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R E Kronauer

Publications and source records attributed to R E Kronauer.

71 records · Page 4Linked to original sources

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↗

Strength and cycle time of high-altitude ventilatory patterns in unacclimatized humans.

Respiration was monitored with magnetometers in 12 healthy supine young adults at sea level and in an altitude chamber at simulated high altitudes of 8,000, 9,000, 11,000, and 14,000 ft. Periodic breathing that was strong enough to include apnea at the time of minimum ventilation was seen in all subjects at high altitude. Cycle time of periodic breathing ranged from 12 to 34 s. On average across the population the incidence of periodic breathing increased with altitude. Cycle time of the periodic pattern increased as strength of the pattern increased. After normalizing to a standard pattern strength, cycle time decreased as altitude increased. The study included two series of experiments, the second occurring 3 wk after the first and involving seven of the same subjects. The standard cycle time at 14,000 ft for each subject in the second series was the same as in the first series to within, on the average, 6%. Each subject studied at 11,000 ft in both series reproduced his cycle time to within, on the average, 9%. The variation of standard cycle time for a given subject is less than the variation across the population, indicating characteristic cycle times for some individuals (one-way analysis of variance, P less than 0.025).

Acclimatization↗

Opponent-movement mechanisms in human vision.

A vertical grating that sinusoidally reverses contrast can be synthesized from two identical component gratings that move with equal velocities in opposite directions (leftward and rightward). Such a counterphase grating is used as a suprathreshold masking pattern. When the mask is of low spatial frequency and is modulated rapidly, a test pattern consisting of an increment of the rightward component and an equivalent simultaneous decrement of the leftward component is highly detectable compared with simultaneous increments or decrements of both components. The visibility of the opponent-movement test signal is strongly facilitated by high-contrast masks. This facilitation is accompanied by a high sensitivity for judging the direction of motion of the test. These results show that certain detection mechanisms are highly sensitive to the difference of the rightward and leftward components. However, when the mask is of threshold contrast, the rightward- and leftward-moving test components appear to be detected independently. A high-contrast grating that rapidly moves in one direction strongly masks gratings moving in the same or opposite direction; this shows that moving patterns are not detected by unidirectional mechanisms when contrast is clearly suprathreshold. The results may be explained by a model with mechanisms that are excited by one direction of motion and inhibited by the opposite direction.

Discrimination Learning↗

Synchronized cycles in ventilation and vocal activity during spontaneous conversational speech.

Ventilation was monitored in 10 human subjects during spontaneous conversational speech to determine whether oscillatory patterns in vocal activity were correlated with oscillatory patterns in ventilation. The 10 subjects were studied as five pairs (or dyads), and spontaneous conversation occurred within each dyad. Patterns with cycle times ranging from 16 to 512 s were studied. Of the 10 subjects in this study, 1 subject showed a very striking pattern of mutual entrainment between low-frequency oscillations in ventilation and in vocal activity, 5 subjects showed somewhat weaker coordination between oscillations in ventilation and vocal activity, and the remaining 4 subjects showed little or no coupling between ventilation and vocal activity. Mutual entrainment between rhythms in ventilation and vocal activity can occur, but this study suggests that there may be great differences among individuals in the degree to which ventilation covaries with vocal activity in spontaneous conversational speech. We hypothesize that degree of entrainment is affected both by the strength of any spontaneous ventilatory patterns and by the extent to which any individual's spontaneous ventilatory pattern matches that of his conversational partner.

Adult↗

Mathematical model of the human circadian system with two interacting oscillators.

Human subjects during extended isolation from environmental time cues show complex variations in timing and duration of sleep with a progressive pattern, which eventually results in rest-activity and body temperature rhythms having different average periods. We present a model where temperature and rest-activity are each governed by an oscillator of the van der Pol type, denoted x and y, respectively. The oscillators affect one another through "velocity" type coupling, the effect of x on y being about four times greater than y on x. Periodic zeitgeber, z, is modeled as forcing only on y. We find that the entire pattern sequence can be realistically reproduced by causing only the intrinsic period of the y oscillator to increase while that of x remains stable. Desynchronization between x and y is the result of the intrinsic periods of the two oscillators becoming so disparate that the coupling is no longer able to enforce synchrony. Prior to desynchronization both human subjects and our model exhibit "phase trapping" wherein the relative phase of x and y is slowly modulated although the average x and y periods match. The model phase relations between temperature and both the timing and duration of sleep are, throughout, in good agreement with entrained and free-running human data. Most importantly, the model shows that the dramatic change in the length of the rest-activity cycle when desynchronization occurs is actually due to a relatively small variation in the governing variable, y.

Body Temperature↗

Factors inducing periodic breathing in humans: a general model.

A general model is developed to account for all kinds of periodic breathing (PB) resulting from instability in respiratory control: in normals during sleep and on acute exposure to high altitude, in sleeping infants, and in patients with cardiovascular or neurologic lesions. It is found that in almost every case the ventilatory oscillation is mediated predominantly by the peripheral controller. System stability is decreased by hypoxia, hypercapnia, increased lung washout times, prolonged lung-chemoreceptor delays, and high controller sensitivity. Stability is enhanced by large lung CO2 and O2 storage volumes but little affected by body tissue stores. Using our own measurements of lung-ear delays, the model predicts that the mean cycle time of PB decreases from about 30 s at sea level to 20 s at 14,000 ft, in excellent agreement with data from other studies. Allometric scaling of the relevant parameters also shows close agreement between model predictions and data obtained on infants.

Adult↗

Spatial adaptation of short-wavelength pathways in humans.

Color-selective spatial adaptation of the short-wavelength, or blue-sensitive, pathway was demonstrated. The adaptation was orientation selective and strongly monocular. Adaptation was assessed by measuring visibility thresholds for monochromatic gratings in subjects adapted to high-contrast violet gratings designed to stimulate only blue-sensitive cones. The results showed spatially selective, adaptable channels within the short-wavelength pathway.

Adaptation, Physiological↗

Methods for identifying respiratory oscillations disclose altitude effects.

The respiration of 16 supine subjects was monitored at sea level and at an altitude of 3,050 m. Breath shape was defined by inspiratory and expiratory durations [TI, TE] and volumes [VI, VE], total breath duration [Tt], and ventilation [.VE(=VE/Tt)]. Dynamic breath pattern analysis revealed that the chronological sequences of these variables often display large sustained oscillations. Depending on their relative phase the simultaneous oscillations in VE and Tt are "compensating" or "reinforcing." The first typifies the traditional view that breath-to-breath changes in VE and Tt oscillate in phase, i.e., deeper breaths are longer thereby reducing ventilatory fluctuations. For the reinforcing oscillations breath-to-breath changes in VE and Tt oscillate out-of-phase, i.e., deeper breaths tend to be shorter, thereby enhancing ventilatory fluctuations. The observed amplitudes of the reinforced .VE oscillations are as large as 75% of the mean .VE level. In five subjects strong reinforcing oscillations with characteristic period of 20.6 +/- 3.5 (SD) s accompanied high-altitude exposure (most prominently after acclimatization) and did not appear to be a function of acid-base balance.

Altitude↗

Using a comb filter to describe time-varying biological rhythmicities.

A very important problem in the analysis of biological data sequences is the detection of oscillations in the presence of random variations (noise). If the oscillations are not stationary, i.e., if they drift in frequency and amplitude, or occur in bursts, traditional analysis techniques utilizing the power spectrum or its time-domain equivalent, the autocorrelation function, can be both misleading and insensitive. Temporal filtering by a "comb" or se of band-pass filters is very effective for identifying and describing nonstationary oscillations. The basic procedures for interpreting the output of a comb filter are presented here, illustrated by examples using predefined input test sequences.

Computers↗

Apparent saturation of blue-sensitive cones occurs at a color-opponent stage.

Response saturation of blue-sensitive cone pathways was studied by measuring increment thresholds for violet test flashes on flashed violet fields in the presence of a steady yellow "auxiliary" field of constant radiance. Adding intense yellow field flashes to the violet field flash could eliminate or reduce response saturation (greatly reduce threshold), whereas "negative" yellow field flashes drove the mechanism to further saturation. The response saturation is thus not, in general, controlled exclusively by independent blue-sensitive cones but by spectrally opponent mechanisms that receive opposite-signed signals from blue-sensitive cones and from green-or red-sensitive cones. These results add to a growing number of studies that demonstrate that detection of signals from blue-sensitive cones is largely through a color-opponent pathway.

Action Potentials↗

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↗