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The Journal of Neuroscience, April 13, 2005, 25(15):3962-3972; doi:10.1523/JNEUROSCI.4250-04.2005

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Behavioral/Systems/Cognitive
Phase Synchrony among Neuronal Oscillations in the Human Cortex

J. Matias Palva,¹ Satu Palva,^2,3 and Kai Kaila¹

¹Department of Biological and Environmental Sciences and ²Cognitive Brain Research Unit, Department of Psychology, University of Helsinki, FIN-00014 Helsinki, Finland, and ³BioMag Laboratory, Helsinki University Central Hospital, FIN-00029 Helsinki, Finland

Synchronization of neuronal activity, often associated with network oscillations, is thought to provide a means for integrating anatomically distributed processing in the brain. Neuronal processing, however, involves simultaneous oscillations in various frequency bands. The mechanisms involved in the integration of such spectrally distributed processing have remained enigmatic. We demonstrate, using magnetoencephalography, that robust cross-frequency phase synchrony is present in the human cortex among oscillations with frequencies from 3 to 80 Hz. Continuous mental arithmetic tasks demanding the retention and summation of items in the working memory enhanced the cross-frequency phase synchrony among (10 Hz), (20 Hz), and (30-40 Hz) oscillations. These tasks also enhanced the "classical" within-frequency synchrony in these frequency bands, but the spatial patterns of , , and synchronies were distinct and, furthermore, separate from the patterns of cross-frequency phase synchrony. Interestingly, an increase in task load resulted in an enhancement of phase synchrony that was most prominent between - and -band oscillations. These data indicate that cross-frequency phase synchrony is a salient characteristic of ongoing activity in the human cortex and that it is modulated by cognitive task demands. The enhancement of cross-frequency phase synchrony among functionally and spatially distinct networks during mental arithmetic tasks posits it as a candidate mechanism for the integration of spectrally distributed processing.

Key words: ; attention; cortex; ; oscillator; parietal; synchrony; working memory

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Received Oct 13, 2004; revised March 4, 2005; accepted March 4, 2005.

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