Rosa M. Salas’s research while affiliated with University of Seville and other places

Evidence suggests that sleep homeostasis is not only dependent on duration of previous wakefulness but also on experience- and/or use-dependent processes. Such homeostatic mechanisms are reflected by selective increases in the duration of a sleep stage, modifications to electrophysiological-metabolic brain patterns in specific sleep states, and/or reactivation to neuronal ensembles in subsequent sleep periods. Use-dependent sleep changes, apparently different from those changes caused by memory consolidation processes, are thought to reflect neuronal restoration processes after the sustained exposure to stimulation during the preceding wakefulness. In the present study, we investigated changes in the brain electrical activity pattern during human sleep after 6 hr of continuous auditory stimulation during previous wakefulness. Poststimulation nights showed a widespread increase of spectral power within the alpha (8-12 Hz) and sleep spindle (12-15 Hz) frequency range during slow-wave sleep (SWS) compared with the baseline night. This effect was mainly attributable to an enhanced EEG amplitude rather than an increase of oscillations, except for temporal (within alpha and sleep spindles) and parietal regions (within sleep spindles) in which both parameters contributed equally to the increase of spectral energy. Power increments were accompanied by a strengthening of the coherence between fronto-temporal cortical regions within a broad frequency range during SWS but to the detriment of the coherence between temporal and parieto-occipital areas, suggesting underlying compensatory mechanisms between temporal and other cortical regions. In both cases, coherence was built up progressively across the night, although no changes were observed within each SWS period. No electrophysiological changes were found in rapid eye movement sleep. These results point to SWS as a critical brain period for correcting the cortical synaptic imbalance produced by the predominant use of specific neuronal populations during the preceding wakefulness, as well as for synaptic reorganization after prolonged exposure to a novel sensory experience.

Evidence suggests that sleep architecture is affected by endogenous homeostatic mechanisms as well as by behavioral and sensory demands during the prior wakefulness. Regarding the auditory system, sensory deprivation has shown to drastically modify the sleep structure, stressing the relevance of such sensory system for sleep organization. Changes in sleep architecture following prolonged auditory stimulation during prior wakefulness would provide additional support to this hypothesis. In the present study, auditory stimulation was administered over a 6 h period prior to sleep. Sleep parameters obtained from visual scoring were quantified across the total sleep period, for each sleep cycle, and for the two halves of the night, separately. Results showed that 6 h of waking-auditory stimulation were followed by an increase in the duration of slow wave sleep, a shortening of the latency between slow wave sleep periods, and a longer sleep onset latency as compared with the baseline night. In contrast, REM sleep parameters were unaffected by the pre-sleep auditory stimulation. These results indicate that sleep architecture depends on auditory demands during the prior wakefulness, suggesting that the local neural activation underlying auditory stimulation may trigger brain control mechanisms selectively involved in both the slow wave sleep maintenance and organization.

Cortical oscillations in the range of alpha activity (8-13 Hz) are one of the fundamental electrophysiological phenomena of the human electroencephalogram (EEG). Evidence from quantitative EEG data has shown that their electrophysiological features, cortical generation mechanisms, and therefore, their functional correlates vary along the sleep-wake continuum. Specifically, spectral microstructure and EEG coherence levels between anterior and posterior cortical regions permit to differentiate among alpha activity spontaneously appearing in relaxed wakefulness with eyes closed, drowsiness period, and REM sleep, by reflecting distinct properties of neural networks involved in its cortical generation as well as a different interplay between cortical generators, respectively. Besides, the dissimilar spatiotemporal features of brain electrical microstates within the alpha range reveals a different geometry of active neural structures underlying each alpha variant or, simply, changes in the stability level of neural networks during each brain state. Studies reviewed in this paper support the hypothesis that two different alpha variants occur during human REM sleep: 'background responsive alpha activity', blocked over occipital regions when rapid eye movements are present, and 'REM-alpha bursts', non modulated by the alteration of tonic and phasic periods. Altogether, evidence suggests that electrophysiological features of human cortical oscillations in the alpha frequency range vary across different behavioural states, as well as within state, reflecting different cerebral phenomena with probably dissimilar functional meaning.

El estudio de los ritmos cerebrales puede ser abordado mediante el análisis de sus propiedades electrofisiológicas usando técnicas de EEG cuantitativo. En aquellos casos donde una misma actividad aparece espontáneamente en diferentes estados de activación cerebral, el estudio de estas propiedades ayudaría a establecer diferencias funcionales asociadas a cada uno de estos estados. El presente trabajo revisa aquellos estudios que han determinado las propiedades electrofisiológicas de las diferentes variantes normales de alfa que aparecen en el continuo vigilia-sueño, más concretamente en vigilia relajada, somnolencia y durante la fase REM del sueño. Los resultados indican que cada variante normal de alfa, aun mostrando una distribución topográfica similar en cada uno de los estados cerebrales, presenta características diferenciales atendiendo a su composición espectral, relaciones funcionales entre regiones corticales, y micro-estados cerebrales subyacentes. De acuerdo con estos hallazgos, cada variante de alfa desempeñaría diferentes funciones cerebrales. El ritmo alfa de vigilia parece ser el resultado de una máxima sincronización neuronal como consecuencia de la ausencia de procesamiento sensorial, mientras que la presencia de actividad alfa durante la somnolencia estaría más asociada al procesamiento de las imágenes hipnagógicas que ocurren al comienzo del sueño. Los brotes de alfa de REM, sin embargo, constituirían un punto de contacto entre el cerebro dormido y el ambiente externo. Esta caracterización electrofisiológica encuentra su campo de aplicación más directo en el diseño de algoritmos para clasificar el sueño de forma automática, así como en el diagnóstico y evaluación de determinadas patologías donde pudieran verse afectados los mecanismos cerebrales de generación de esta actividad a lo largo del continuo vigilia-sueño.

Normal electroencephalographic (EEG) alpha variants appear during relaxed wakefulness with closed eyes, drowsiness period at sleep onset, and rapid eye movement (REM) sleep in bursts without arousal signals. Previous results revealed that fronto-occipital and fronto-frontal alpha coherences became weaker from wakefulness to drowsiness, and finally to REM sleep. The present work was aimed at determining whether a generalized or a unidirectional deactivation of the long fronto-occipital fasciculi, previously proposed to be involved in the alpha rhythm generation, could explain the above-mentioned results. Polynomial regression analyses, applied to the change of alpha coherence with distance along the antero-posterior axis, suggested that the anterior and posterior local circuits show a similar level of activation in all brain states. Bivariate partial correlation analyses between local alpha coherences revealed that such local circuits maintain a reciprocal dependency during wakefulness, but unidirectional during drowsiness (anterior-to-posterior, A-P) and REM sleep (posterior-to-anterior, P-A). From these findings, both anterior and posterior cortical structures are suggested as being involved in the generation of the three alpha variants. If the implication of a double cortical generation source (anterior and posterior) of alpha variants is assumed, these two generators seem to maintain a mutual inter-dependency during wakefulness, whereas during the transition to human sleep, the anterior areas work quite independently of the posterior regions. Finally, the occipital structures may be the driving force for the REM-alpha bursts generation, since involvement of frontal regions demonstrated a high dependence on the posterior neural circuits in the genesis of this sleep event.

Evidence suggests that an important contribution of spectral power in the alpha range is characteristic of human REM sleep. This contribution is, in part, due to the appearance of well-defined bursts of alpha activity not associated with arousals during both tonic and phasic REM fragments. The present study aims at determining if the REM-alpha bursts constitute a different alpha variant from the REM background alpha activity. Since previous findings showed a selective suppression of background alpha activity over occipital regions during phasic REM fragments and, on the other hand, the density of alpha bursts seem to be independent of the presence or absence of rapid eye movements, one expects to find the same spectral power contribution of alpha bursts in tonic and phasic REM fragments. The results indicated that REM-alpha bursts showed a similar power contribution and topographic distribution (maximum energy over occipital regions) both in tonic and phasic REM fragments. This suggests that two variants of alpha activity with different functional roles are present during the human REM sleep: i) background alpha activity, modulated over occipital regions by the presence of rapid eye movements, which may be an electrophysiological correlate of the visual dream contents; and ii) REM-alpha bursts, independent of the presence of rapid eye movements, which could be facilitating the connection between the dreaming brain and the external world, working as a micro-arousal in this brain state.

The functional relationships between the brain areas supposedly involved in the generation of the alpha activity were quantified by means of INTRA- and INTER-hemispheric coherences during different arousal states (relaxed wakefulness, drowsiness at sleep onset, and rapid eye movement sleep) where such an activity can be clearly detectable in the human EEG. A significant decrease in the fronto-occipital as well as in the inter-frontal coherence values in the alpha range was observed with the falling of the vigilance level, which suggests that the brain mechanisms underlying these coherences are state dependent. Making fronto-frontal coherence values in the alpha frequency band useful indexes to discern between brain functional states characterized by a different arousal level.

Alpha activity attenuation (blocking) over occipital regions is an electrophysiological index of cortical activation associated with visual attention and waking mental imagery. The present work focused on exploring whether the human REM background alpha activity was modulated, attending to tonic- (without rapid oculomotor activity) and phasic-REM periods (with a prominent burst of REMs). The obtained results revealed that the background alpha activity showed a decreased spectral power over occipital brain regions during phasic-REM in comparison with tonic-REM periods. This result suggests an active visual processing caused by the complex mental imagery generated during periods of oculomotor activity in human REM sleep.