doi-asserted-by":"crossref","first-page":"234","DOI":"10.3389\/fnagi.2019.00234","article-title":"The effect of aging on resting state connectivity of predefined networks in the brain","volume":"11","author":"E Varangis","year":"2019","journal-title":"Front Aging Neurosci."},{"issue":"1","key":"pcbi.1013093.ref098","doi-asserted-by":"crossref","DOI":"10.1111\/psyp.14159","article-title":"The older adult brain is less modular, more integrated, and less efficient at rest: a systematic review of large-scale resting-state functional brain networks in aging","volume":"60","author":"HA Deery","year":"2023","journal-title":"Psychophysiology."},{"issue":"7","key":"pcbi.1013093.ref099","doi-asserted-by":"crossref","first-page":"547","DOI":"10.1016\/j.tics.2019.04.012","article-title":"Neural dedifferentiation in the aging brain","volume":"23","author":"JD Koen","year":"2019","journal-title":"Trends Cogn Sci."},{"issue":"46","key":"pcbi.1013093.ref100","article-title":"Decreased segregation of brain systems across the healthy adult lifespan","volume":"111","author":"MY Chan","year":"2014","journal-title":"Proc Natl Acad Sci U S A."},{"issue":"3","key":"pcbi.1013093.ref101","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1093\/geront\/gnu057","article-title":"Training older adults to use tablet computers: does it enhance cognitive function?","volume":"56","author":"MY Chan","year":"2016","journal-title":"Gerontologist."},{"issue":"18","key":"pcbi.1013093.ref102","doi-asserted-by":"crossref","first-page":"5543","DOI":"10.1002\/hbm.26030","article-title":"Interrelating differences in structural and functional connectivity in the older adult\u2019s brain","volume":"43","author":"J Stumme","year":"2022","journal-title":"Hum Brain Mapp."},{"issue":"2","key":"pcbi.1013093.ref103","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1037\/a0039260","article-title":"Cognitive dedifferentiation with increasing age and proximity of death: within-person evidence from the seattle longitudinal study","volume":"30","author":"G H\u00fcl\u00fcr","year":"2015","journal-title":"Psychol Aging."},{"issue":"1","key":"pcbi.1013093.ref104","first-page":"30","article-title":"Functional dedifferentiation and altered connectivity in older adults: neural accounts of cognitive aging","volume":"2","author":"JOS Goh","year":"2011","journal-title":"Aging Dis."},{"issue":"27","key":"pcbi.1013093.ref105","doi-asserted-by":"crossref","first-page":"9253","DOI":"10.1523\/JNEUROSCI.0853-10.2010","article-title":"Neural specificity predicts fluid processing ability in older adults","volume":"30","author":"J Park","year":"2010","journal-title":"J Neurosci."},{"key":"pcbi.1013093.ref106","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/j.neubiorev.2018.09.005","article-title":"Adolescence as a neurobiological critical period for the development of higher-order cognition","volume":"94","author":"B Larsen","year":"2018","journal-title":"Neurosci Biobehav Rev."},{"issue":"1","key":"pcbi.1013093.ref107","doi-asserted-by":"crossref","first-page":"3511","DOI":"10.1038\/s41467-024-47748-w","article-title":"Functional connectivity development along the sensorimotor-association axis enhances the cortical hierarchy","volume":"15","author":"AC Luo","year":"2024","journal-title":"Nat Commun."},{"issue":"4","key":"pcbi.1013093.ref108","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1016\/j.cger.2013.07.002","article-title":"Normal cognitive aging","volume":"29","author":"CN Harada","year":"2013","journal-title":"Clin Geriatr Med."},{"issue":"3","key":"pcbi.1013093.ref109","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1162\/netn_a_00377","article-title":"Reorganization of structural connectivity in the brain supports preservation of cognitive ability in healthy aging","volume":"8","author":"J Neudorf","year":"2024","journal-title":"Netw Neurosci."}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1013093","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,9,10]],"date-time":"2025-09-10T00:00:00Z","timestamp":1757462400000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1013093","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,9,10]],"date-time":"2025-09-10T17:58:38Z","timestamp":1757527118000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1013093"}},"subtitle":[],"editor":[{"given":"Elishai","family":"Ezra Tsur","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2025,9,2]]},"references-count":109,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2025,9,2]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1013093","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.04.28.650949","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,9,2]]}}}