Neural oscillations subserve many human perceptual and cognitive operations. Accordingly, brain functional connectivity is not static in time, but fluctuates dynamically following the synchronization and desynchronization of neural populations. This dynamic functional connectivity has recently been demonstrated in spontaneous fluctuations of the Blood Oxygen Level-Dependent (BOLD) signal, measured with functional Magnetic Resonance Imaging (fMRI). We analyzed temporal fluctuations in BOLD connectivity and their electrophysiological correlates, by means of long (≈50 min) joint electroencephalographic (EEG) and fMRI recordings obtained from two populations: 15 awake subjects and 13 subjects undergoing vigilance transitions. We identified positive and negative correlations between EEG spectral power (extracted from electrodes covering different scalp regions) and fMRI BOLD connectivity in a network of 90 cortical and subcortical regions (with millimeter spatial resolution). In particular, increased alpha (8–12 Hz) and beta (15–30 Hz) power were related to decreased functional connectivity, whereas gamma (30–60 Hz) power correlated positively with BOLD connectivity between specific brain regions. These patterns were altered for subjects undergoing vigilance changes, with slower oscillations being correlated with functional connectivity increases. Dynamic BOLD functional connectivity was reflected in the fluctuations of graph theoretical indices of network structure, with changes in frontal and central alpha power correlating with average path length. Our results strongly suggest that fluctuations of BOLD functional connectivity have a neurophysiological origin. Positive correlations with gamma can be interpreted as facilitating increased BOLD connectivity needed to integrate brain regions for cognitive performance. Negative correlations with alpha suggest a temporary functional weakening of local and long-range connectivity, associated with an idling state.

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The integration of segregated brain functional modules is a prerequisite for conscious awareness during wakeful rest. Here, we test the hypothesis that temporal integration, measured as longterm memory in the history of neural activity, is another important quality underlying conscious awareness. For this aim, we study the temporal memory of blood oxygen level-dependent signals across the human nonrapid eye movement sleep cycle. Results reveal that this property gradually decreases from wakefulness to deep nonrapid eye movement sleep and that such decreases affect areas identified with default mode and attention networks. Although blood oxygen level-dependent spontaneous fluctuations exhibit nontrivial spatial organization, even during deep sleep, they also display a decreased temporal complexity in specific brain regions. Conversely, this result suggests that long-range temporal dependence might be an attribute of the spontaneous conscious mentation performed during wakeful rest.T he human brain displays complex spatiotemporal patterns of energy-consuming activity, even in the absence of an explicit task or stimulation (1). Large efforts have been devoted to the study of spontaneous neural activity encoded in the slow (∼0.1 Hz) fluctuations of the blood oxygen level-dependent (BOLD) signal, which are measured with functional MRI (fMRI). Nontrivial spatial organization of functional brain activity in resting state networks (RSNs) was consistently shown (2-4), comprising brain regions with high BOLD signal coherence and anatomical consistency with systems activated during task performance or stimulation (5).Remarkably, although human nonrapid eye movement (NREM) sleep is characterized by impaired awareness and reduced conscious mentation, organization into RSNs is preserved in light sleep (6) and to a large extent, deeper sleep stages (7, 8) (SI Appendix, Fig. S8.1). In particular, the default mode network (DMN; a set of task-deactivated regions implied with internal conscious cognitive processes) (9, 10) was repeatedly observed during deep sleep, albeit with reduced frontal connectivity (11,12). Although brain modules are preserved, even in the absence of conscious awareness, their functional integration is greatly impaired (8,13,14), which was predicted by an information integration account of consciousness (15). These results suggest that ongoing conscious mentation is not the only origin of RSN activity, whereas the level of consciousness is reflected in the interaction of functional networks.However, brain activity is not completely characterized in the spatial domain only. fMRI BOLD signals display rich temporal organization, including scale-free 1/f power spectra and long-range temporal autocorrelations (16)(17)(18), with activity at any given time being influenced by the previous history of the system up to several minutes into the past. These landmarks of complex information processing and rapid adaptability are shared by many systems found in nature (19,20). Evidence for such properties is also manifest...

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