High-frequency gamma electroencephalogram activity in association with sleep-wake states and spontaneous behaviors in the rat.

The occurrence of high-frequency gamma activity (30-60 Hz) and its relationship to other frequency band activities were examined by spectral analysis of the electroencephalogram in association with sleep wake states and spontaneous behaviors in the rat. In the electroencephalogram, gamma wave activity was evident in unfiltered and high-frequency filtered recordings, in which it was prominent during attentive or active Wake episodes and during Paradoxical Sleep, when theta-like activity was also apparent. In amplitude spectra from these episodes, multiple peaks were evident within the gamma frequency band, indicating broad-band high-frequency activity, in association with a single low-frequency peak in the theta band. gamma peaks were attenuated during quiet Waking, in association with a low-frequency peak between theta and delta, and during Slow Wave Sleep, in association with a low-frequency peak in the delta band. In coherence spectra from ipsilateral cortical leads, peaks were also present within the gamma range and were significantly higher in Waking moving and Paradoxical Sleep than in Waking quiet and Slow Wave Sleep. In measures of frequency band amplitude, gamma activity (30.5-58.0 Hz) varied significantly across the sleep waking cycle, being similarly high during Wake and Paradoxical Sleep and lowest during Slow Wave Sleep. Across these states, gamma was negatively correlated with delta (1.5-4.0 Hz). In contrast, high beta (19.0-30.0 Hz) was significantly lower in Wake than in Slow Wave Sleep and was positively correlated with delta. gamma differed significantly across specific behaviors, being highest in Paradoxical Sleep with twitches and during Waking eating and moving behaviors, slightly lower in Waking attentive, lower in Waking grooming and as low in Waking quiet as during Slow Wave Sleep. These results indicate that the reciprocal variation of high-frequency gamma activity (and not beta) with low-frequency delta activity reflects the sleep waking cycle of the rat. Moreover, gamma activity reflects the degree of behavioral arousal, since it is high during active Waking, when the electromyogram is high, and low during quiet Waking, when the electromyogram is low. It also reflects cortical arousal, independent of motor activity, since it attains high levels in association with attentive immobility and maximal levels only during particular active behaviors (eating and moving and not grooming), and it also attains maximal levels during Paradoxical Sleep, when the nuchal electromyogram is minimal, but small twitches evidence dreaming. The co-variation of gamma and a slow oscillation in the theta band across states and behaviors suggests that a common system may modulate these fast and slow electroencephalogram rhythms, and that such modulation, potentially emanating from the basal forebrain, could predominate during certain states or behaviors, such as Paradoxical Sleep.