Section of Psychopharmacology and Sleep Research
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University of Zürich - Institute of Pharmacology and Toxicology Section of Psychopharmacology and Sleep Research |
EEG slow-wave activity (SWA; spectral power in the 0.75-4.5 Hz band) is a function of the duration of prior waking and, thereby, an indicator of sleep homeostasis. We present a model that accounts for both the declining trend of SWA during sleep and for its variation within the successive nonrapid eye movement (non-REM) sleep episodes. The values of the model parameters were estimated by an optimization procedure in which empirical SWA of baseline nights (16 subjects, 26 nights) served as a reference. A sensitivity analysis revealed the model to be quite robust to small changes (+/- 5%) of the parameter values. The estimated parameter values were used to simulate data sets from three different experimental protocols (sleep in the evening or sleep in the morning after prolonged waking, or extended sleep initiated at the habitual bedtime; n = 8 or 9). The timing of the REM trigger parameter was derived from the empirical data. A close fit was obtained between the simulated and empirical SWA data, and even the occasional late SWA peaks during extended sleep could be reproduced. Minor discrepancies suggest indirect or direct circadian influences on SWA. The simulations demonstrate that the concept of sleep homeostasis as proposed in the two-process model of sleep regulation can be refined to account in quantitative terms for empirical data and to predict the changes induced by the prolongation of waking or sleep.
Brain Res. Bull. 31 (1993): 97-113.
The dynamics of the sleep EEG was investigated by all-night spectral analysis of 51 sleep records. Power density was calculated for 1-Hz bins in the 0.25-25.0 Hz range. Values in non-rapid-eye-movement sleep (NREMS) were higher than in REMS in the 0.25-16.0 Hz range, and lower in the 18.25-22.0 Hz range. Power density in the 0.25-12.0 Hz range showed a declining trend over the first four NREMS episodes, which, depending on the frequency bin, could be approximated by non-linear or linear decay functions. In the frequency range of sleep spindles (12.25-15.0 Hz), power density in the 13.25-15.0 Hz band showed an increasing trend between NREMS episode 2 and NREMS episode 4. A correlation matrix of 25 1-Hz bins revealed for NREMS a negative correlation between slow- wave activity (SWA; 0.25-4.0 Hz) and activity in the spindle frequency range. This negative correlation was highest in the first NREMS episode and diminished progressively over the subsequent NREMS episodes. Within NREMS episodes, the values in the spindle frequency range showed a U-shaped time course, the trough coinciding with a high level of SWA. By contrast, in both the early and late part of the episode the two types of activity changed in the same direction. The results are consistent with recent electrophysiological studies indicating that the establishment of NREMS is associated with a progressive hyperpolarization of thalamocortical neurons during which the membrane potential exhibits oscillations first in the spindle frequency range and then in the range of SWA.
J. Sleep Res. 2 (1993): 70-81.
The effect of repeated partial sleep deprivation on sleep stages and electroencephalogram (EEG) power spectra during sleep and wakefulness was investigated in nine healthy young subjects. Three baseline nights of 8 hours (2300-0700 hours) were followed by four nights with 4 hours of sleep (2300-0300 hours) and three recovery nights of 8 hours (2300-0700 hours). Sleep restriction curtailed sleep stages 1 and 2 as well as rapid eye movement (REM) sleep, but left slow wave sleep largely unaffected. In the first two recovery nights, total sleep time and REM sleep were enhanced, and sleep latency was shortened. Slow wave sleep was increased only in the first recovery night. In accordance with the prediction of the two-process model of sleep regulation, slow wave activity (SWA; spectral power density in the 0.75-4.5-Hz range) in nonrapid eye movement (NREM) sleep increased by approximately 20% in the first night following sleep restriction, remained at this level in the subsequent 3 nights and decreased immediately after the first recovery night. In contrast to these immediate changes, progressive and more persistent changes were seen in the EEG activity of higher frequencies. Thus, activity in the upper delta band tended to gradually increase from night to night during the sleep restriction period, whereas after an initial increase, activity in the theta-alpha band changed in the opposite direction. The progressive changes were also present in the EEG spectra of REM sleep and wakefulness. Because the time course of these changes paralleled the cumulative deficit in REM sleep, they may represent a correlate of REM sleep pressure.
Sleep 16 (1993): 100-113.
The predictions of the two-process model of sleep regulation were tested by 12-h sleep deprivation (SDEP) and 12-h cold exposure (cold, 4 degrees C) in the rat, both carried out in the 12-h dark period. The analysis was based on recordings of vigilance states, electroencephalogram (EEG) power spectra (0.25- 25.0 Hz), and cortical temperature (Tcrt). During recovery from SDEP, slow-wave activity (SWA; EEG power density in the 0.75-4.0 Hz band) in non-REM sleep was higher and the number of brief awakenings (nBA) lower than in the corresponding baseline period. During recovery from cold, a moderate and delayed increase in SWA was present. During SDEP, Tcrt was above baseline, and in some intervals of recovery below baseline. In those intervals waking was also decreased. During recovery from cold, Tcrt and waking were not affected. It is concluded that 12-h SDEP causes an intensification of sleep, as indicated by the enhanced SWA and the reduced nBA, whereas 12-h cold has only marginal effects. The time course of SWA in both experiments could be accurately predicted by a computer simulation based on the assumption of the two-process model and the parameter values determined in a previous experiment.
Physiol. Behav. 54 (1993): 885-894.
Both period-amplitude analysis (PAA) and power spectral analysis (PSA) were performed on all-night human sleep EEG recordings obtained from 11 subjects. The comparison of the two methods was based on the PAA variables time in band (a wave incidence measure) and rectified amplitude, and on the PSA variables spectral power density and spectral amplitude (the square root of power). The mean time course of these variables was determined for the first 4 nonREM-REM sleep cycles. Spectral power density and spectral amplitude in the delta range were high in nonREM sleep and low in REM sleep, and showed a declining trend over consecutive nonREM sleep episodes. In the frequency range below 2 Hz, rectified amplitude was highly correlated with both time in band and spectral amplitude, and there was no evidence for a dissociation between wave amplitude and wave incidence measures. However, in frequencies above 2 Hz, the modulation of time in band was a mirror image of that below 2 Hz. This result does not reflect a property of the data, but is inherent to the methodology applied. The reversal point of modulation was merely shifted when the high-pass filter settings were changed. It is concluded that band-pass filtering is necessary prior to PAA even for the analysis of the lowest frequency range, and that the indiscriminate use of PAA may give rise to spurious results.
J. Sleep Res. 2 (1993): 121-129.
In the guinea pig, sleep and slow-wave activity (SWA) are evenly distributed over 24 h in contrast to the sleep pattern in other rodents where a daily preference for sleep and a decline of SWA within the main sleep period is typical. SWA is regulated as a function of prior waking and is assumed to reflect sleep intensity. We investigated sleep homeostasis and its possible circadian modulation by performing sleep deprivation (SD) at two phases of the light-dark cycle. SD induced an increase in SWA which was similar under both conditions. We conclude that sleep is homeostatically regulated in the guinea pig and is not subject to a circadian modulation.
Neurosci. Lett. 164 (1993): 105-108.
Vigilance states, electroencephalogram (EEG) power spectra (0.25-25.0 Hz), and cortical temperature (TCRT) were obtained in nine guinea pigs for 24 h in a 12:12-h light-dark (LD 12:12) schedule. Sleep was markedly polyphasic and fragmented and amounted to 32% of recording time, which is a low value compared with sleep in other rodents. There was 6.8% more sleep in the light period than in the dark period. EEG power density in non-rapid eye movement (NREM) sleep showed no significant temporal trend within the light or the dark period. The homeostatic aspects of sleep regulation, as proposed in the two-process model, can account for the slow-wave activity (SWA) pattern also in the guinea pig: The small 24-h amplitude of the sleep-wakefulness pattern resulted in a small, 12% decline of SWA within the light period. In contrast to more distinctly nocturnal rodents, SWA in the dark period was not higher than in the light period. TCRT showed no difference between the light and the dark period. TCRT in REM sleep and waking was higher than TCRT in NREM sleep. TCRT increased after the transition from NREM sleep to either REM sleep or waking, and decreased in the last minute before the transition and after the transition from waking to NREM sleep. Motor activity measured in six animals for 11 days in constant darkness showed no apparent rhythm in three animals and a significant circadian rhythm in three others. Our data support the notion that guinea pigs exhibit only a weak circadian rest-activity rhythm.
Am. J. Physiol. 264 (1993): R1125-R1132.