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S Daan

Publications and source records attributed to S Daan.

At least 55 records · Page 3Linked to original sources

A single pacemaker can produce different rates of reentrainment in different overt rhythms.

Using a simple Van der Pol oscillator, we show that periodic events triggered by different states of the oscillator can reentrain at different rates following a phase shift of the Zeitgeber. Such differences can emerge due to waveform distortion during reentrainment, due to masking and due to fitting procedures (e.g. acrophase determination). Therefore, different rates of reentrainment are consistent with single pacemaker models of circadian systems. Likewise, "Fractional Desynchronization" can yield different "ranges of entrainment" in different overt rhythms generated by the same pacemaker.

Journal Article↗

Generation of activity-rest patterns by dual circadian pacemaker systems: a model.

Activity-rest patterns displayed by an animal under various circumstances are suggested to result from the combined influences of two virtually identical circadian pacemaker components. Increased output of each component proportionally increases the probability of activity of the animal. Such a dual circadian pacemaker model explains much of the phenomenology of activity-rest records. Under normal light-dark schedules, the two components of the model have a similar phase relationship to the Zeitgeber. Differential synchronization of the components to dawn and dusk is not assumed.

Journal Article↗

Warming up for sleep? Ground squirrels sleep during arousals from hibernation.

Hypothermia during mammalian hibernation is periodically interrupted by arousals to euthermy, the function of which is unknown. We report that arctic ground squirrels (Spermophilus parryii) consistently sleep during these arousals, and that their EEG shows the decrease in slow wave activity (delta power) that is characteristic of a declining requirement for sleep. These results are consistent with the novel hypothesis that the need for sleep slowly accumulates during torpor, and that returning to euthermy is periodically required to allow sleep. Sleep thus seems to be energetically expensive for a hibernating mammal, and cannot be considered solely a strategy for saving energy.

Animals↗

Effect of clomipramine on sleep and EEG power spectra in the diurnal rodent Eutamias sibiricus.

Sleep was recorded in the diurnal rodent Eutamias sibiricus, chronically implanted with EEG and EMG electrodes. The tricyclic antidepressant drug clomipramine suppressed the duration of REM sleep and EEG power density in the frequencies between 1.5 and 13.5 Hz in nonREM sleep. During the administration of clomipramine, 24 h of sleep deprivation by forced locomotion significantly reduced the duration of waking and increased the duration of nonREM sleep. During the first 2 h of recovery sleep, EEG power density of the frequencies between 2.5 and 6 Hz was enhanced. These effects of sleep deprivation were to a large extent similar to those of sleep deprivation under drug free conditions. It is concluded that clomipramine affects the EEG in nonREM sleep but does not interfere substantially with nonREM sleep regulatory processes, which are activated by sleep deprivation.

Animals↗

Avian basal metabolic rates: their association with body composition and energy expenditure in nature.

Measurements of basal metabolic rate (BMR), body water, fat, and lean dry mass of different organs were obtained in 22 bird species, ranging from 10.8 to 1,253 g body mass. Residuals of BMR (after subtracting BMR allometrically predicted from body mass) were positively correlated with residuals of lean dry heart and kidney mass. Measurements of both BMR and the daily energy expenditure of parent birds (DEEpar) during the period of nestling care as assessed by labeled-water turnover were collected from the literature for 26 altricial bird species. The allometric relationships with body mass in this data set were: log BMR (W) = -1.385 + 0.684 log mass (g) [fraction of variance (r2) = 0.973] and log DEEpar (W) = -0.797 + 0.659 log mass (g) (r2 = 0.967). Residuals of log BMR and log DEEpar were positively correlated with each other. The parallel regressions and correlation of residuals lead to reduced variance in the ratio of BMR/DEEpar (mean 0.301; SD 0.086). We suggest that natural selection has led to an adjustment of the size of organs (such as heart and kidney) involved in sustaining energy metabolism at the DEE maximized during parental care and that size-independent variation in BMR reflects the relative size of this highly metabolically active machinery. These relationships of BMR lead to new interpretations of the decline in mass-specific BMR with increasing body size and decreasing latitude and of the difference in mass-specific BMR between birds and mammals.

Animals↗

The two-oscillator circadian system of tree shrews (Tupaia belangeri) and its response to light and dark pulses.

The wheel-running activity rhythm of tree shrews (tupaias; Tupaia belangeri) housed in constant darkness (DD) phase-advanced following a 3-hr light pulse at circadian time (CT) 21. Dark pulses of 3 hr presented to tupaias in bright constant light (LL) did not induce significant phase shifts of the free-running activity rhythm, irrespective of the CT. In dim LL, tupaias showed simultaneous splitting of their circadian rhythm of wheel-running activity, nest-box activity, and feeding behavior. Light pulses of 6 hr and 2300 lux were presented to 13 tupaias with split wheel-running activity rhythms. These light pulses induced immediate phase shifts in the two components of the split rhythm in opposite directions. No differences were observed between the light-pulse phase response curves of the two components. Equally large immediate phase advances were induced in both components by light pulses of 230 lux, but not by 23 lux. The final phase shifts were small at all CTs. In two tupaias, activity rhythms transiently split and re-fused. Analysis of the relative position of the components in one of these indicates asymmetry in the coupling between the components.

Animals↗

Differential elimination of circadian and ultradian rhythmicity by hypothalamic lesions in the common vole, Microtus arvalis.

Effects of hypothalamic lesions on the ultradian and circadian organization of wheel running and feeding were studied in the common vole, Microtus arvalis. Circadian organization broke down within 30 days in continuous darkness in 24% of intact voles (n = 135). Ultradian rhythmicity of feeding (period 2-3 hr) persisted in constant conditions in all intact voles. Following lesions of the suprachiasmatic nuclei (SCN), circadian rhythmicity disappeared when lesions were complete (n = 8) or more extensive than 25% of the total SCN volume (n = 5). Absence of circadian rhythmicity was also found in animals with substantial lesions in the diencephalic paraventricular area (PVA) and in the retrochiasmatic area (RCA) and/or adjacent arcuate nucleus (Arc). Complete loss of ultradian and circadian organization occurred in eight voles with damage to the RCA and/or Arc. In three of these, the SCN was intact. The SCN is a likely candidate for a circadian pacemaker in voles (as in other rodents), while the loss of circadian rhythmicity following PVA and RCA/Arc lesions may be due to destruction of efferent pathways from the SCN. The RCA/Arc area is apparently necessary for the expression of ultradian rhythms. The intact SCN is neither necessary nor sufficient for the generation of ultradian rhythmicity.

Activity Cycles↗

Effects of seganserin, a 5-HT2 antagonist, and temazepam on human sleep stages and EEG power spectra.

The effects of seganserin, a specific 5HT2 antagonist, on human sleep were assessed in two experiments and compared to the effects of temazepam and sleep deprivation. During daytime recovery sleep after sleep deprivation, seganserin did not significantly enhance visually scored slow wave sleep (SWS, stages 3 + 4) or the EEG power density in the delta frequencies. Under these conditions temazepam reduced the power density in the delta and theta frequencies. During nighttime sleep after a nap in the evening, seganserin caused an increase in SWS, a reduction in intermittent wakefulness, and an enhancement of the power density in the delta and theta frequencies during non-rapid eye movement (NREM) sleep. Temazepam induced a reduction in the power density in the delta and theta frequencies. It is concluded that the 5HT2 antagonist, seganserin, can induce SWS. However, since the spectral results showed that the changes in the sleep EEG were not identical to those induced by sleep deprivation it seems premature to conclude that 5HT2 receptors are primarily involved in NREM sleep regulation.

Adult↗

Sleep EEG spectral analysis in a diurnal rodent: Eutamias sibiricus.

1. Sleep was studied in the diurnal rodent Eutamias sibiricus, chronically implanted with EEG and EMG electrodes. Analysis of the distribution of wakefulness, nonrapid eye movement (NREM) sleep, and rapid eye movement (REM) sleep over the 24 h period (LD 12:12) showed that total sleep time was 27.5% of recording time during the 12 h light period and 74.4% during the 12 h dark period. Spectral analysis of the sleep EEG revealed a progressive decay in delta power density in NREM sleep during darkness. Power density of the higher frequencies increased at the end of darkness. Power density of the higher frequencies decreased and that of the lower frequencies increased during light. 2. Analysis of the distribution of vigilance states under three different photoperiods (LD 18:6; 12:12; 6:18) revealed that changes in daylength mainly resulted in a redistribution of sleep and wakefulness over light and darkness. Under long days the percentage of sleep during light was enhanced. The time course of delta power density in NREM sleep was characterized by a long rising part and a short falling part under long days, while a reversed picture emerged under short days. As a consequence, the power density during days. As a consequence, the power density during light was relatively high under long days. 3. After 24 h sleep deprivation by forced activity, no significant changes in the percentages of wakefulness and NREM were observed, whereas REM sleep was slightly enhanced. EEG power density, however, was significantly increased by ca. 50% in the 1.25-10.0 Hz range in the first 3 h of recovery sleep. This increase gradually decayed over the recovery night. 4. The same 24 h sleep deprivation technique led to a ca. 25% increase in oxygen consumption during recovery nights. While the results of the EEG spectral analysis are compatible with the hypothesis that delta power density reflects the 'intensity' of NREM sleep as enhanced by prior wakefulness and reduced by prior sleep, such enhanced sleep depth after sleep deprivation is not associated with reduced energy expenditure as might be anticipated by some energy conservation hypotheses on sleep function.

Animals↗

Bright morning light advances the human circadian system without affecting NREM sleep homeostasis.

Eight male subjects were exposed to either bright light or dim light between 0600 and 0900 h for 3 consecutive days each. Relative to the dim light condition, the bright light treatment advanced the evening rise in plasma melatonin and the time of sleep termination (sleep onset was held constant) for an average approximately 1 h. The magnitude of the advance of the plasma melatonin rise was dependent on its phase in dim light. The reduction in sleep duration was at the expense of rapid-eye-movement (REM) sleep. Spectral analysis of the sleep electroencephalogram (EEG) revealed that the advance of the circadian pacemaker did not affect EEG power densities between 0.25 and 15.0 Hz during either non-REM or REM sleep. The data show that shifting the human circadian pacemaker by 1 h does not affect non-REM sleep homeostasis. These findings are in accordance with the predictions of the two-process model of sleep regulation.

Adult↗

Intraspecific allometry of basal metabolic rate: relations with body size, temperature, composition, and circadian phase in the kestrel, Falco tinnunculus.

The relationship between body size and basal metabolic rate (BMR) in homeotherms has been treated in the literature primarily by comparison between species of mammals or birds. This paper focuses on the intraindividual changes in BMR when body mass (W) varies with different maintenance regimens. BMR varied in individual kestrels in proportion to W1.67, which is considerably steeper than the mass exponents for homomorphic change (0.667; Heusner, 1984) for interspecific comparison among all birds (0.677) or raptors (0.678), for interindividual comparison of kestrels on ad libitum maintenance regimens (0.786), and for mass proportionality (1.00). The circadian range of telemetered core temperature also varied more strongly with intraindividual than with interspecific (Aschoff, 1981a) variation in mass. This was due to reduced nocturnal core temperature at low-maintenance regimens, which was, however, insufficient to account for the excessive reduction in BMR. kidney lean mass at Carcass analysis of eight birds sacrificed revealed a disproportionate reduction in heart and kidney lean mass at low-maintenance regimens. We surmise that variation in BMR primarily reflects variation in these metabolically highly active tissues. This may account for positive correlations found between heart, kidney, and BMR residuals relative to interspecific allometric prediction, and between alpha and rho residuals, as expected on the basis of the constant excess of BMR during alpha above BMR during rho (Aschoff & Pohl, 1970a).

Animals↗

Reduction of human sleep duration after bright light exposure in the morning.

In 8 subjects the spontaneous termination of sleep was determined after repetitive exposure to either bright or dim light, between 6.00 and 9.00 h, on 3 days preceding sleep assessment. Sleep duration was significantly shorter following bright light than following dim light. During sleep the time course of EEG energy was not affected by the light treatment. Analysis of the time course of body temperature during sleep indicated an earlier rise of body temperature following the bright light treatment. In terms of the two-process model of sleep regulation this can be interpreted as a direct effect of light on the circadian phase of the wake up threshold.

Adult↗

Quantitative analysis of the effects of slow wave sleep deprivation during the first 3 h of sleep on subsequent EEG power density.

The relation between EEG power density during slow wave sleep (SWS) deprivation and power density during subsequent sleep was investigated. Nine young male adults slept in the laboratory for 3 consecutive nights. Spectral analysis of the EEG on the 2nd (baseline) night revealed an exponential decline in mean EEG power density (0.25-15.0 Hz) over successive nonrapid eye movement--rapid eye movement sleep cycles. During the first 3 h of the 3rd night the subjects were deprived of SWS by means of acoustic stimuli, which did not induce wakefulness. During SWS deprivation an attenuation of EEG power densities was observed in the delta frequencies, as well as in the theta band. In the hours of sleep following SWS deprivation both the power densities in the frequency range from 1 to 7 Hz and the amount of SWS were enhanced, relative to the same period of the baseline night. Both the amount of EEG energy accumulating subsequent to SWS deprivation and its time course could be predicted accurately from the EEG energy deficit caused by SWS deprivation. The data show that the level of integral EEG power density during a certain period after sleep onset depends on the amount of EEG energy accumulated during the preceding sleep rather than on the time elapsed since sleep onset. In terms of the two-process model of sleep regulation (Borbély 1982; Daan et al. 1984) this finding indicates that EEG power density reflects the rate of decay of the regulating variable, S, rather than S itself, as was originally postulated.

Acoustic Stimulation↗

EEG power density during nap sleep: reflection of an hourglass measuring the duration of prior wakefulness.

The relation between the duration of prior wakefulness and EEG power density during sleep in humans was assessed by means of a study of naps. The duration of prior wakefulness was varied from 2 to 20 hr by scheduling naps at 1000 hr, 1200 hr, 1400 hr, 1600 hr, 1800 hr, 2000 hr, and 0400 hr. In contrast to sleep latencies, which exhibited a minimum in the afternoon, EEG power densities in the delta and theta frequencies were a monotonic function of the duration of prior wakefulness. The data support the hypothesis that EEG power density during non-rapid eye movement sleep is only determined by the prior history of sleep and wakefulness and is not determined by clock-like mechanisms.

Adult↗

Commentary on the mutual interaction model of McCarley and Massaquoi for REM-NREM cycle.

McCarley and Massaquoi successfully simulated human REM-NREM cycle characteristics by extending the McCarley-Hobson model with two sets of assumptions, one creating limit cycle behavior, the other introducing two sources of circadian variation. We argue that the limit cycle assumptions, due to freedom in choosing parameter values, suffice to explain variation in REM across the night. Nonmonotonic circadian variation in REM latency requires a circadian cycle dependence only of initial conditions at sleep onset.

Animals↗

The timing of sleep in depression: theoretical considerations.

Endogenously depressed subjects frequently show severe sleep problems. In this article sleep time in depression is discussed in relation to a recently developed model for sleep timing in healthy subjects. In terms of the model, two parameter sets survive a qualitative comparison with the empirical data. These are a deficient increase of sleep need (process S) and, alternatively, an increased amplitude of random fluctuations in the wake threshold (process C). Distinct predictions based on these alternatives are discussed.

Circadian Rhythm↗

Distribution of REM latencies and other sleep phenomena in depression as explained by a single ultradian rhythm disturbance.

The McCarley-Hobson model, describing the alternation of NREM and REM sleep in the cat, was applied to human electroencephalographic data. The influence of initial conditions on oscillatory behavior was especially emphasized. It appears that the distribution of REM latency in depression, the abnormal accumulation of REM sleep, the variability of NREM-REM cycle duration, the frequent stage shifts, and frequent awakenings can be explained in this model by means of a decrease in the initial value of a single variable, which may be regarded as representing the strength of REM inhibition. The observation of slow wave sleep deficiency in depression may well be another reflection of this parameter.

Biological Clocks↗

Timing of human sleep: recovery process gated by a circadian pacemaker.

A model for the timing of human sleep is presented. It is based on a sleep-regulating variable (S)--possibly, but not necessarily, associated with a neurochemical substance--which increases during wakefulness and decreases during sleep. Sleep onset is triggered when S approaches an upper threshold (H); awakening occurs when S reaches a lower threshold (L). The thresholds show a circadian rhythm controlled by a single circadian pacemaker. Time constants of the S process were derived from rates of change of electroencephalographic (EEG) power density during regular sleep and during recovery from sleep deprivation. The waveform of the circadian threshold fluctuations was derived from spontaneous wake-up times after partial sleep deprivation. The model allows computer simulations of the main phenomena of human sleep timing, such as 1) internal desynchronization in the absence of time cues, 2) sleep fragmentation during continuous bed rest, and 3) circadian phase dependence of sleep duration during isolation from time cues, recovery from sleep deprivation, and shift work. The model shows that the experimental data are consistent with the concept of a single circadian pacemaker in humans. It has implications for the understanding of sleep as a restorative process and its timing with respect to day and night.

Bed Rest↗