Ultradian, circahoral and circadian structures in endothermic vertebrates and humans

1. For more than 30 years many studies have been carried out concerning rhythms with periods approaching 24 hr (circadian rhythms). 2. The latter have been demonstrated as resulting from environmental 24 hr synchronizers (zeitgebers), but they usually persist in the absence of a 24 hr synchronization, which proves their endogenous nature. 3. Biological rhythms with periods less than 20 hr (ultradian rhythms) and particularly those approaching 1 hr (circahoral rhythms) have been determined: for motility, rest-activity, sleep phases, endocrine secretions and other physiological functions. 4. These ultradian and circahoral rhythms have been found in rodents, birds, monkeys and humans. 5. Existing at all stages of ontogeny, they have been proved to be endogenous and species and strain specific. 6. As these ultradian rhythms can be influenced by environmental factors and sometimes by circadian rhythms they are not truly periodic, so therefore cannot be computed by the usual processes of mathematical time analysis.     

Small bowel and plasma gastrin rhythms in cats

The purpose of this experiment was to study the possible role of the gastric antrum and small bowel in the rhythm(s) of plasma gastrin. The cat was used as the laboratory animal. Three groups of cats were provided with a gastric fistula for the study of gastric acid and plasma gastrin rhythms. The first group (N = 7) served as controls. A second group (N = 3) was antrectomized and later subjected to a 80% small bowel resection. Gastric acid secretions were collected every 30 min from 0800 to 2400. Blood samples for determination of gastrin were drawn every 2 hr from 0800 to 2400. In control animals a circadian (i.e. approximately 24 hr) and 3 ultradian (i.e. less than 24 hr) rhythms were detected for acid output. In the antrectomized cats, circadian and ultradian rhythms were documented. After small bowel resection circadian and ultradian rhythms in gastric acid secretion were observed. For plasma gastrin, circadian and ultradian rhythms were found in the control cats. In the antrectomized cats no rhythms were observed. After small bowel resection an ultradian rhythm reappeared in these antrectomized cats. Removal of the antrum in the cat induces disappearance of circadian and ultradian rhythms of plasma gastrin but fails to modify the acid rhythms. Small bowel resection results in the reappearance of an ultradian rhythm for plasma gastrin and a shift in acrophase for the circadian rhythm in acid secretion.     

Ultradian rhythm unmasked in the Pdf clock mutant of Drosophila

A diverse range of organisms shows physiological and behavioural rhythms with various periods. Extensive studies have been performed to elucidate the molecular mechanisms of circadian rhythms with an approximately 24 h period in both Drosophila and mammals, while less attention has been paid to ultradian rhythms with shorter periods. We used a video-tracking method to monitor the movement of single flies, and clear ultradian rhythms were detected in the locomotor behaviour of wild type and clock mutant flies kept under constant dark conditions. In particular, the Pigment-dispersing factor mutant (Pdf ⁰¹ ) demonstrated a precise and robust ultradian rhythmicity, which was not temperature compensated. Our results suggest that Drosophila has an endogenous ultradian oscillator that is masked by circadian rhythmic behaviours.     

Free-Running Locomotor Activity Rhythms in Cave-Dwelling Catfishes,Trichomycterus sp., from Brazil (Teleostei,Siluriformes)

Locomotor activity of troglobitic (cave restricted) catfishes, Trichomycterus sp., was recorded in the laboratory under constant darkness for ten consecutive days. Infra-red photocell beam crossings were totalled every thirty minutes and stored for later analysis. Spectral analysis of the data followed by a statistical test designed for the detection of significant components was then performed. The results of 14 individuals, seven pigmented and seven albino fishes, are reported here. In eight individuals, four pigmented and four albinos, we were not able to detect significant components in the circadian range, although ultradian and/or infradian significant components were found in all cases. Former studies on other two Brazilian cave catfishes, Pimelodella kronei and Imparfinis sp., showed similar results, suggesting a picture of gradual loss of circadian rhythmicity and persistence of ultradian and infradian rhythms in the locomotor activity of troglobitic populations. The proportion of specimens without significant circadian components was higher in Trichomycterus sp., supposedly less specialized to cave life than the other two species. This is tentatively correlated with a higher environmental stability during the time of evolutionary isolation in the subterranean biotope. These results provide strong arguments for the importance of external selection over circadian rhythmicity.     

Free-Running Locomotor Activity Rhythms in Cave-Dwelling Catfishes, Trichomycterus sp., from Brazil (Teleostei, Siluriformes)

Locomotor activity of troglobitic (cave restricted) catfishes, Trichomycterus sp., was recorded in the laboratory under constant darkness for ten consecutive days. Infra-red photocell beam crossings were totalled every thirty minutes and stored for later analysis. Spectral analysis of the data followed by a statistical test designed for the detection of significant components was then performed. The results of 14 individuals, seven pigmented and seven albino fishes, are reported here. In eight individuals, four pigmented and four albinos, we were not able to detect significant components in the circadian range, although ultradian and/or infradian significant components were found in all cases. Former studies on other two Brazilian cave catfishes, Pimelodella kronei and Imparfinis sp., showed similar results, suggesting a picture of gradual loss of circadian rhythmicity and persistence of ultradian and infradian rhythms in the locomotor activity of troglobitic populations. The proportion of specimens without significant circadian components was higher in Trichomycterus sp., supposedly less specialized to cave life than the other two species. This is tentatively correlated with a higher environmental stability during the time of evolutionary isolation in the subterranean biotope. These results provide strong arguments for the importance of external selection over circadian rhythmicity.     

THE OCCURRENCE AND FUNCTIONS OF ULTRADIAN RHYTHMS

Ultradian oscillations with periods between 5 min and 4 h have been described in cell-free extracts, single-celled eukaryotes, cultured cells and embryos. Whereas some of these potentially oscillatory systems (e.g. glycolysis) may only exhibit this type of behaviour rarely if at all in vivo , other ultradian oscillators in lower eukaryotes are rhythms and probably have timekeeping functions. Rhythms with ultradian periods of 10 min to 20 h in oxygen consumption and carbon dioxide production have also been studied in endotherm animals: these rhythms may be modified by variations of environmental parameters and by circadian and infradian synchronizers. Interspecies and interstrain differences strongly suggest that these rhythms are endogenous and have a genetic origin. We suggest that the temporal organization of biochemical and physiological processes facilitates optimization of thermodynamic maintenance of the organism within the random fluctuations of its physicochemical environment and contributes to genetic selection.     

Cellular oscillators: rhythmic gene expression and metabolism

Many biological processes are driven by biological clocks that, depending on the frequency they generate, are classified into ultradian, circadian and infradian oscillators. In virtually all light-sensitive organisms from cyanobacteria to humans, a circadian timing system adapts cyclic physiology to geophysical time. Recent evidence suggests that even in mammals circadian oscillators function in a cell-autonomous manner. In yeast, an ultradian oscillator regulates cyclic respiratory activity and global gene expression. Circadian oscillators and the ultradian yeast respiratory clock share at least four properties: they follow limit-cycle kinetics, interweave with cellular metabolism, are temperature-compensated and influence the cell division clock.     

Chronobiology: anatomy in time

Chronobiology is that branch of science that objectively explores and quantifies mechanisms of biological time structure, including the important rhythmic manifestations of life. It is the study of biological rhythms. This paper introduces chronobiology and some of its vocabulary, principles, and techniques. A circadian rhythm is a regularly repetitive, quantitative physiological change with a period of about 24 hr (20-28), but the spectrum of rhythms includes those with periods less than 20 hr (ultradian) and longer than 28 hr (infradian). These rhythms are ubiquitous among the eukaryotes, innate and endogenous; their periods are precisely controlled by synchronizers in the environment. Rhythms can be manipulated by altering their synchronizers or by introducing more dominant ones. When organisms are removed from their environment and placed in constant conditions, rhythms revert to their natural frequencies and free-run. All of an organism's rhythms operate simultaneously, but their peaks and troughs do not necessarily occur at the same time. There are rhythms in susceptibility to drugs; a fixed dose may have a therapeutic effect at one point along the 24 hr time scale and a harmful one at another. Knowledge of these rhythms can be important when designing experimental or treatment protocols and interpreting results. Examples are provided to show that single-time-point sampling can lead to erroneous results, unless biological periodicity is taken into consideration.     

Oscillations in joint synchrony of reproductive hormones in healthy men

Negative-feedback (inhibitory) and positive-feedforward (stimulatory) processes regulate physiological systems. Whether such processes are themselves rhythmic is not known. Here, we apply cross-approximate entropy (cross-ApEn), a noninvasive measurement of joint (pairwise) signal synchrony, to inferentially assess hypothesized circadian and ultradian variations in feedback coupling. The data comprised simultaneous measurements of three pituitary and one peripheral hormone (LH, FSH, prolactin, and testosterone) in 12 healthy men each sampled every 10 min for 4 days (5,760 min). Ergodicity, due to the time series stationarity of the measurements over the 4 days, allows for effective estimation of parameters based upon the 12 subjects. Cross-ApEn changes were quantified via moving-window estimates applied to 4-day time series pairs. The resultant ordered windowed cross-ApEn series (in time) were subjected to power spectrum analysis. Rhythmicity was assessed against the null hypothesis of randomness using 1,000 simulated periodograms derived by shuffling the interpulse-interval hormone-concentration segments and redoing cross-ApEn windows and spectral analysis. By forward cross-ApEn analysis, paired LH-testosterone, LH-prolactin, and LH-FSH synchrony maintained dominant rhythms with periodicities of 18-22.5, 18, and 22.5 h, respectively (each P < 0.001). By reverse (feedback) cross-ApEn analysis, testosterone-LH, testosterone-prolactin, and testosterone-FSH synchrony cycles were 30, 18, and 30-45 h, respectively (each P ≤ 0.001). Significant 8- or 24-h rhythms were also detected in most linkages, and maximal bihormonal synchrony occurred consistently at ～0400-0500. Collectively, these analyses demonstrate significant ultradian (<24 h), circadian (～24 h), and infradian (>24 h) oscillations in pituitary-testis synchrony, wherein maximal biglandular coordination is strongly constrained to the early morning hours.     

Effect of geomagnetic activity on the rhythm of resistance of rats to hypoxia

The resistance of male Wistar rats to acute hypoxia was estimated from the lifetime at an "altitude" of 11.5 km above sea level from 13 to 21 p.m. and in different seasons of a year. Identical types of rhythms (circaseptan, circasemiseptan, infradian, circadian, and ultradian) of geomagnetic activity and lifetime were revealed. It was found that the periods of basal rhythms either coincide or are multiple. It was shown that the rhythms of geomagnetic activity (including nonbasal rhythms) affect lifetime rhythms (especially ultradian rhythms). As the periods of the rhythms decrease, the number of rhythms for both parameters increase (ultradian rhythms are most numerous), and relative differences in the values of periods (they are minimal for ultradian rhythms) and the amplitudes of rhythms decrease.     

Small Bowel and Plasma Gastrin Rhythms in Cats

The purpose of this experiment was to study the possible role of the gastric antrum and small bowel in the rhythm(s) of plasma gastrin. The cat was used as the laboratory animal. Three groups of cats were provided with a gastric fistula for the study of gastric acid and plasma gastrin rhythms. The first group (N = 7) served as controls. A second group (N = 3) was antrectomized and later subjected to a 80% small bowel resection. Gastric acid secretions were collected every 30 min from 0800 to 2400. Blood samples for determination of gastrin were drawn every 2hr from 0800 to 2400. In control animals a circadian (i.e.<24hr) and 3 ultradian (i.e.<24 hr) rhythms were detected for acid output. In the antrectomized cats, circadian and ultradian rhythms were documented. After small bowel resection circadian and ultradian rhythms in gastric acid secretion were observed. For plasma gastrin, circadian and ultradian rhythms were found in the control cats. In the antrectomized cats no rhythms were observed. After small bowel resection an ultradian rhythm reappeared in these antrectomized cats. Removal of the antrum in the cat induces disappearance of circadian and ultradian rhythms of plasma gastrin but fails to modify the acid rhythms. Small bowel resection results in the reappearance of an ultradian rhythm for plasma gastrin and a shift in acrophase for the circadian rhythm in acid secretion.     

Behavioral analysis of circadian rhythms: Entraining the circadian clock and determining the food-entrainable oscillator mechanism

Sleep and Biological Rhythms - Circadian rhythms are oscillations in behavior and physiological functions that are regulated by internal biological clocks. In mammals, the circadian rhythms can be...     

From Ultradian to Infradian Rhythms: LH Releases Patterns In Vitro

In the present study, we examined in vitro luteinizing hormone (LH) release patterns from pituitaries and from pituitary cell cultures (3 and 7 days in culture) to elucidate the endogenous period generated by the gonadotroph cell population and to evaluate the relationship between the basic period generated at the cellular level and the output pattern observed at the organ level. In addition, we examined the effect of photic environmental signals perceived by the animals on LH release patterns from pituitaries in vitro. When the animals were exposed to circadian photoperiodic signals, the in vitro LH release pattern from the pituitaries exhibited ultradian, circadian, and infra-dian frequencies. When the animals were exposed to continuous illumination, the in vitro patterns exhibited only ultradian and infradian frequencies. Furthermore, free running is a process, not a state. This process is driven by a change in the relative dominance of different frequencies that construct the pattern without changing the basic period length. Evaluation of the relative dominance of the different frequencies that construct the pattern indicates that, although infradian oscillators may take part in shaping the output pattern, the basic rhythm generated by the pituitary cells is in the ultradian domain. The results obtained from the examined system suggest that an endogenous oscillator is a cellular entity with ultradian periodicity, and that the rhythmic output of many biological variables is structured by various ultradian components that construct the circadian and infradian output rhythms.     

From Ultradian to Infradian Rhythms: LH Releases Patterns In Vitro

In the present study, we examined in vitro luteinizing hormone (LH) release patterns from pituitaries and from pituitary cell cultures (3 and 7 days in culture) to elucidate the endogenous period generated by the gonadotroph cell population and to evaluate the relationship between the basic period generated at the cellular level and the output pattern observed at the organ level. In addition, we examined the effect of photic environmental signals perceived by the animals on LH release patterns from pituitaries in vitro. When the animals were exposed to circadian photoperiodic signals, the in vitro LH release pattern from the pituitaries exhibited ultradian, circadian, and infra-dian frequencies. When the animals were exposed to continuous illumination, the in vitro patterns exhibited only ultradian and infradian frequencies. Furthermore, free running is a process, not a state. This process is driven by a change in the relative dominance of different frequencies that construct the pattern without changing the basic period length. Evaluation of the relative dominance of the different frequencies that construct the pattern indicates that, although infradian oscillators may take part in shaping the output pattern, the basic rhythm generated by the pituitary cells is in the ultradian domain. The results obtained from the examined system suggest that an endogenous oscillator is a cellular entity with ultradian periodicity, and that the rhythmic output of many biological variables is structured by various ultradian components that construct the circadian and infradian output rhythms.     

TEMPORAL PATTERN OF LH SECRETION: REGULATION BY MULTIPLE ULTRADIAN OSCILLATORS VERSUS A SINGLE CIRCADIAN OSCILLATOR

The possibility that the 24h rhythm output is the composite expression of ultradian oscillators of varying periodicities was examined by assessing the effect of external continuously or pulsed (20-minute) Gonadotropinreleasing hormone (GnRH) infusions on in vitro luteinizing hormone (LH) release patterns from female mouse pituitaries during 38h study spans. Applying stepwise analyses (spectral, cosine fit, best-fit curve, and peak detection analyses) revealed the waveform shape of LH release output patterns over time is composed of several ultradian oscillations of different periods. The results further substantiated previous observations indicating the pituitary functions as an autonomous clock. The GnRH oscillator functions as a pulse generator and amplitude regulator, but it is not the oscillator that drives the ultradian LH release rhythms. At different stages of the estrus cycle, the effect of GnRH on the expression of ultradian periodicities varies, resulting in the modification of their amplitudes but not their periods. The functional output from the system of ultradian oscillators may superimpose a “circadian or infradian phenotype” on the observed secretion pattern. An “amplitude control” hypothesis is proposed: The temporal pattern of LH release is governed by several oscillators that function in conjunction with one another and are regulated by an amplitude-controlled mechanism. Simulated models show that such a mechanism results in better adaptive response to environmental requirements than does a single circadian oscillator. (Chronobiology International, 18(3), 399–412, 2001)     

Temporal pattern of LH secretion: regulation by multiple ultradian oscillators versus a single circadian oscillator

The possibility that the 24h rhythm output is the composite expression of ultradian oscillators of varying periodicities was examined by assessing the effect of external continuously or pulsed (20-minute) Gonadotropinreleasing hormone (GnRH) infusions on in vitro luteinizing hormone (LH) release patterns from female mouse pituitaries during 38h study spans. Applying stepwise analyses (spectral, cosine fit, best-fit curve, and peak detection analyses) revealed the waveform shape of LH release output patterns over time is composed of several ultradian oscillations of different periods. The results further substantiated previous observations indicating the pituitary functions as an autonomous clock. The GnRH oscillator functions as a pulse generator and amplitude regulator, but it is not the oscillator that drives the ultradian LH release rhythms. At different stages of the estrus cycle, the effect of GnRH on the expression of ultradian periodicities varies, resulting in the modification of their amplitudes but not their periods. The functional output from the system of ultradian oscillators may superimpose a “circadian or infradian phenotype” on the observed secretion pattern. An “amplitude control” hypothesis is proposed: The temporal pattern of LH release is governed by several oscillators that function in conjunction with one another and are regulated by an amplitude-controlled mechanism. Simulated models show that such a mechanism results in better adaptive response to environmental requirements than does a single circadian oscillator. (Chronobiology International, 18(3), 399-412, 2001)     

Changes in Body Temperature Rhythm after Traumatic Brain Injury

We performed this study to determine whether in head injured patients body temperature rhythmicity exists outside the usual spectrum. Temperature data of in total 22 patients with head injury were analyzed using the Regressive and Iterative Cosinor methods. We found that circadian rhythm often remained, and usually was combined with rhythms in ultradian or infradian ranges. Tau shifts over consecutive days were observed in three severely head injured patients (Glasgow Coma Scale score ≤ 8). To validate the results we used surrogate data. Detection of temperature rhythms in this study may serve to estimate the clinical importance of biological rhythms in head injury.     

Changes in Body Temperature Rhythm after Traumatic Brain Injury

We performed this study to determine whether in head injured patients body temperature rhythmicity exists outside the usual spectrum. Temperature data of in total 22 patients with head injury were analyzed using the Regressive and Iterative Cosinor methods. We found that circadian rhythm often remained, and usually was combined with rhythms in ultradian or infradian ranges. Tau shifts over consecutive days were observed in three severely head injured patients (Glasgow Coma Scale score ≤ 8). To validate the results we used surrogate data. Detection of temperature rhythms in this study may serve to estimate the clinical importance of biological rhythms in head injury.     

Circadian and Ultradian Variations in the Plasma Level of Maturational Gonadotropin (GTH II) in the Indian Catfish, Clarias batrachus

Analysis of changes in the plasma levels of GTH II over a 24-h day/night cycle revealed statistically significant circadian or ultradian variation or both in its circulating level during the gonadally active months of the reproductive cycle. The 24-h average and amplitude of circadian or ultradian rhythm in GTH II increased with the advancement of ovarian development and maturation. In April, a single peak in GTH II level was noticed in the night at 20.6 h. In contrast, in May, June and July a biphasic pattern (ultradian rhythm) was noticed with two characteristic peaks, one in the photophase and the other in the scotophase. In addition, in May a statistically significant circadian rhythm in plasma GTH II was validated with a peak located at 20.1 h. These rhythms seem to have physiological significance. The ultradian rhythm with two peaks during the reproductively active phase appears to provide a suitable physiological milieu for the temporally different yet synchronous population of oocytes for the secretion of steroids. Thus, the observed temporal organization in GTH II may have physiological consequences leading to accomplishment of reproductive process at appropriate time of the year.     

Biological rhythms of cells of the lymphoid series in the thymus-dependent zone of lymph nodes

Biological rhythms of density in the lymphoid line cell populations (small and middle lymphocytes, immunoblasts, plasmoblasts, immature and mature plasmocytes) have been investigated in the thymus-dependent zone of the mouse axillary lymph nodes. The observations have been performed for 572 h. The amount of the cells is counted over the nodes of the morphometrical net. The results of the observations are treated by means of the spectral cosinor-analysis. Circadian or similar to it rhythmicity is peculiar for all the types of the cells, besides plasmoblasts. Within the ultradian range rhythmicity of the small lymphocytes, immunoblasts and immature plasmocytes is presented by two components, and that of the plasmoblasts--by one. The complexity of the rhythm components demonstrates that the course of the process during the observation time does not repeat strictly every day running, since periods of the circadian components somewhat fluctuate from 24 h, and periods of the ultracircadian ones--do not make multiple parts from the duration of the circadian periods. The time of picks and troughs of the amount of the cells during successive 24 h also do not coincide with each other, as the circadian and ultradian components interact. During one 24-hour's period the curve does not possess a regular sinusoid form and additional picks and troughs are revealed, since certain oscillations of a higher frequency participate in the process. These data are necessary to estimate reactions at T-immunological response.