Sleep-Wake Related Discharge Properties of Basal Forebrain Neurons Recorded With Micropipettes in Head-Fixed Rats

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Abstract
The basal forebrain has been shown to play an important role in cortical activation of wake and paradoxical sleep (PS), yet has also been posited to play a role in slow wave sleep (SWS). In an effort to determine whether these different roles may be fulfilled by different cell groups, including cholinergic and GABAergic cells, we recorded from 123 units in waking-sleeping, head-fixed rats using micropipettes to allow juxtacellular labeling. Functional sets of intermingled cell groups emerged as units whose discharge was as follows: 1) maximum in active wake (aW) and positively or not correlated with EEG gamma activity, while positively correlated with nuchal EMG activity, and thus potentially facilitatory for waking and behavioral arousal (12%); 2) maximum in SWS or SWS-PS and positively correlated with delta EEG activity, while not or negatively correlated with EMG activity, and thus potentially promotive for sleep with cortical slow wave activity and/or accompanying behavioral changes (16%); 3) maximum in PS or PS and aW and positively correlated with gamma and theta EEG activity, while negatively or not correlated with EMG activity, and thus potentially promotive for cortical activation during PS or PS and W (62%); and 4) equivalent across all states and thus not involved in state regulation (∼10%). Units of each group also manifested different firing patterns typified as slow tonic (19.5%), fast tonic (32.5%), or fast phasic (48%), including rhythmic bursting (6%). Through these diverse cell groups, the basal forebrain has the capacity to modulate cortical activity, behavior, and/or related physiological processes across the sleep-waking cycle and thereby regulate the sleep-wake state of the animal.