Three parsimonious spatiotemporal patterns in cerebellum reveal individual traits in function and behavior | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Three parsimonious spatiotemporal patterns in cerebellum reveal individual traits in function and behavior Guoyuan Yang, Shuo Lv, Jinlong Li, Ruoqi Yang, Xinyu Wu, Zhiming Wang, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7559401/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted You are reading this latest preprint version Abstract The cerebellum plays a critical role in large-scale brain functional connectivity structure and synchronizing global waves propagation. Despite extensive studies on cortical spatiotemporal architecture, particularly in zero-lag and time-lag synchrony, leaving the parsimonious cerebellar representational structure largely overlooked. Using complex principal component analysis (CPCA) on resting-state fMRI data from the Human Connectome Project, we delineate three dominant low-dimensional spatiotemporal patterns within the cerebellum, manifesting as unimodal-to-transmodal progression, functional antagonism, and hemispheric asymmetry. These patterns correlate with various empirical functional connectivity topographies, including functional connectome network structure, quasiperiodic pattern, functional connectivity gradients and brain network dynamics hierarchy. Importantly, functional connectivity reconstructed from these patterns preserves the intrinsic structural modularity, proving the robustness of the sparse cerebellar dynamics. Moreover, personalized CPCA-derived features enable accurate sex classification and prediction of cognitive, emotional, alertness and personality traits. Our findings position the cerebellum as an active, trait-sensitive integrator of brain-wide spatiotemporal organization, suggesting propagation-based motifs constitute a foundational mode of individual brain intrinsic function. Biological sciences/Neuroscience/Computational neuroscience/Dynamical systems Biological sciences/Neuroscience/Cognitive neuroscience Full Text Additional Declarations There is NO Competing Interest. Supplementary Files supplementary.docx SI Cite Share Download PDF Status: Under Review Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7559401","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":530756211,"identity":"1d89eb0e-b9ee-444f-94ba-a01cb33db47d","order_by":0,"name":"Guoyuan 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