Ultradian and circadian compartmentalization of respiratory and metabolic exchanges in small laboratory vertebrates

Carbon dioxide emission (VCO2) taken as an index of respiratory and metabolic exchanges, was continuously recorded during 4-30 consecutive days in 100 quail, 87 chicks, 347 rats, 665 mice and 70 guinea-pigs which were under controlled environmental parameters. Harmonic analysis, fast Fourier transform, chi-square periodograms, peak and trough intervals were computed with VCO2 values obtained with CO2 concentrations sampled every 20 min on the CO2 recordings. In LD 12:12 alternation, circadian rhythms were observed in all quail, chicks, rats and mice, but only in 80% of the guinea-pigs. Ultradian VCO2 rhythms, with periods which show statistically significant interspecies differences, were assessed. For each of the 5 species these computed periods, which were the same in LL and DD, were: 1.17 h for quail and chickens, 1.25 h for rats, 1.50 h for mice and 1.0 h for guinea-pigs. In LD 12:12 these periods were different during L and D in quail, chicks, rats and mice, but not in guinea-pigs. The amplitudes of these ultradian variations were, according to the species, 10-20% of their mean VCO2 levels. These ultradian rhythms persist in the absence (or masking) of circadian rhythms, e.g. in LD 12:12 in 20% of guinea-pigs and in LL in 87% of Japanese quail and in 23% of Sprague-Dawley rats. Moreover, these ultradian rhythms persist during starvation, locomotor activity restraint and ageing. These ultradian VCO2 cycles which are related to rest-activity variations appear to be basic physiological rhythms with a genetic origin.     

Species-specific evolution of the FcR family in endothermic vertebrates

In primates and rodents, the extended FcR family is comprised of three subsets: classical FcRs, structurally diverse cell surface receptors currently designated FCRL1-FCRL6, and intracellular proteins FCRLA and FCRLB. Using bioinformatic analysis, we revealed the FcR-like genes of the same three subsets in the genome of dog, another representative of placental mammals, and in the genome of short-tailed opossum, a representative of marsupials. In contrast, a single FcR-like gene was found in the current version of the chicken genome. This in silico finding was confirmed by the gene cloning and subsequent Southern blot hybridization. The chicken FCRL gene encodes a cell surface receptor with the extracellular region composed of four Ig-like domains of the D1-, D2-, D3-, and D4-subtypes. The gene is expressed in lymphoid and non-lymphoid tissues. Phylogenetic analysis of the mammalian and chicken genes suggested that classical FcRs, FCRLA, and FCRLB emerged after the mammalian-avian split but before the eutherian-marsupial radiation. The data obtained show that the repertoire of the classical FcRs and surface FcR-like proteins in mammalian species was shaped by an extensive recombination process, which resulted in domain shuffling and species-specific gain and loss of distinct exons or entire genes.     

The interaction of circahoral and circadian rhythms. A cybernetic model]

Parameters of ideal circadian cycle were compared with those of a circadian cycle composed of near-hour fluctuations. The integral cycle was optimized by the algorithm of matrix random search studying the approximation of its parameters to the ideal cycle after the phase shift of the latter. Computer calculations revealed a low efficiency of the search by fluctuation amplitude. The search by phase and frequency was effective in a narrow range of changes and needed time. The average level of fluctuations tuned up practically immediately: ideal and optimized curves coincided in all points. Examples of interactions between day, near-hour and shorter cycles are cited.     

Extraocular photoreception and circadian entrainment in nonmammalian vertebrates

In mammals both the regulation of circadian rhythms and photoperiodic responses depend exclusively upon photic information provided by the lateral eyes; however, nonmammalian vertebrates can also rely on multiple extraocular photoreceptors to perform the same tasks. Extraocular photoreceptors include deep brain photoreceptors located in several distinct brain sites and the pineal complex, involving intracranial (pineal and parapineal) and extracranial (frontal organ and parietal eye) components. This review updates the research field of the most recent acquisitions concerning the roles of extraocular photoreceptors on circadian physiology and behavior, particularly photic entrainment and sun compass orientation.     

Extraocular phototransduction and circadian timing systems in vertebrates

It is widely accepted that, for organisms with eyes, the daily regulation of circadian rhythms is made possible by light transduction through those organs. Yet, it has been demonstrated repeatedly in recent years that ocular light receptors that mediate vision, at least in mammals, are not the same photoreceptors involved in circadian regulation. Moreover, it has been recognized for many years that circadian regulation can occur in organisms without eyes. In fact, extraocular circadian phototransduction (EOCP) appears to be a phylogenetic rule for the vast majority of species. EOCP has been reported in every nonmammalian species studied to date. In mammals, however, the story is very different. This paper presents findings from studies that have examined specifically the capacity for EOCP in vertebrate species. In addition, the literature addressing noncircadian aspects of extraocular phototransduction is briefly discussed. Finally, possible mechanisms underlying EOCP are discussed, as are some of the implications of the presence, or absence, of EOCP across phylogeny. (Chronobiology International, 18(2), 137-172, 2001)     

Cellular and biochemical mechanisms underlying circadian rhythms in vertebrates

Circadian clocks organize neural processes, such as motor activities, into near 24-hour oscillations and adaptively synchronize these rhythms to the solar cycle. Recently, the first mammalian clock genes have been found. Unpredicted diversity in signaling pathways and clock-controlled gating of signals that modulate timekeeping has been discovered. A diffusible clock output has been found to control some behavioral rhythms. Consensus is emerging that circadian mechanisms are conserved across phylogeny, but that mammals have developed a great complexity of controls.     

Ultradian and circadian rhythms of sleep-wake and food-intake behavior during early infancy

The early development of sleep-wake and food-intake rhythms in human infants is reviewed. The development of a 24h day-night rhythm contains two observable developmental processes: the alterations in the periodic structure of behavior (decreased ultradian, increased circadian components) and the process of synchronization to external time (entrainment). The authors present the results of their studies involving 26 German children and compare them with previous investigations. In their research, it became evident that, during the first weeks of life, the time pattern of sleep-wake and food-intake behavior is characterized by different ultradian periodicities, ranging from 2h to 8h. In the course of further ontogenesis, the share of ultradian rhythms in the sleep-wake behavior decreases, while it remains dominant for food-intake behavior. The circadian component is established as early as the first weeks of life and increases in the months that follow. Besides, the authors' study supports the notion of broad interindividual variation in ultradian rhythms and in the development of a day-night rhythm. Examples of free-running rhythms of sleep-wake and food-intake behavior by various authors are strong indicators of the endogenous nature of the circadian rhythms in infants and show that the internal clock is already functioning at birth. It is still uncertain when the process of synchronization to external and social time cues begins and how differences in the maturation of perceptive organs affect the importance of time cues for the entrainment. Prepartally, the physiological maternal entrainment factors and mother-fetus interactions may be most important; during the first weeks of life, the social time cues gain importance, while light acts as a dominant “zeitgeber” at a later time only.     

Ultradian, circadian and seasonal variations of plasma progesterone and LH concentrations during the luteal phase

The circadian variations in plasma progesterone (P) and LH concentrations were investigated in six women, aged 23-40 years. All were studied in the mid-luteal phase (7 +/- 2 days after LH mid-cycle surge). Experiments were conducted in autumn and in spring. Blood samples were obtained every 15 min for 24 hr. Plasma P and LH concentrations were measured by RIA. Each subject's time-series was analysed using three methods; visual inspection (chronogram), spectral analysis to estimate component periods of rhythms (tau) and cosinor analysis to quantify the rhythms parameters. Marked temporal variations in plasma P concentration were observed in each subject. The maximal variations over a 24-hr period, ranged between 13-58.5 mmol/l. Differences related to sampling time were statistically validated by ANOVA (p less than 0.00001). Significant harmonic periods were detected by spectral analysis but differed among subjects. In all subjects but one, a circadian rhythm was detected. The acrophase location was similar (about 0700 hr) in the four subjects studied in autumn, but ranged from 1940 to 0320 hr in those studied in spring. An ultradian rhythm with tau = 8 hr was also validated in six time-series with similar acrophases (about 0200, 1000, and 1800 hr). Cosinor analysis of pooled data revealed that the 24-hr, 12-hr, and 8-hr rhythms were statistically significant (p = 0.001) in autumn. algebraic sum of these three cosine functions yielded a circadian waveform with peak-times occurring near 0300 and 1130 hr and a trough-time about 2200 hr. In spring, the circadian pattern appeared quite different, and peak-times were found near 0700 and 2000 hr, and trough-times near 0300 and 1500 hr. Furthermore, the 24-hr mean of P was higher in autumn (28.9 +/- 0.4 nmol/l) than in spring (17.2 +/- 0.4 nmol/l), p from ANOVA less than 0.00001. The evidence for a similar circadian LH pattern is not as strong. Seasonal, circadian and ultradian rhythms characterize the physiologic time structure of plasma P concentration in mid-luteal phase.     

The evolution of sleep circadian cycle and telencephalo-diencephalic interaction in vertebrates

The modem representations of wakefulness-sleep cycle evolution and the data about dynamics of reactivity of activating and inhibition neurotransmitter systems of the forebrain, converging in striatum, in cold- and warm-blooded vertebrates are considered. The data about dynamics of immune reactivity of vasopressin- and oxytocinergic cells of paraventricular and supraoptical hypothalamic nuclei is presented. On the basis of the obtained results, the idea of the leading role of telencephalo-diencephalic interactions in activation of somnogenic processes and their possible mechanisms is advanced.     

Amino acid composition in endothermic vertebrates is biased in the same direction as in thermophilic prokaryotes

Background

Among bacteria and archaea, amino acid usage is correlated with habitat temperatures. In particular, protein surfaces in species thriving at higher temperatures appear to be enriched in amino acids that stabilize protein structure and depleted in amino acids that decrease thermostability. Does this observation reflect a causal relationship, or could the apparent trend be caused by phylogenetic relatedness among sampled organisms living at different temperatures? And do proteins from endothermic and exothermic vertebrates show similar differences?

Results

We find that the observed correlations between the frequencies of individual amino acids and prokaryotic habitat temperature are strongly influenced by evolutionary relatedness between the species analysed; however, a proteome-wide bias towards increased thermostability remains after controlling for phylogeny. Do eukaryotes show similar effects of thermal adaptation? A small shift of amino acid usage in the expected direction is observed in endothermic ('warm-blooded') mammals and chicken compared to ectothermic ('cold-blooded') vertebrates with lower body temperatures; this shift is not simply explained by nucleotide usage biases.

Conclusion

Protein homologs operating at different temperatures have different amino acid composition, both in prokaryotes and in vertebrates. Thus, during the transition from ectothermic to endothermic life styles, the ancestors of mammals and of birds may have experienced weak genome-wide positive selection to increase the thermostability of their proteins.

     

Ultradian and circadian CO2 emission variations in nocturnal and diurnal animals exposed to a light stimulus

1. Carbon dioxide emission (VCO2) has been continuously recorded in three laboratory animal species (Sprague-Dawley rats, Japanese quail, Hartley guinea-pigs) which differ by their nocturnal and diurnal activities. A 100 lux stimulus has been delivered at various time intervals. 2. A regular alternation of 12, 3 or 1.5 hr light (L) and darkness (D) gives VCO2 circadian and ultradian rhythms of 24, 6 or 3 hr periods, respectively, in quail and rats. 3. Such circadian and ultradian LD rhythms are not induced in all guinea-pigs. 4. The amplitudes of the VCO2 responses are greatest at D----L when the animals have a maximum diurnal activity and at L----D when their maximum activity is nocturnal. 5. Interactions between circadian and ultradian rhythms are seen in all LD experiments, as well as in continuous light (LL) or continuous dark (DD). 6. No more well-marked or even inverted VCO2 responses to the light stimuli may occur after several days of exposure to these LD alternations.     

Quantifying light-dependent circadian disruption in humans and animal models

Although circadian disruption is an accepted term, little has been done to develop methods to quantify the degree of disruption or entrainment individual organisms actually exhibit in the field. A variety of behavioral, physiological and hormonal responses vary in amplitude over a 24-h period and the degree to which these circadian rhythms are synchronized to the daily light–dark cycle can be quantified with a technique known as phasor analysis. Several studies have been carried out using phasor analysis in an attempt to measure circadian disruption exhibited by animals and by humans. To perform these studies, species-specific light measurement and light delivery technologies had to be developed based upon a fundamental understanding of circadian phototransduction mechanisms in the different species. When both nocturnal rodents and diurnal humans, experienced different species-specific light–dark shift schedules, they showed, based upon phasor analysis of the light–dark and activity–rest patterns, similar levels of light-dependent circadian disruption. Indeed, both rodents and humans show monotonically increasing and quantitatively similar levels of light-dependent circadian disruption with increasing shift-nights per week. Thus, phasor analysis provides a method for quantifying circadian disruption in the field and in the laboratory as well as a bridge between ecological measurements of circadian entrainment in humans and parametric studies of circadian disruption in animal models, including nocturnal rodents.     

Temporal orientation: circadian clocks in animals and humans

By evolutionary adaptation to the regular day-night changes in environmental conditions, most organisms have acquired a temporal programme which matches the 24-h day. It rests on periodic processes which have the characteristics of self-sustaining oscillations, and which, in constant conditions, free-run with periods slightly deviating from 24 h. When entrained to 24 h, these circadian rhythms can be used by the organism as a clock for temporal orientation, e.g. in the occupation of one of the temporal niches provided by the environment or in the coordination of activities among individuals and species. The circadian clock also provides the basis for using the sun as a compass in spatial orientation, and for the recognition of the time of day as exemplified by the time sense in honey bees. Within the human organism, almost every function is modulated in a circadian fashion. Usually, all rhythms keep a distinct phase-relationship t to each other, providing a high degree of temporal order within the organism. When living in an isolation unit without time cues, most subjects develop free-running rhythms with periods close to 25 h in all functions. Sometimes, however, the sleep-wake cycle is lengthened to 30 h and more, or shortened to less than 22 h. In those instances, other rhythms such as that of body temperature become uncoupled from the sleep-wake cycle and continue to free-run with a period of about 25 h. During such states of ‘internal desynchronization’, subjects can be awake continuously for about 32 h, and sleep without interruption for 16 h. Nevertheless, they experience their ‘days’ as equal to 24 h. This is due to a profound change in time estimation: the intervals of 1-h estimates made by the subject are positively correlated with the duration of wakefulness. In contrast, short-time estimates (in the range of seconds) remain unaffected by desynchronization, indicating that short- and long-time estimates are based on different mechanisms.     

Ultradian and circadian activity-rest rhythms of preterm neonates compared to full-term neonates using actigraphic monitoring

During the first weeks of life, preterm neonates show fewer circadian rhythms in their physiological parameters than full-term neonates. To determine whether preterm neonates differ in their temporal adaptation to the daynight cycle from full-term neonates at the early age of 1 week, we compared activity-rest behavior of both groups. Activity-rest behavior of 10 neurologically healthy preterm neonates (born in 34th to 36th week of gestation) and 10 neurologically healthy full-term neonates (born in 37th to 42nd week of gestation) was monitored longitudinally for 8 successive days in the first 2 weeks of life. Actigraphy was used to register and display time patterns of activity and rest in neonates by using small actometers, which resemble a wristwatch. Nursing/feeding was recorded using the actometer's integrated event marker button. Recordings for preterm neonates were conducted in the hospital; recordings for full-term neonates were carried out in the hospital and in their homes. In addition to the actigraphic recordings, a standardized diary was kept regularly. To assess periodic characteristics, frequency components of activity-rest behavior were analyzed using fast Fourier transformation (FFT). Amounts of daily sleep time, nightly sleep time, and sleep time during 24h were compared. Nursing/feeding epochs were also analyzed for 5 preterm and 5 full-term neonates to compare their food intake behavior. The majority of preterm neonates showed a multitude of ultradian frequencies in their spectra. In contrast, several full-term neonates showed a distinct circadian frequency. In preterm neonates, average nightly sleep and average daily sleep of all recorded days were very similar, but after the fourth day of life, only average nightly sleep increased. In full-term neonates, average nightly and daily sleep time of all recorded days differed by about 1h. Average sleep time during 24h for preterm and full-term neonates was similar. Preterm neonates showed longer intervals between events of food intake than full-term neonates. The circadian peaks in the frequency spectra of full-term neonates may indicate the initial adaptation in the first week of life to a 24h day. This is in agreement with our results concerning the different durations of nightly and daily sleep. The increase in nightly sleep time of preterm neonates may be attributed to the progressing adaptation to a circadian activity- rest pattern. (Chronobiology International, 18(4), 697-708, 2001)     

Immunoglobulin-containing cells in chick embryo urogenital tissues: a new site for early B lineage cells in endothermic vertebrates.

We have employed histological and immunofluorescent staining procedures in order to characterize the distribution of mu + B lineage cells in tissue sections prepared from developing chicken embryo urogenital tissues (UGTs) between 14 and 21 days of incubation. B lineage cells were observed in tissue sections prepared from developing UGTs, especially the mesonephros and its associated tissue, throughout the sample period. The highest densities of mu + B lineage cells were observed in tissue sections prepared from 18 day embryos. The mu + UGT cells were distributed singly and in clusters in subcapsular regions and within the peritubular interstitium of the mesonephros. These observations (1) are consistent with those which suggest nonbursal site(s) for origin of cells in B lineage, (2) may help account for the varying effects of embryonic caudectomy performed between the second and third days of incubation and surgical bursectomy performed close to hatching, (3) may help provide new insights on the effects of sex hormones on B cell development, and (4) suggest fundamental ontogenetic and phylogenetic similarities between developing vertebrate immune systems.     

Chaos and Ultradian Rhythms

Systems in a chaotic state have apparently random outputs despite a simple underlying kinetic mechanism. For instance, the interaction of two coupled oscillators (the mitotic oscillator and the ultradian clock) can produce chaotic behaviour over a limited range of parameter values. Mathematical modelling shows that physiologically realistic characteristics are thereby exhibited. Cell division cycles of lower eukaryotes (protozoa and yeasts) show both deterministic and stochastic properties. Both dispersion of cell cycle times and quantized values can be generated, as a deterministic chaotic consequence of oscillator interaction rather than from noisy limit cycles. Advantages may stem from chaotic operation; a controlled chaotic attractor could provide multifrequency outputs that determine rhythmic behaviour on different time scales ( e.g. ultradian and circadian) with the facility for rapid state changes from one periodicity to another.