The mathematical structure of the human sleep-wake cycle.
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Circadian Rhythm↗
Biomedical subjects
P H Gander
Publications and source records attributed to P H Gander.
Regular 48-hour cycling of sleep duration and mood in a 35-year-old woman: use of lithium in time isolation.
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Adult↗
Masking of the circadian rhythms of heart rate and core temperature by the rest-activity cycle in man.
Heart rate and core temperature are elevated by physical activity and reduced during rest and/or sleep. These masking effects may confound interpretation of rhythm waveforms, particularly in situations where the rest-activity rhythm has a different period from that of the core temperature rhythm. Such desynchronization often occurs temporarily as an individual adjusts to a new work shift or to a new time zone following rapid transmeridian travel, making it difficult to assess the impact of such schedule changes on the circadian system. The present experiments were designed to estimate the magnitude of these masking effects, by monitoring the heart rate, rectal temperature, and nondominant wrist activity (2-min samples) of 12 male subjects during 6 days of normal routine outside the lab and during 6 days of strict bedrest. Subjects also kept sleep, dietary, and exercise logs throughout the study. Average (20-min) waveforms were computed for each subject and each rhythm, at home and in bedrest. In addition, data were partitioned according to self-reported sleep and wake times and were analyzed separately for each state. Average waveform comparisons indicated that about 45% of the range of the circadian heart rate rhythm during normal routine was attributable to the masking effects of activity during wake, which also produced a 16% elevation in mean heart rate during wake and an 11% increase in mean heart rate overall. (Analysis of variance indicated that mean heart rate during sleep at home was not significantly different from the mean during sleep in bedrest.) On average, about 14% of the range of the circadian temperature rhythm during normal routine was attributable to the effects of activity masking. However, the change in range of the temperature rhythm, from home to bedrest, was very variable between subjects (-41% to +13%). This variability was not accounted for by age or by reported frequency of exercise at home. Normal activity during wake increased the mean temperature during wake by an average of 0.16 degrees C and the overall mean by about 0.12 degrees C. (Analysis of variance indicated that mean temperature during sleep at home was not significantly different from the mean during sleep in bedrest.) A 10-hr "night" (lights-off from 2200 to 0800 hr) was provided during bedrest, within which subjects could select their own sleep times. Times of sleep onset and wake onset were not significantly different between home and bedrest.(ABSTRACT TRUNCATED AT 400 WORDS)
Adult↗
International aircrew sleep and wakefulness after multiple time zone flights: a cooperative study.
An international research team has carried out an electroencephalographic study of sleep and wakefulness in flight crews operating long-haul routes across seven or eight time zones. Following baseline recordings, volunteer crews (n = 56) from four airlines spent their first outbound layover at a sleep laboratory. This paper provides an overview of the project's history, its research design, and the standardization of procedures. The overall results are remarkably consistent among the four participating laboratories and strongly support the feasibility of cooperative international sleep research in the operational arena.
Adult↗
Circadian rhythmicity and fatigue in flight operations.
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Aerospace Medicine↗
Effectiveness of cyclic intragastric feeding as a circadian zeitgeber in the squirrel monkey.
To investigate the mechanism of circadian synchronization by oral food consumption, we tested the effectiveness of cyclic intragastric feeding in synchronizing squirrel monkeys to a 24-hour schedule. Five monkeys were prepared with intragastric catheters and were studied free-ranging within a cage in an environmentally controlled chamber with continuous monitoring of body temperature (every 15 min) and locomotor activity (every 30 min). A balanced mixture of dextrose, amino acids, fat emulsion, vitamins and minerals was infused from 0800-2000 every day via the gastrostomy, first during a 12-hour light and 12-hour dark cycle (lights on 0800-2000 hours), and then subsequently in constant illumination. Serial phase analysis was performed by computing cycle-by-cycle acrophases of best-fitting sinusoids. One animal was synchronized by the feeding regimen while four others exhibited relative coordination. We conclude from these results that the timing of intragastic feeding may have only a weak synchronizing effect on primate circadian rhythms and that pre-gastric cues associated with oral food ingestion contribute to entrainment by feeding schedules.
Animals↗
Forced internal desynchronization between circadian temperature and activity rhythms in squirrel monkeys.
In an attempt to force internal desynchronization between the rest-activity rhythm and the body temperature rhythm of the squirrel monkey (Saimiri sciureus), five animals were studied in a 14:14 light-dark cycle. In four animals a 28-h spectral component was found to predominate in the rest-activity rhythm, whereas an unentrained circadian component (tau = 25.9 +/- 0.4 h) predominated in the body temperature rhythm. Plots of the cycle-by-cycle acrophases of the two rhythms confirm that they desynchronize, due to the failure of the temperature rhythm to entrain to the light-dark cycle. These data from intact animals provide further support for the hypothesis that the squirrel monkey circadian timing system has at least two pacemakers. A rhythm for which the supra-chiasmatic nuclei (SCN) have previously been shown to be essential (rest-activity) simultaneously exhibited a different period from a rhythm (body temperature) that has been shown to persist after destruction of the SCN.
Animals↗
Phase shifting two coupled circadian pacemakers: implications for jet lag.
Flights across time zones produce an abrupt displacement of the environmental time cues (zeitgebers), and the endogenous circadian timing system resynchronizes only gradually to the new schedule. A coupled two-oscillator model can simulate the human circadian system in temporal isolation and in artificial zeitgeber cycles. The model is here shown to explain the major features of resynchronization of circadian rhythms after time zone shifts, i.e., the rate of adjustment depends on the rhythm being measured, the number of time zones crossed, the flight direction (eastward or westward), and the strength of the zeitgebers in the new time zone. Investigations of the contribution of different model parameters to system performances suggest that intersubject differences in pacemaker periods may be a major factor in the observed variability in the effects of time zone shifts on circadian rhythms. With individualized period estimate the models can simulate case studies in which four subjects recorded their sleep-wake and core body temperature rhythms throughout simple and complex patterns of transmeridian flights.
Body Temperature↗
Role of the suprachiasmatic nuclei in the circadian timing system of the squirrel monkey. I. The generation of rhythmicity.
The circadian organization of squirrel monkey (Saimiri sciureus) drinking behavior was evaluated before and after the placement of radiofrequency lesions which completely destroyed the suprachiasmatic nuclei (SCN) in 4 monkeys and partially ablated the SCN in another 4 animals. In continuous illumination (LL: 600 lux) prior to surgery, each monkey had a precise free-running circadian rhythm of drinking behavior with a period of 25.31 +/- 0.21 h (means +/- S.E.M.). By 4-6 weeks following the lesions, the temporal organization of drinking behavior had become disrupted, but a statistically significant free-running circadian rhythm was still detectable by time series analyses. Subsequently, the circadian organization of drinking behavior in 7 out of 8 monkeys gradually decayed with either no statistically significant rhythmicity or only weak circadian and/or ultradian rhythmicity detected by time series analyses by 10-92 weeks post-lesion. The remaining animal which maintained a statistically reliable free-running rhythm in drinking behavior received the least damage (less than 50%) to the SCN. Despite the major alterations in the temporal patterning of behavior, the overall amount of drinking behavior per 24 h was unchanged. The SCN are thus essential for maintaining the circadian organization of squirrel monkey drinking behavior. However, the existence of residual circadian rhythmicity following SCN lesions and the gradual decay of circadian organization thereafter suggest that the SCN may coordinate the activity of circadian oscillators which lie outside its borders.
Animals↗
Simulating the action of zeitgebers on a coupled two-oscillator model of the human circadian system.
Our two-oscillator model was originally designed to describe the circadian rhythms of human subjects maintained in temporal isolation. The performance of this model in response to simulated environmental synchronizing cycles (zeitgebers) is examined here. Six distinct types of synchronization are demonstrated between the x oscillator (postulated to regulate the core temperature rhythm), the y oscillator (postulated to regulate the rest-activity rhythm), and z (the zeitgeber). Four types of synchronization are identifiable, if we consider only the periods of the three oscillators. Both x and y may be synchronized by z; either may synchronize with z while the other exhibits a different period; or x, y, and z may each show different periods. Two further classes of synchronization are discernible when phase criteria are taken into account. When either x or y is on the verge of desynchronizing from the other two oscillators, it undergoes periodic phase modulations while retaining the common overall period. The type of synchronization observed depends on the periods of x, y, and z and on the strength of the z drive. The effects of modifying each of these parameters have been systematically investigated by simulation, and model performance is summarized in terms of range of entrainment "maps." These constitute extensive sets of predictions about expected patterns of entrainment of the core temperature and rest-activity rhythms of human subjects exposed to various environmental zeitgebers. Experimental data are available against which model predictions can be tested.
Body Temperature↗
Modeling the action of zeitgebers on the human circadian system: comparisons of simulations and data.
In our two-oscillator model for the human circadian timing system, the effect of an environmental synchronizing cycle (zeitgeber) is determined by the periods of the two oscillators and of the zeitgeber and by the zeitgeber strength. The oscillators x and y are postulated to regulate the core temperature and rest-activity rhythms, respectively. From published examples of experiments in which human subjects were exposed to artificial zeitgebers, it is possible to derive estimates of the periods of the core temperature and/or the rest-activity rhythms and the zeitgeber. Two strengths of zeitgeber input to the model that correspond to two of the common zeitgeber regimes used in human entrainment studies have now been elucidated. Thus specific model simulations can be generated for each of the available examples of entrainment of human subjects by artificial zeitgebers. Such simulations indicate that the model can reproduce, with remarkable subtlety, the types of full and partial entrainment observed experimentally. The model is thus demonstrated to accommodate the period and phase control of endogenous rhythms by environmental zeitgebers, which is a crucial functional attribute of circadian timing systems.
Activity Cycles↗
Phase-resetting action of light on the circadian activity rhythm of Rattus exulans.
The phase resetting action of light on the circadian rhythm of locomotor activity has been examined in wild-caught Polynesian rats (Rattus exulans). Phase-response curves to 4-, 8-, and 16-h light pulses have been derived. All three curves conform to the generalization that pulses occurring during late subjective day and early subjective night produce delays, whereas advances occur in response to pulses coinciding with the late subjective night and early subjective day. Weak (type 1) phase resetting is observed in response to 4-h pulses and perhaps 8-h pulses, whereas strong (type 0) resetting apparently occurs in response to 16-h pulses. These data evidently constitute the first report of strong phase resetting in mammalian circadian rhythms. The phase relationship between an entrained activity rhythm and the light-dark cycle is dependent on the photoperiod and, in 24-h cycles, on the period difference between the rhythm and the zeitgeber. In longer zeitgeber cycles, activity onset is delayed by a direct masking effect of light. A primarily nonparametric action of light in natural entrainment is consistent with these data and with field observations.
Animals↗
Light-dark masking of circadian temperature and activity rhythms in squirrel monkeys.
In addition to the synchronizing effects of light-dark cycles on the circadian timing system, light per se can have a direct masking effect on the rhythmic variables monitored as indicators of the circadian system. To evaluate the masking action of light on the rest-activity and body temperature rhythms of squirrel monkeys (Saimiri sciureus), animals were exposed to cycles of 2 h of light alternating with 2 h of darkness (LD 2:2). There was evidence of phase-dependent masking responses in both rhythms, with the light-induced elevation of activity levels and temperature greatest around the times of the circadian maxima of the respective rhythms. The temporal distribution of activity was more strongly affected by the masking action of the 2:2 cycle than was the temporal pattern of temperature variation. The increase in activity observed during lights on was evidently a major factor contributing to the corresponding increase in temperature.
Animals↗
Gradual decay of circadian drinking organization following lesions of the suprachiasmatic nuclei in primates.
The circadian organization of squirrel monkey drinking behavior was examined in constant illumination following lesions which totally destroyed the suprachiasmatic nuclei. Although these lesions eventually led to a severe disruption in the temporal organization of drinking behavior, a statistically significant circadian pattern of drinking was observed in each monkey for at least 4-6 weeks after lesioning.
Animals↗
Task variables determine which biological clock controls circadian rhythms in human performance.
There are circadian (approximately 24 h) rhythms for a wide range of human physiological and psychological functions including mood and performance efficiency. These rhythms are self-sustaining in conditions of temporal isolation, indicating that internal oscillators (or biological clocks) control them. Recent research has proposed an endogenous two-oscillator model of the human circadian system, with one oscillator indicated by the core body temperature rhythm and a second oscillator responsible for the daily cycle of sleep and wakefulness. The present study was designed to produce a desynchronization in period between the two oscillators, to assess directly the behaviour of the rhythms of different performance tasks. The results, reported here, indicate that a simple manual dexterity task is almost entirely under the control of the temperature rhythm oscillator, whereas a more complex cognitive task demonstrates a periodicity which appears to be influenced by those oscillators controlling temperature and the sleep/wake cycle.
Adult↗