{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T18:07:02Z","timestamp":1790100422594,"version":"4.0.1"},"reference-count":131,"publisher":"Oxford University Press (OUP)","issue":"9","license":[{"start":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T00:00:00Z","timestamp":1790035200000},"content-version":"vor","delay-in-days":21,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Leverhulme Early Career Fellowship","award":["ECF-2021-628"],"award-info":[{"award-number":["ECF-2021-628"]}]},{"name":"Leverhulme Early Career Fellowship","award":["ECF-2021-628"],"award-info":[{"award-number":["ECF-2021-628"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Rhythmic light stimulation offers solutions to innumerable cognitive and neurological disorders. However, like any neuromodulatory technique, responses to rhythmic light stimulation are highly variable, producing challenges in replicating lab-based studies and translating findings to the clinic. Across 3 magnetoencephalography (MEG) and electroencephalography (EEG) experiments, we show that this variability can, in part, be attributed to rhythmic light stimulation eliciting multiple, coexisting neural responses which have separable influences on behavior. Specifically, we find that rhythmic light stimulation produces neural responses at both the fundamental (f) and second harmonic (2f) frequencies, and that these responses are differentially shaped by endogenous oscillatory dynamics that vary across participants. Importantly, these harmonic responses separably contribute to perception, with gamma-band responses supporting the representation of stimulus-specific information, and alpha-band responses causally contributing to near-threshold visual perception. We reproduce these effects across datasets, paradigms, and oscillatory bands, suggesting that the multiple, concurrent oscillatory responses elicited by rhythmic light stimulation are a robust and pervasive phenomenon. We propose that the complexity of neural responses to rhythmic stimulation can explain why there is substantial variability between studies using these techniques, and that understanding these complex responses may help advance neuromodulatory technologies for both fundamental and clinical neuroscience.<\/jats:p>","DOI":"10.1093\/cercor\/bhag152","type":"journal-article","created":{"date-parts":[[2026,9,4]],"date-time":"2026-09-04T11:28:40Z","timestamp":1788521320000},"source":"Crossref","is-referenced-by-count":0,"title":["Rhythmic stimulation elicits multiple, concurrent neural responses"],"prefix":"10.1093","volume":"36","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-8600-4480","authenticated-orcid":false,"given":"Benjamin J","family":"Griffiths","sequence":"first","affiliation":[{"name":"Centre for Human Brain Health, University of Birmingham , Birmingham B15 2TT ,","place":["United Kingdom"]},{"name":"School of Psychology, University of Nottingham , Nottingham NG7 2RD ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katharina","family":"Duecker","sequence":"additional","affiliation":[{"name":"Department of Neuroscience, Brown University , Providence, Rhode Island 02906 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Camille","family":"Fakche","sequence":"additional","affiliation":[{"name":"Lyon Neuroscience Research Center, Universit\u00e9 de Lyon , Lyon 69500 ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura","family":"Dugu\u00e9","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Paris Cit\u00e9, CNRS, Integrative Neuroscience and Cognition Center , Paris 75006 ,","place":["France"]},{"name":"Institut Universitaire de France (IUF) , Paris ,","place":["France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew","family":"Quinn","sequence":"additional","affiliation":[{"name":"Centre for Human Brain Health, University of Birmingham , Birmingham B15 2TT ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ole","family":"Jensen","sequence":"additional","affiliation":[{"name":"Centre for Human Brain Health, University of Birmingham , Birmingham B15 2TT ,","place":["United Kingdom"]},{"name":"Oxford Centre for Human Brain Activity, Department of Psychiatry, University of Oxford , Oxford OX3 7JX ,","place":["United Kingdom"]},{"name":"Department of Experimental Psychology, University of Oxford , Oxford OX1 3EL ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,22]]},"reference":[{"key":"2026092213411070600_ref1","doi-asserted-by":"publisher","first-page":"929","DOI":"10.1016\/j.neuron.2019.04.011","article-title":"Gamma entrainment binds higher-order brain regions and offers neuroprotection","volume":"102","author":"Adaikkan","year":"2019","journal-title":"Neuron"},{"key":"2026092213411070600_ref2","doi-asserted-by":"publisher","first-page":"eabj9782","DOI":"10.1126\/sciadv.abj9782","article-title":"Supramodality of neural entrainment: rhythmic visual stimulation causally enhances auditory working memory performance","volume":"8","author":"Albouy","year":"2022","journal-title":"Sci Adv"},{"key":"2026092213411070600_ref3","doi-asserted-by":"publisher","first-page":"11262","DOI":"10.1523\/JNEUROSCI.5867-12.2013","article-title":"Transcranial alternating current stimulation modulates large-scale cortical network activity by network resonance","volume":"33","author":"Ali","year":"2013","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref4","doi-asserted-by":"publisher","first-page":"3328","DOI":"10.1016\/j.celrep.2018.02.091","article-title":"Hippocampal theta-gamma coupling reflects state-dependent information processing in decision making","volume":"22","author":"Amemiya","year":"2018","journal-title":"Cell Rep"},{"key":"2026092213411070600_ref5","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1101\/005926","article-title":"Untangling cross-frequency coupling in neuroscience","volume":"31","author":"Aru","year":"2014","journal-title":"Curr Opin Neurobiol"},{"key":"2026092213411070600_ref6","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/S0167-8760(00)00145-8","article-title":"Gamma, alpha, delta, and theta oscillations govern cognitive processes","volume":"39","author":"Ba\u015far","year":"2001","journal-title":"Int J Psychophysiol"},{"key":"2026092213411070600_ref7","doi-asserted-by":"publisher","first-page":"423","DOI":"10.1523\/JNEUROSCI.4122-11.2012","article-title":"Cross-frequency phase\u2013phase coupling between theta and gamma oscillations in the hippocampus","volume":"32","author":"Belluscio","year":"2012","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref8","doi-asserted-by":"publisher","first-page":"ENEURO.0182-17.2017","DOI":"10.1523\/eneuro.0182-17.2017","article-title":"Prestimulus EEG power predicts conscious awareness but not objective visual performance","volume":"4","author":"Benwell","year":"2017","journal-title":"eNeuro"},{"key":"2026092213411070600_ref9","doi-asserted-by":"publisher","first-page":"3125","DOI":"10.1111\/ejn.15166","article-title":"Low pre-stimulus EEG alpha power amplifies visual awareness but not visual sensitivity","volume":"55","author":"Benwell","year":"2021","journal-title":"Eur J Neurosci"},{"key":"2026092213411070600_ref10","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1162\/jocn_a_01591","article-title":"Inferring causality from noninvasive brain stimulation in cognitive neuroscience","volume":"33","author":"Bergmann","year":"2021","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref11","doi-asserted-by":"publisher","first-page":"670","DOI":"10.1016\/j.neuron.2014.03.013","article-title":"Slow and fast gamma rhythms coordinate different spatial coding modes in hippocampal place cells","volume":"82","author":"Bieri","year":"2014","journal-title":"Neuron"},{"key":"2026092213411070600_ref12","doi-asserted-by":"publisher","first-page":"16","DOI":"10.3389\/fninf.2015.00016","article-title":"The PREP pipeline: standardized preprocessing for large-scale EEG analysis","volume":"9","author":"Bigdely-Shamlo","year":"2015","journal-title":"Front Neuroinform"},{"key":"2026092213411070600_ref13","doi-asserted-by":"publisher","first-page":"3156","DOI":"10.1038\/s41467-024-47263-y","article-title":"Multisensory flicker modulates widespread brain networks and reduces interictal epileptiform discharges","volume":"15","author":"Blanpain","year":"2024","journal-title":"Nat Commun"},{"key":"2026092213411070600_ref14","doi-asserted-by":"publisher","first-page":"ENEURO.0282-24.2024","DOI":"10.1523\/ENEURO.0282-24.2024","article-title":"Visual processing by hierarchical and dynamic multiplexing","volume":"11","author":"Bonnefond","year":"2024","journal-title":"eNeuro"},{"key":"2026092213411070600_ref15","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1038\/nn.4497","article-title":"Dynamic models of large-scale brain activity","volume":"20","author":"Breakspear","year":"2017","journal-title":"Nat Neurosci"},{"key":"2026092213411070600_ref16","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1177\/0049124104268644","article-title":"Multimodel inference: understanding AIC and BIC in model selection","volume":"33","author":"Burnham","year":"2004","journal-title":"Sociol Methods Res"},{"key":"2026092213411070600_ref17","doi-asserted-by":"publisher","first-page":"7869","DOI":"10.1523\/jneurosci.0113-09.2009","article-title":"The phase of ongoing EEG oscillations predicts visual perception","volume":"29","author":"Busch","year":"2009","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref18","doi-asserted-by":"publisher","first-page":"203","DOI":"10.1146\/annurev-neuro-062111-150444","article-title":"Mechanisms of gamma oscillations","volume":"35","author":"Buzs\u00e1ki","year":"2012","journal-title":"Annu Rev Neurosci"},{"key":"2026092213411070600_ref19","doi-asserted-by":"publisher","first-page":"e14543","DOI":"10.1371\/journal.pone.0014543","article-title":"Steady-state visual evoked potentials can be explained by temporal superposition of transient event-related responses","volume":"6","author":"Capilla","year":"2011","journal-title":"PLoS One"},{"key":"2026092213411070600_ref20","doi-asserted-by":"publisher","first-page":"e0278412","DOI":"10.1371\/journal.pone.0278412","article-title":"Gamma frequency sensory stimulation in mild probable Alzheimer\u2019s dementia patients: results of feasibility and pilot studies","volume":"17","author":"Chan","year":"2022","journal-title":"PLoS One"},{"key":"2026092213411070600_ref21","doi-asserted-by":"publisher","first-page":"3143","DOI":"10.1016\/j.cub.2017.09.001","article-title":"Theta phase synchronization is the glue that binds human associative memory","volume":"27","author":"Clouter","year":"2017","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref22","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/9609.001.0001","volume-title":"Analyzing neural time series data: theory and practice","author":"Cohen","year":"2014"},{"key":"2026092213411070600_ref23","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.neuroimage.2016.11.036","article-title":"Rhythmic entrainment source separation: optimizing analyses of neural responses to rhythmic sensory stimulation","volume":"147","author":"Cohen","year":"2017","journal-title":"NeuroImage"},{"key":"2026092213411070600_ref24","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1016\/j.tics.2016.12.008","article-title":"Brain oscillations and the importance of waveform shape","volume":"21","author":"Cole","year":"2017","journal-title":"Trends Cogn Sci"},{"key":"2026092213411070600_ref25","doi-asserted-by":"publisher","first-page":"4830","DOI":"10.1523\/JNEUROSCI.2208-16.2017","article-title":"Nonsinusoidal beta oscillations reflect cortical pathophysiology in Parkinson\u2019s disease","volume":"37","author":"Cole","year":"2017","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref26","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1038\/nrn.2016.21","article-title":"Rhythms of the hippocampal network","volume":"17","author":"Colgin","year":"2016","journal-title":"Nat Rev Neurosci"},{"key":"2026092213411070600_ref27","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1038\/nature08573","article-title":"Frequency of gamma oscillations routes flow of information in the hippocampus","volume":"462","author":"Colgin","year":"2009","journal-title":"Nature"},{"key":"2026092213411070600_ref28","doi-asserted-by":"publisher","first-page":"1869","DOI":"10.1152\/jn.1998.79.4.1869","article-title":"Synchronous period-doubling in flicker vision of salamander and man","volume":"79","author":"Crevier","year":"1998","journal-title":"J Neurophysiol"},{"key":"2026092213411070600_ref29","doi-asserted-by":"publisher","first-page":"1655","DOI":"10.1038\/s41593-020-00744-x","article-title":"Parameterizing neural power spectra into periodic and aperiodic components","volume":"23","author":"Donoghue","year":"2020","journal-title":"Nat Neurosci"},{"key":"2026092213411070600_ref30","doi-asserted-by":"publisher","first-page":"026003","DOI":"10.1088\/1741-2552\/acbc4a","article-title":"How to entrain a selected neuronal rhythm but not others: open-loop dithered brain stimulation for selective entrainment","volume":"20","author":"Duchet","year":"2023","journal-title":"J Neural Eng"},{"key":"2026092213411070600_ref31","doi-asserted-by":"publisher","first-page":"6684","DOI":"10.1523\/JNEUROSCI.3134-20.2021","article-title":"No evidence for entrainment: endogenous gamma oscillations and responses to rhythmic visual stimulation coexist in visual cortex","volume":"41","author":"Duecker","year":"2021","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref32","doi-asserted-by":"publisher","first-page":"e1234242024","DOI":"10.1523\/JNEUROSCI.1234-24.2024","article-title":"Challenges and approaches in the study of neural entrainment","volume":"44","author":"Duecker","year":"2024","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref33","doi-asserted-by":"publisher","first-page":"11889","DOI":"10.1523\/JNEUROSCI.1161-11.2011","article-title":"The phase of ongoing oscillations mediates the causal relation between brain excitation and visual perception","volume":"31","author":"Dugu\u00e9","year":"2011","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref34","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1016\/0013-4694(71)90183-0","article-title":"Analysis of the interrelations between frequency bands of the EEG by means of the bispectrum a preliminary study","volume":"31","author":"Dumermuth","year":"1971","journal-title":"Electroencephalogr Clin Neurophysiol"},{"key":"2026092213411070600_ref35","doi-asserted-by":"publisher","first-page":"1670","DOI":"10.1152\/jn.00315.2021","article-title":"Automatic decomposition of electrophysiological data into distinct non-sinusoidal oscillatory modes","volume":"126","author":"Fabus","year":"2021","journal-title":"J Neurophysiol"},{"key":"2026092213411070600_ref36","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1162\/jocn_a_02075","article-title":"Perceptual cycles travel across retinotopic space","volume":"36","author":"Fakche","year":"2024","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref37","doi-asserted-by":"publisher","volume-title":"ENeuro","author":"Fakche","DOI":"10.1523\/ENEURO.0244-21.2022"},{"key":"2026092213411070600_ref38","doi-asserted-by":"publisher","first-page":"1542","DOI":"10.1016\/j.neuroimage.2003.10.045","article-title":"The temporal frequency tuning of human visual cortex investigated using synthetic aperture magnetometry","volume":"21","author":"Fawcett","year":"2004","journal-title":"Neuroimage"},{"key":"2026092213411070600_ref39","doi-asserted-by":"publisher","first-page":"119047","DOI":"10.1016\/j.neuroimage.2022.119047","article-title":"FLUX: a pipeline for MEG analysis","volume":"253","author":"Ferrante","year":"2022","journal-title":"Neuroimage"},{"key":"2026092213411070600_ref40","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/j.brs.2022.01.001","article-title":"tACS phase-specifically biases brightness perception of flickering light","volume":"15","author":"Fiene","year":"2022","journal-title":"Brain Stimul"},{"key":"2026092213411070600_ref41","doi-asserted-by":"publisher","first-page":"220","DOI":"10.1016\/j.neuron.2015.09.034","article-title":"Rhythms for cognition: communication through coherence","volume":"88","author":"Fries","year":"2015","journal-title":"Neuron"},{"key":"2026092213411070600_ref42","doi-asserted-by":"publisher","first-page":"18556","DOI":"10.1523\/JNEUROSCI.2164-11.2011","article-title":"Oscillatory sensory selection mechanisms during intersensory attention to rhythmic auditory and visual inputs: a human electrocorticographic investigation","volume":"31","author":"Gomez-Ramirez","year":"2011","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref43","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1371\/journal.pone.0060035","article-title":"Alpha-band rhythms in visual task performance: phase-locking by rhythmic sensory stimulation","volume":"8","author":"Graaf","year":"2013","journal-title":"PLoS One"},{"key":"2026092213411070600_ref44","doi-asserted-by":"publisher","first-page":"267","DOI":"10.3389\/fnins.2013.00267","article-title":"MEG and EEG data analysis with MNE-Python","volume":"7","author":"Gramfort","year":"2013","journal-title":"Front Neurosci"},{"key":"2026092213411070600_ref45","doi-asserted-by":"publisher","first-page":"832","DOI":"10.1016\/j.tins.2023.07.003","article-title":"Gamma oscillations and episodic memory","volume":"46","author":"Griffiths","year":"2023","journal-title":"Trends Neurosci"},{"key":"2026092213411070600_ref46","doi-asserted-by":"publisher","first-page":"21834","DOI":"10.1073\/pnas.1914180116","article-title":"Directional coupling of slow and fast hippocampal gamma with neocortical alpha\/beta oscillations in human episodic memory","volume":"166","author":"Griffiths","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026092213411070600_ref47","doi-asserted-by":"publisher","first-page":"118454","DOI":"10.1016\/j.neuroimage.2021.118454","article-title":"Disentangling neocortical alpha\/beta and hippocampal theta\/gamma oscillations in human episodic memory formation","volume":"242","author":"Griffiths","year":"2021","journal-title":"Neuroimage"},{"key":"2026092213411070600_ref48","doi-asserted-by":"publisher","first-page":"3736","DOI":"10.1038\/s41467-022-31407-z","article-title":"Rhythmic interactions between the mediodorsal thalamus and prefrontal cortex precede human visual perception","volume":"13","author":"Griffiths","year":"2022","journal-title":"Nat Commun"},{"key":"2026092213411070600_ref49","doi-asserted-by":"publisher","DOI":"10.1101\/2023.07.21.550057","article-title":"Imperceptible gamma-band sensory stimulation enhances episodic memory retrieval","author":"Griffiths","year":"2023"},{"key":"2026092213411070600_ref50","doi-asserted-by":"publisher","first-page":"H1","DOI":"10.1152\/ajpheart.1988.254.1.H1","article-title":"Phase-locked rhythms in periodically stimulated heart cell aggregates","volume":"254","author":"Guevara","year":"1988","journal-title":"Am J Phys Heart Circ Phys"},{"key":"2026092213411070600_ref51","doi-asserted-by":"publisher","first-page":"10173","DOI":"10.1523\/JNEUROSCI.1163-17.2017","article-title":"Individual alpha peak frequency predicts 10 Hz flicker effects on selective attention","volume":"37","author":"Gulbinaite","year":"2017","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref52","doi-asserted-by":"publisher","first-page":"116146","DOI":"10.1016\/j.neuroimage.2019.116146","article-title":"Attention differentially modulates the amplitude of resonance frequencies in the visual cortex","volume":"203","author":"Gulbinaite","year":"2019","journal-title":"Neuroimage"},{"key":"2026092213411070600_ref53","doi-asserted-by":"publisher","first-page":"e3001466","DOI":"10.1371\/journal.pbio.3001466","article-title":"Multiple gamma rhythms carry distinct spatial frequency information in primary visual cortex","volume":"19","author":"Han","year":"2021","journal-title":"PLoS Biol"},{"key":"2026092213411070600_ref54","doi-asserted-by":"publisher","first-page":"904","DOI":"10.1016\/j.cub.2014.03.007","article-title":"Entrainment of prefrontal beta oscillations induces an endogenous echo and impairs memory formation","volume":"24","author":"Hanslmayr","year":"2014","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref55","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.tins.2015.11.004","article-title":"Oscillations and episodic memory\u2014addressing the synchronization\/desynchronization conundrum","volume":"39","author":"Hanslmayr","year":"2016","journal-title":"Trends Neurosci"},{"key":"2026092213411070600_ref56","doi-asserted-by":"publisher","first-page":"485","DOI":"10.1016\/j.tins.2019.04.004","article-title":"Modulating human memory via entrainment of brain oscillations","volume":"42","author":"Hanslmayr","year":"2019","journal-title":"Trends Neurosci"},{"key":"2026092213411070600_ref57","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1017\/S0952523800008154","article-title":"Temporal-frequency selectivity in monkey visual cortex","volume":"13","author":"Hawken","year":"1996","journal-title":"Vis Neurosci"},{"key":"2026092213411070600_ref58","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/j.biopsycho.2012.04.008","article-title":"Presence of strong harmonics during visual entrainment: a magnetoencephalography study","volume":"91","author":"Heinrichs-Graham","year":"2012","journal-title":"Biol Psychol"},{"key":"2026092213411070600_ref59","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.copsyc.2018.12.016","article-title":"Neural entrainment and network resonance in support of top-down guided attention","volume":"29","author":"Helfrich","year":"2019","journal-title":"Curr Opin Psychol"},{"key":"2026092213411070600_ref60","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.tics.2014.12.009","article-title":"Gamma oscillations in visual cortex: the stimulus matters","volume":"19","author":"Hermes","year":"2015","journal-title":"Trends Cogn Sci"},{"key":"2026092213411070600_ref61","doi-asserted-by":"publisher","first-page":"2951","DOI":"10.1093\/cercor\/bhu091","article-title":"Stimulus dependence of gamma oscillations in human visual cortex","volume":"25","author":"Hermes","year":"2015","journal-title":"Cereb Cortex"},{"key":"2026092213411070600_ref62","doi-asserted-by":"publisher","first-page":"346","DOI":"10.1007\/s002210100682","article-title":"Human EEG responses to 1\u2013100 Hz flicker: resonance phenomena in visual cortex and their potential correlation to cognitive phenomena","volume":"137","author":"Herrmann","year":"2001","journal-title":"Exp Brain Res"},{"key":"2026092213411070600_ref63","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.ijpsycho.2015.02.003","article-title":"EEG oscillations: from correlation to causality","volume":"103","author":"Herrmann","year":"2016","journal-title":"Int J Psychophysiol"},{"key":"2026092213411070600_ref64","doi-asserted-by":"publisher","first-page":"119462","DOI":"10.1016\/j.neuroimage.2022.119462","article-title":"The relationship between frequency content and representational dynamics in the decoding of neurophysiological data","volume":"260","author":"Higgins","year":"2022","journal-title":"Neuroimage"},{"key":"2026092213411070600_ref65","doi-asserted-by":"publisher","first-page":"903","DOI":"10.1098\/rspa.1998.0193","article-title":"The empirical mode decomposition and the Hilbert spectrum for nonlinear and non-stationary time series analysis","volume":"454","author":"Huang","year":"1998","journal-title":"Proc R Soc Lond A: Math Phys Eng Sci"},{"key":"2026092213411070600_ref66","doi-asserted-by":"publisher","first-page":"e100238","DOI":"10.7554\/eLife.100238","article-title":"The gamma rhythm as a guardian of brain health","volume":"13","author":"Ichim","year":"2024","journal-title":"Elife"},{"key":"2026092213411070600_ref67","doi-asserted-by":"publisher","first-page":"1","DOI":"10.7554\/eLife.43620.001","article-title":"Multiple mechanisms link prestimulus neural oscillations to sensory responses","volume":"8","author":"Iemi","year":"2019","journal-title":"Elife"},{"key":"2026092213411070600_ref68","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1073\/pnas.1714058114","article-title":"Direct electrical stimulation of the amygdala enhances declarative memory in humans","volume":"115","author":"Inman","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2026092213411070600_ref69","doi-asserted-by":"publisher","first-page":"422","DOI":"10.1162\/jocn_a_00494","article-title":"Direct control of visual perception with phase-specific modulation of posterior parietal cortex","volume":"26","author":"Jaegle","year":"2014","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref70","volume-title":"JASP (version 0.9) [computer software]","author":"JASP-Team","year":"2018"},{"key":"2026092213411070600_ref71","doi-asserted-by":"publisher","first-page":"18","DOI":"10.1007\/s10548-009-0078-5","article-title":"Saccade related gamma-band activity in intracerebral EEG: dissociating neural from ocular muscle activity","volume":"22","author":"Jerbi","year":"2009","journal-title":"Brain Topogr"},{"key":"2026092213411070600_ref72","doi-asserted-by":"publisher","first-page":"20210088","DOI":"10.1098\/rsfs.2021.0088","article-title":"Principles, mechanisms and functions of entrainment in biological oscillators","volume":"12","author":"Jim\u00e9nez","year":"2022","journal-title":"Interface Focus"},{"key":"2026092213411070600_ref73","doi-asserted-by":"publisher","first-page":"956","DOI":"10.1016\/j.cels.2024.09.001","article-title":"Entrainment and multi-stability of the p53 oscillator in human cells","volume":"15","author":"Jim\u00e9nez","year":"2024","journal-title":"Cell Syst"},{"key":"2026092213411070600_ref74","doi-asserted-by":"publisher","first-page":"3119","DOI":"10.1523\/JNEUROSCI.1633-18.2019","article-title":"Stimulus-driven brain rhythms within the alpha band: the attentional-modulation conundrum","volume":"39","author":"Keitel","year":"2019","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref75","doi-asserted-by":"publisher","first-page":"2991","DOI":"10.1111\/ejn.15740","article-title":"Rhythms in cognition: the evidence revisited","volume":"55","author":"Keitel","year":"2022","journal-title":"Eur J Neurosci"},{"key":"2026092213411070600_ref76","doi-asserted-by":"publisher","first-page":"3383","DOI":"10.1016\/j.cub.2018.08.065","article-title":"An optimal oscillatory phase for pattern reactivation during memory retrieval","volume":"28","author":"Kerr\u00e9n","year":"2018","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref77","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1101\/2022.05.07.490872","article-title":"Phase separation of competing memories along the human hippocampal theta rhythm","volume":"11","author":"Kerr\u00e9n","year":"2023","journal-title":"Elife"},{"key":"2026092213411070600_ref78","doi-asserted-by":"publisher","first-page":"1875","DOI":"10.1162\/jocn.2010.21536","article-title":"Differential roles of frequency-following and frequency-doubling visual responses revealed by evoked neural harmonics","volume":"23","author":"Kim","year":"2011","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref79","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1162\/jocn_a_01058","article-title":"Power and phase of alpha oscillations reveal an interaction between spatial and temporal visual attention","volume":"29","author":"Kizuk","year":"2017","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref80","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1038\/s41582-020-00426-z","article-title":"Technology of deep brain stimulation: current status and future directions","volume":"17","author":"Krauss","year":"2021","journal-title":"Nat Rev Neurol"},{"key":"2026092213411070600_ref81","doi-asserted-by":"publisher","first-page":"33","DOI":"10.3389\/fnsys.2015.00033","article-title":"Modulating pathological oscillations by rhythmic non-invasive brain stimulation\u2014a therapeutic concept?","volume":"9","author":"Krawinkel","year":"2015","journal-title":"Front Syst Neurosci"},{"key":"2026092213411070600_ref82","doi-asserted-by":"publisher","first-page":"R890","DOI":"10.1016\/j.cub.2019.07.075","article-title":"A new unifying account of the roles of neuronal entrainment","volume":"29","author":"Lakatos","year":"2019","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref83","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1038\/s41467-018-08080-2","article-title":"Evidence that neural information flow is reversed between object perception and object reconstruction from memory","volume":"10","author":"Linde-Domingo","year":"2019","journal-title":"Nat Commun"},{"key":"2026092213411070600_ref84","doi-asserted-by":"publisher","first-page":"1002","DOI":"10.1016\/j.neuron.2013.03.007","article-title":"The theta-gamma neural code","volume":"77","author":"Lisman","year":"2013","journal-title":"Neuron"},{"key":"2026092213411070600_ref85","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.conb.2018.07.009","article-title":"Gamma oscillations in cognitive disorders","volume":"52","author":"Mably","year":"2018","journal-title":"Curr Opin Neurobiol"},{"key":"2026092213411070600_ref86","doi-asserted-by":"publisher","first-page":"256","DOI":"10.1016\/j.cell.2019.02.014","article-title":"Multi-sensory gamma stimulation ameliorates Alzheimer\u2019s-associated pathology and improves cognition","volume":"177","author":"Martorell","year":"2019","journal-title":"Cell"},{"key":"2026092213411070600_ref87","doi-asserted-by":"publisher","first-page":"2725","DOI":"10.1523\/JNEUROSCI.3963-08.2009","article-title":"To see or not to see: prestimulus \u03b1 phase predicts visual awareness","volume":"29","author":"Mathewson","year":"2009","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref88","doi-asserted-by":"publisher","first-page":"2321","DOI":"10.1162\/jocn_a_00288","article-title":"Making waves in the stream of consciousness: entraining oscillations in EEG alpha and fluctuations in visual awareness with rhythmic visual stimulation","volume":"24","author":"Mathewson","year":"2012","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref89","doi-asserted-by":"publisher","first-page":"e13029","DOI":"10.1111\/psyp.13029","article-title":"Evaluating the feasibility of the steady-state visual evoked potential (SSVEP) to study temporal attention","volume":"55","author":"Mora-Cortes","year":"2018","journal-title":"Psychophysiology"},{"key":"2026092213411070600_ref90","doi-asserted-by":"publisher","first-page":"138","DOI":"10.3389\/fnhum.2013.00138","article-title":"High-frequency brain activity and muscle artifacts in MEG\/EEG: a review and recommendations","volume":"7","author":"Muthukumaraswamy","year":"2013","journal-title":"Front Hum Neurosci"},{"key":"2026092213411070600_ref91","doi-asserted-by":"publisher","first-page":"4","DOI":"10.1167\/15.6.4","article-title":"The steady-state visual evoked potential in vision research: a review","volume":"15","author":"Norcia","year":"2015","journal-title":"J Vis"},{"key":"2026092213411070600_ref92","doi-asserted-by":"publisher","first-page":"10","DOI":"10.3389\/fnhum.2016.00010","article-title":"Modification of brain oscillations via rhythmic light stimulation provides evidence for entrainment but not for superposition of event-related responses","volume":"10","author":"Notbohm","year":"2016","journal-title":"Front Hum Neurosci"},{"key":"2026092213411070600_ref93","doi-asserted-by":"publisher","first-page":"2255","DOI":"10.1038\/s41467-019-09569-0","article-title":"Coordinated representational reinstatement in the human hippocampus and lateral temporal cortex during episodic memory retrieval","volume":"10","author":"Estefan","year":"2019","journal-title":"Nat Commun"},{"key":"2026092213411070600_ref94","doi-asserted-by":"publisher","first-page":"1899","DOI":"10.1093\/cercor\/bhl098","article-title":"Topography of cortical activation differs for fundamental and harmonic frequencies of the steady-state visual-evoked responses. An EEG and PET H215O study","volume":"17","author":"Pastor","year":"2007","journal-title":"Cereb Cortex"},{"key":"2026092213411070600_ref95","doi-asserted-by":"publisher","first-page":"4156","DOI":"10.1093\/cercor\/bhab472","article-title":"Theta:gamma phase coupling and evoked gamma activity reflect the fidelity of mental templates during memory matching in visual perception","volume":"32","author":"Peylo","year":"2022","journal-title":"Cereb Cortex"},{"key":"2026092213411070600_ref96","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9780511755743","volume-title":"Synchronization: a universal concept in nonlinear sciences","author":"Pikovsky","year":"2001","edition":"1"},{"key":"2026092213411070600_ref97","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1038\/s44271-024-00158-6","article-title":"Sensory stimulation enhances visual working memory capacity","volume":"2","author":"Pileckyte","year":"2024","journal-title":"Commun Psychol"},{"key":"2026092213411070600_ref98","doi-asserted-by":"publisher","first-page":"174","DOI":"10.1038\/s41593-017-0054-4","article-title":"Studying and modifying brain function with non-invasive brain stimulation","volume":"21","author":"Polan\u00eda","year":"2018","journal-title":"Nat Neurosci"},{"key":"2026092213411070600_ref99","doi-asserted-by":"publisher","first-page":"2977","DOI":"10.21105\/joss.02977","article-title":"EMD: empirical mode decomposition and Hilbert\u2013Huang spectral analyses in python","volume":"6","author":"Quinn","year":"2021","journal-title":"J Open Source Softw"},{"key":"2026092213411070600_ref100","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1162\/jocn_a_01763","article-title":"Harmonic amplitude summation for frequency-tagging analysis","volume":"33","author":"Retter","year":"2021","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref101","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1080\/17588928.2018.1521386","article-title":"Entrainment enhances theta oscillations and improves episodic memory","volume":"9","author":"Roberts","year":"2018","journal-title":"Cogn Neurosci"},{"key":"2026092213411070600_ref102","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.cortex.2019.05.012","article-title":"The relevance of alpha phase in human perception","volume":"120","author":"Ruzzoli","year":"2019","journal-title":"Cortex"},{"key":"2026092213411070600_ref103","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1162\/jocn.2009.21126","article-title":"Optimizing functional accuracy of TMS in cognitive studies: a comparison of methods","volume":"21","author":"Sack","year":"2009","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref104","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.concog.2017.02.005","article-title":"Prestimulus alpha-band power biases visual discrimination confidence, but not accuracy","volume":"54","author":"Samaha","year":"2017","journal-title":"Conscious Cogn"},{"key":"2026092213411070600_ref105","doi-asserted-by":"publisher","DOI":"10.1016\/j.tics.2020.05.004","article-title":"Spontaneous brain oscillations and perceptual decision-making","volume":"24","author":"Samaha","year":"2020","journal-title":"Trends Cogn Sci"},{"key":"2026092213411070600_ref106","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1162\/imag_a_00018","article-title":"Overcoming harmonic hurdles: genuine beta-band rhythms vs. contributions of alpha-band waveform shape","volume":"1","author":"Schaworonkow","year":"2023","journal-title":"Imaging Neurosci"},{"key":"2026092213411070600_ref107","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1177\/23982128241255798","article-title":"Does theta synchronicity of sensory information enhance associative memory? Replicating the theta-induced memory effect","volume":"8","author":"Serin","year":"2024","journal-title":"Brain Neurosci Adv"},{"key":"2026092213411070600_ref108","doi-asserted-by":"publisher","first-page":"1412","DOI":"10.1016\/j.brs.2023.08.026","article-title":"Sub-harmonic entrainment of cortical gamma oscillations to deep brain stimulation in Parkinson\u2019s disease: model based predictions and validation in three human subjects","volume":"16","author":"Sermon","year":"2023","journal-title":"Brain Stimul"},{"key":"2026092213411070600_ref109","doi-asserted-by":"publisher","first-page":"488","DOI":"10.1016\/j.brs.2024.02.017","article-title":"Pre-existing oscillatory activity as a condition for sub-harmonic entrainment of finely tuned gamma in Parkinson\u2019s disease","volume":"17","author":"Sermon","year":"2024","journal-title":"Brain Stimul"},{"key":"2026092213411070600_ref110","doi-asserted-by":"publisher","first-page":"1553000","DOI":"10.3389\/fnbeh.2025.1553000","article-title":"Theta-gamma coupling: nonlinearity as a universal cross-frequency coupling mechanism","volume":"19","author":"Sheremet","year":"2025","journal-title":"Front Behav Neurosci"},{"key":"2026092213411070600_ref111","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1038\/s41398-023-02565-5","article-title":"Closing the loop between brain and electrical stimulation: towards precision neuromodulation treatments","volume":"13","author":"Soleimani","year":"2023","journal-title":"Transl Psychiatry"},{"key":"2026092213411070600_ref112","doi-asserted-by":"publisher","first-page":"570","DOI":"10.1038\/s41593-023-01270-2","article-title":"Forty-hertz light stimulation does not entrain native gamma oscillations in Alzheimer\u2019s disease model mice","volume":"26","author":"Soula","year":"2023","journal-title":"Nat Neurosci"},{"key":"2026092213411070600_ref113","doi-asserted-by":"publisher","first-page":"3536","DOI":"10.1523\/JNEUROSCI.4385-13.2014","article-title":"Local entrainment of alpha oscillations by visual stimuli causes cyclic modulation of perception","volume":"34","author":"Spaak","year":"2014","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref114","doi-asserted-by":"publisher","first-page":"2266","DOI":"10.1016\/j.clinph.2005.06.011","article-title":"Nonlinear dynamical analysis of EEG and MEG: review of an emerging field","volume":"116","author":"Stam","year":"2005","journal-title":"Clin Neurophysiol"},{"key":"2026092213411070600_ref115","doi-asserted-by":"publisher","first-page":"6445","DOI":"10.1523\/JNEUROSCI.1128-16.2016","article-title":"Gamma oscillations in the hyperkinetic state detected with chronic human brain recordings in Parkinson\u2019s disease","volume":"36","author":"Swann","year":"2016","journal-title":"J Neurosci"},{"key":"2026092213411070600_ref116","doi-asserted-by":"publisher","first-page":"R658","DOI":"10.1016\/j.cub.2012.06.061","article-title":"The functional importance of rhythmic activity in the brain","volume":"22","author":"Thut","year":"2012","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref117","doi-asserted-by":"publisher","first-page":"289","DOI":"10.3389\/fnins.2020.00289","article-title":"MVPA-light: a classification and regression toolbox for multi-dimensional data","volume":"14","author":"Treder","year":"2020","journal-title":"Front Neurosci"},{"key":"2026092213411070600_ref118","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1016\/j.tics.2024.12.003","article-title":"Processes and measurements: a framework for understanding neural oscillations in field potentials","volume":"29","author":"Van Bree","year":"2025","journal-title":"Trends Cogn Sci"},{"key":"2026092213411070600_ref119","doi-asserted-by":"publisher","first-page":"813","DOI":"10.3758\/s13423-020-01798-5","article-title":"The JASP guidelines for conducting and reporting a Bayesian analysis","volume":"28","author":"Van Doorn","year":"2021","journal-title":"Psychon Bull Rev"},{"key":"2026092213411070600_ref120","doi-asserted-by":"publisher","first-page":"995","DOI":"10.1016\/j.cub.2012.03.050","article-title":"Perceptual echoes at 10 Hz in the human brain","volume":"22","author":"VanRullen","year":"2012","journal-title":"Curr Biol"},{"key":"2026092213411070600_ref121","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/0013-4694(49)90164-9","article-title":"The central effects of rhythmic sensory stimulation","volume":"1","author":"Walter","year":"1949","journal-title":"Electroencephalogr Clin Neurophysiol"},{"key":"2026092213411070600_ref122","doi-asserted-by":"publisher","first-page":"1054","DOI":"10.1126\/science.1252900","article-title":"Targeted enhancement of cortical-hippocampal brain networks and associative memory","volume":"345","author":"Wang","year":"2014","journal-title":"Science"},{"key":"2026092213411070600_ref123","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1016\/j.tics.2003.10.012","article-title":"Synchronous neural oscillations and cognitive processes","volume":"7","author":"Ward","year":"2003","journal-title":"Trends Cogn Sci"},{"key":"2026092213411070600_ref124","doi-asserted-by":"publisher","first-page":"e49458","DOI":"10.7554\/eLife.49458","article-title":"Stimulating the hippocampal posterior-medial network enhances task-dependent connectivity and memory","volume":"8","author":"Warren","year":"2019","journal-title":"Elife"},{"key":"2026092213411070600_ref125","doi-asserted-by":"publisher","first-page":"e07886","DOI":"10.7554\/eLife.07886","article-title":"Phase-amplitude coupling supports phase coding in human ECoG","volume":"4","author":"Watrous","year":"2015","journal-title":"Elife"},{"key":"2026092213411070600_ref126","doi-asserted-by":"publisher","first-page":"1392","DOI":"10.1162\/jocn_a_01422","article-title":"Alpha frequency entrainment reduces the effect of visual distractors","volume":"31","author":"Wiesman","year":"2019","journal-title":"J Cogn Neurosci"},{"key":"2026092213411070600_ref127","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1016\/S0306-4522(00)00579-0","article-title":"Frequency-specific effects of flicker on recognition memory","volume":"104","author":"Williams","year":"2001","journal-title":"Neuroscience"},{"key":"2026092213411070600_ref128","doi-asserted-by":"publisher","first-page":"652393","DOI":"10.3389\/fnhum.2021.652393","article-title":"Improving the effect of transcranial alternating current stimulation (tACS): a systematic review","volume":"15","author":"Wu","year":"2021","journal-title":"Front Hum Neurosci"},{"key":"2026092213411070600_ref129","doi-asserted-by":"publisher","first-page":"3178","DOI":"10.1111\/ejn.15138","article-title":"Pre-stimulus alpha-band power and phase fluctuations originate from different neural sources and exert distinct impact on stimulus-evoked responses","volume":"55","author":"Zazio","year":"2021","journal-title":"Eur J Neurosci"},{"key":"2026092213411070600_ref130","doi-asserted-by":"publisher","first-page":"1718","DOI":"10.1007\/s12264-021-00781-x","article-title":"Precise modulation strategies for transcranial magnetic stimulation: advances and future directions","volume":"37","author":"Zhong","year":"2021","journal-title":"Neurosci Bull"},{"key":"2026092213411070600_ref131","doi-asserted-by":"publisher","first-page":"95","DOI":"10.3389\/fnins.2018.00095","article-title":"The involvement of endogenous neural oscillations in the processing of rhythmic input: more than a regular repetition of evoked neural responses","volume":"12","author":"Zoefel","year":"2018","journal-title":"Front Neurosci"}],"container-title":["Cerebral Cortex"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/cercor\/article-pdf\/36\/9\/bhag152\/71319355\/bhag152.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/cercor\/article-pdf\/36\/9\/bhag152\/71319355\/bhag152.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T17:41:18Z","timestamp":1790098878000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/cercor\/article\/doi\/10.1093\/cercor\/bhag152\/8828956"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,1]]},"references-count":131,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2026,9,1]]}},"URL":"https:\/\/doi.org\/10.1093\/cercor\/bhag152","relation":{},"ISSN":["1047-3211","1460-2199"],"issn-type":[{"value":"1047-3211","type":"print"},{"value":"1460-2199","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,9]]},"published":{"date-parts":[[2026,9,1]]},"article-number":"bhag152"}}