{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T06:50:05Z","timestamp":1766040605474,"version":"3.48.0"},"reference-count":49,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T00:00:00Z","timestamp":1766016000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Physiol."],"abstract":"<jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>Cross-frequency coupling (CFC), particularly phase\u2013amplitude coupling (PAC), reflects hierarchical interactions between neural oscillations and plays a critical role in sensorimotor integration. However, its functional relevance during balance control under sensory perturbations remains insufficiently understood.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Objective<\/jats:title>\n                    <jats:p>This study aimed to investigate PAC characteristics during postural control tasks of varying difficulty in elite freestyle aerial skiers versus non-athlete controls.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>EEG signals and center of pressure (COP) data were recorded from participants performing six standing balance tasks on stable and unstable surfaces. Postural control was assessed using center of pressure data, which represent the point location of the body\u2019s vertical ground reaction force vector and are commonly used to quantify sway and balance performance during stance. Mean Vector Length Modulation Index (MVLmi) and PAC analyses were applied to assess oscillatory interactions.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Surface instability significantly modulated PAC strength across frequency bands (\n                      <jats:italic>P<\/jats:italic>\n                      &amp;lt; 0.05). Athletes exhibited task-specific enhancements in alpha-gamma and delta-gamma coupling during single-leg and double-leg stance. These coupling patterns were more spatially localized and showed trends consistent with superior postural control. In contrast, non-athletes showed widespread PAC increases under perturbation, but with less effective balance performance. Hemispheric asymmetries were observed during single-leg stance: athletes demonstrated contralateral dominance during right-leg tasks and ipsilateral coupling shifts during unstable left-leg stance, indicating dynamic lateralized control shaped by training. Across conditions, athletes showed higher PAC strength and lower sample entropy, reflecting more efficient and adaptable cortical strategies for postural regulation.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>PAC strength is closely linked to postural performance and varies with task complexity and surface condition. These findings highlight the role of training-induced neuroplasticity in modulating cortical dynamics for balance control, offering new insights for targeted neuromodulatory interventions and neurofeedback-based training strategies.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fphys.2025.1700523","type":"journal-article","created":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T06:45:05Z","timestamp":1766040305000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Task-dependent cross-frequency neural coupling during postural perturbation: insights from EEG-based assessment in elite freestyle aerial skiers"],"prefix":"10.3389","volume":"16","author":[{"given":"Yuqi","family":"Cheng","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ao","family":"Fu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Youcai","family":"Guo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jie","family":"Gao","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yongxia","family":"Chen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Qianrong","family":"Qi","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Feng","family":"Guo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xin","family":"Wang","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2025,12,18]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"579","DOI":"10.3390\/e20080579","article-title":"Sample entropy of human gait center of pressure displacement: a systematic methodological analysis","volume":"20","author":"Ahmadi","year":"2018","journal-title":"Entropy"},{"key":"B2","doi-asserted-by":"publisher","first-page":"527","DOI":"10.1007\/bf01797193","article-title":"\u00dcber das elektroenkephalogramm des menschen","volume":"87","author":"Berger","year":"1929","journal-title":"Arch. f\u00fcr Psychiatr. Nervenkrankh."},{"key":"B3","doi-asserted-by":"publisher","first-page":"1193","DOI":"10.1007\/s00221-021-06035-6","article-title":"Stance leg and surface stability modulate cortical activity during human single leg stance","volume":"239","author":"B\u00fcchel","year":"2021","journal-title":"Exp. Brain Res."},{"key":"B4","doi-asserted-by":"publisher","first-page":"1926","DOI":"10.1126\/science.1099745","article-title":"Neuronal oscillations in cortical networks","volume":"304","author":"Buzs\u00e1ki","year":"2004","journal-title":"Science"},{"key":"B5","doi-asserted-by":"publisher","first-page":"506","DOI":"10.1016\/j.tics.2010.09.001","article-title":"The functional role of cross-frequency coupling","volume":"14","author":"Canolty","year":"2010","journal-title":"Trends Cognitive Sciences"},{"key":"B6","doi-asserted-by":"publisher","first-page":"814","DOI":"10.1016\/j.bbe.2024.09.003","article-title":"Probabilistic and explainable modeling of phase\u2013phase cross-frequency coupling patterns in EEG. Application to dyslexia diagnosis","volume":"44","author":"Castillo-Barnes","year":"2024","journal-title":"Biocybern. Biomed. Eng."},{"key":"B7","doi-asserted-by":"publisher","first-page":"746599","DOI":"10.3389\/fneur.2021.746599","article-title":"Tai chi and yoga for improving balance on one leg: a neuroimaging and biomechanics study","volume":"12","author":"Chen","year":"2021","journal-title":"Front. Neurology"},{"key":"B8","doi-asserted-by":"publisher","first-page":"679412","DOI":"10.3389\/fnagi.2021.679412","article-title":"Visual effect on brain connectome that scales feedforward and feedback processes of aged postural system during unstable stance","volume":"13","author":"Chen","year":"2021","journal-title":"Front. Aging Neurosci."},{"key":"B9","doi-asserted-by":"publisher","first-page":"875","DOI":"10.1162\/jocn.2009.21062","article-title":"Good vibrations: cross-frequency coupling in the human nucleus accumbens during reward processing","volume":"21","author":"Cohen","year":"2009","journal-title":"J. Cognitive Neuroscience"},{"key":"B10","doi-asserted-by":"publisher","first-page":"1544","DOI":"10.1016\/j.neuroimage.2008.05.061","article-title":"Is there a \u201cneural efficiency\u201d in athletes? A high-resolution EEG study","volume":"42","author":"Del Percio","year":"2008","journal-title":"Neuroimage"},{"key":"B11","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1016\/j.brainresbull.2009.02.001","article-title":"\u201cNeural efficiency\u201d of athletes\u2019 brain for upright standing: a high-resolution EEG study","volume":"79","author":"Del Percio","year":"2009","journal-title":"Brain Research Bulletin"},{"key":"B12","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.neuroscience.2018.10.040","article-title":"Electroencephalographic correlates of continuous postural tasks of increasing difficulty","volume":"395","author":"Edwards","year":"2018","journal-title":"Neuroscience"},{"key":"B13","doi-asserted-by":"publisher","first-page":"704","DOI":"10.1038\/35094565","article-title":"Dynamic predictions: oscillations and synchrony in top\u2013down processing","volume":"2","author":"Engel","year":"2001","journal-title":"Nat. Rev. Neurosci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"474","DOI":"10.1016\/j.tics.2005.08.011","article-title":"A mechanism for cognitive dynamics: neuronal communication through neuronal coherence","volume":"9","author":"Fries","year":"2005","journal-title":"Trends Cognitive Sciences"},{"key":"B15","doi-asserted-by":"publisher","first-page":"525","DOI":"10.1152\/jn.00067.2019","article-title":"Age-related changes in leg proprioception: implications for postural control","volume":"122","author":"Henry","year":"2019","journal-title":"J. Neurophysiology"},{"key":"B16","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/j.humov.2016.11.002","article-title":"Cortical activation during balancing on a balance board","volume":"51","author":"Herold","year":"2017","journal-title":"Hum. Movement Science"},{"key":"B49","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1016\/j.jbiomech.2005.12.016","article-title":"The equations of motion for a standing human reveal three mechanisms for balance[J]","volume":"40","author":"Hof","year":"2007","journal-title":"J. biomech."},{"key":"B17","doi-asserted-by":"publisher","first-page":"575","DOI":"10.1152\/jn.2001.86.2.575","article-title":"Somatosensory loss increases vestibulospinal sensitivity","volume":"86","author":"Horak","year":"2001","journal-title":"J. Neurophysiology"},{"key":"B18","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.neulet.2015.02.049","article-title":"Cortical processes associated with continuous balance control as revealed by EEG spectral power","volume":"592","author":"H\u00fclsd\u00fcnker","year":"2015","journal-title":"Neurosci. Letters"},{"key":"B19","doi-asserted-by":"publisher","first-page":"308","DOI":"10.1016\/j.jbiomech.2013.10.002","article-title":"The effect of leg preference on postural stability in healthy athletes","volume":"47","author":"Huurnink","year":"2014","journal-title":"J. Biomechanics"},{"key":"B20","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1016\/j.tins.2015.09.001","article-title":"Neural cross-frequency coupling: connecting architectures, mechanisms, and functions","volume":"38","author":"Hyafil","year":"2015","journal-title":"Trends Neurosciences"},{"key":"B21","doi-asserted-by":"publisher","first-page":"3294","DOI":"10.1519\/JSC.0000000000002210","article-title":"Sex-related differences in the accuracy of estimating target force using percentages of maximal voluntary isometric contractions vs. ratings of perceived exertion during isometric muscle actions","volume":"32","author":"Keller","year":"2018","journal-title":"J. Strength & Cond. Res."},{"key":"B22","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1016\/j.bbe.2020.12.008","article-title":"A model of human postural control inspired by separated human sensory systems","volume":"41","author":"Keyvanara","year":"2021","journal-title":"Biocybern. Biomed. Eng."},{"key":"B23","doi-asserted-by":"publisher","first-page":"e13628","DOI":"10.1016\/j.heliyon.2023.e13628","article-title":"Single-leg stance on a challenging surface can enhance cortical activation in the right hemisphere\u2013A case study","volume":"9","author":"Khan","year":"2023","journal-title":"Heliyon"},{"key":"B24","doi-asserted-by":"publisher","first-page":"4692","DOI":"10.1016\/j.physa.2012.05.034","article-title":"Evaluation of the temporal structure of postural sway fluctuations based on a comprehensive set of analysis tools","volume":"391","author":"Kirchner","year":"2012","journal-title":"Stat. Mech. Its Appl."},{"key":"B25","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.neuroimage.2009.02.006","article-title":"Automated cortical projection of EEG sensors: anatomical correlation via the international 10\u201310 system","volume":"46","author":"Koessler","year":"2009","journal-title":"Neuroimage"},{"key":"B26","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/j.neuroscience.2020.01.029","article-title":"Modulations of inter-hemispherical phase coupling in human single leg stance","volume":"430","author":"Lehmann","year":"2020","journal-title":"Neuroscience"},{"key":"B27","doi-asserted-by":"publisher","first-page":"3008","DOI":"10.1109\/tsp.2018.2821627","article-title":"Mutual information in frequency and its application to measure cross-frequency coupling in epilepsy","volume":"66","author":"Malladi","year":"2018","journal-title":"IEEE Trans. Signal Processing"},{"key":"B28","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1007\/s00221-005-0256-y","article-title":"Multisensory control of human upright stance","volume":"171","author":"Maurer","year":"2006","journal-title":"Exp. Brain Research"},{"key":"B29","doi-asserted-by":"publisher","first-page":"272","DOI":"10.3389\/fnbeh.2015.00272","article-title":"Changes in cortical activity associated with adaptive behavior during repeated balance perturbation of unpredictable timing","volume":"9","author":"Mierau","year":"2015","journal-title":"Front. Behav. Neurosci."},{"key":"B30","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12984-018-0465-9","article-title":"On the use of approximate entropy and sample entropy with centre of pressure time-series","volume":"15","author":"Montesinos","year":"2018","journal-title":"J. Neuroengineering Rehabilitation"},{"key":"B31","doi-asserted-by":"publisher","first-page":"407","DOI":"10.1123\/jab.2013-0230","article-title":"One-leg standing performance and muscle activity: are there limb differences?","volume":"30","author":"Muehlbauer","year":"2014","journal-title":"J. Appl. Biomechanics"},{"key":"B32","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1016\/j.brainresrev.2009.11.006","article-title":"Characteristics of the athletes' brain: evidence from neurophysiology and neuroimaging","volume":"62","author":"Nakata","year":"2010","journal-title":"Brain Research Reviews"},{"key":"B33","doi-asserted-by":"publisher","first-page":"117","DOI":"10.5277\/ABB-01060-2017-02","article-title":"Quiet standing postural sway of 10-to 13-year-old,national-level, female acrobatic gymnasts","volume":"20","author":"Opala-Berdzik","year":"2018","journal-title":"Acta Bioengineering Biomechanics"},{"key":"B34","doi-asserted-by":"publisher","first-page":"e3990","DOI":"10.1371\/journal.pone.0003990","article-title":"Gamma power is phase-locked to posterior alpha activity","volume":"3","author":"Osipova","year":"2008","journal-title":"PloS One"},{"key":"B35","doi-asserted-by":"publisher","first-page":"ENEURO.0207","DOI":"10.1523\/ENEURO.0207-18.2018","article-title":"Differentiation in theta and beta electrocortical activity between visual and physical perturbations to walking and standing balance","volume":"5","author":"Peterson","year":"2018","journal-title":"Eneuro"},{"key":"B36","doi-asserted-by":"publisher","first-page":"H2039","DOI":"10.1152\/ajpheart.2000.278.6.H2039","article-title":"Physiological time-series analysis using approximate entropy and sample entropy","volume":"278","author":"Richman","year":"2000","journal-title":"Am. J. Physiology-Heart Circulatory Physiology"},{"key":"B37","doi-asserted-by":"publisher","first-page":"125","DOI":"10.3389\/fnins.2019.00125","article-title":"Cross-frequency coupling based neuromodulation for treating neurological disorders","volume":"13","author":"Salimpour","year":"2019","journal-title":"Front. Neuroscience"},{"key":"B38","doi-asserted-by":"publisher","first-page":"e20515","DOI":"10.7554\/eLife.20515","article-title":"On cross-frequency phase-phase coupling between theta and gamma oscillations in the hippocampus","volume":"5","author":"Scheffer-Teixeira","year":"2016","journal-title":"Elife"},{"key":"B39","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.tins.2008.09.012","article-title":"Low-frequency neuronal oscillations as instruments of sensory selection","volume":"32","author":"Schroeder","year":"2009","journal-title":"Trends Neurosciences"},{"key":"B40","doi-asserted-by":"publisher","first-page":"188","DOI":"10.1016\/j.biopsycho.2005.10.005","article-title":"Neural underpinning of postural responses to visual field motion","volume":"72","author":"Slobounov","year":"2006","journal-title":"Biol. Psychol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"3454","DOI":"10.1093\/cercor\/bhac283","article-title":"Midfrontal theta dynamics index the monitoring of postural stability","volume":"33","author":"Stokkermans","year":"2023","journal-title":"Cereb. Cortex"},{"key":"B42","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1016\/j.gaitpost.2019.05.011","article-title":"Balance and the brain: a review of structural brain correlates of postural balance and balance training in humans","volume":"71","author":"Surgent","year":"2019","journal-title":"Gait & Posture"},{"key":"B43","doi-asserted-by":"publisher","first-page":"e1001936","DOI":"10.1371\/journal.pbio.1001936","article-title":"Dynamic changes in phase-amplitude coupling facilitate spatial attention control in fronto-parietal cortex","volume":"12","author":"Szczepanski","year":"2014","journal-title":"PLoS Biology"},{"key":"B44","doi-asserted-by":"publisher","first-page":"1492","DOI":"10.1016\/j.neuroimage.2011.05.078","article-title":"Long-term effects of motor training on resting-state networks and underlying brain structure","volume":"57","author":"Taubert","year":"2011","journal-title":"Neuroimage"},{"key":"B45","doi-asserted-by":"publisher","first-page":"919184","DOI":"10.3389\/fphys.2022.919184","article-title":"Adaptations of postural sway dynamics and cortical response to unstable stance with stroboscopic vision in older adults","volume":"13","author":"Tsai","year":"2022","journal-title":"Front. Physiology"},{"key":"B46","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1016\/j.neulet.2014.06.017","article-title":"Frequency characteristics of cortical activity associated with perturbations to upright stability","volume":"578","author":"Varghese","year":"2014","journal-title":"Neurosci. Letters"},{"key":"B47","article-title":"Relationship between leg dominance tests and type of task","author":"Velotta","year":"2011","journal-title":"ISBS-Conference Proc. Arch."},{"key":"B48","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1016\/j.physa.2016.02.012","article-title":"Spurious cross-frequency amplitude\u2013amplitude coupling in nonstationary, nonlinear signals","volume":"454","author":"Yeh","year":"2016","journal-title":"Phys. A Stat. Mech. Its Appl."}],"container-title":["Frontiers in Physiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphys.2025.1700523\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,18]],"date-time":"2025-12-18T06:45:06Z","timestamp":1766040306000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphys.2025.1700523\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,18]]},"references-count":49,"alternative-id":["10.3389\/fphys.2025.1700523"],"URL":"https:\/\/doi.org\/10.3389\/fphys.2025.1700523","relation":{},"ISSN":["1664-042X"],"issn-type":[{"value":"1664-042X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,18]]},"article-number":"1700523"}}