Effects of constant darkness and constant light on circadian organization and reproductive responses in the ram

The relationship between circadian rhythms in the blood plasma concentrations of melatonin and rhythms in locomotor activity was studied in adult male sheep (Soay rams) exposed to 16-week periods of short days (8 hr of light and 16 hr of darkness; LD 8:16) or long days (LD 16:8) followed by 16-week periods of constant darkness (dim red light; DD) or constant light (LL). Under both LD 8:16 and LD 16:8, there was a clearly defined 24-hr rhythm in plasma concentrations of melatonin, with high levels throughout the dark phase. Periodogram analysis revealed a 24-hr rhythm in locomotor activity under LD 8:16 and LD 16:8. The main bouts of activity occurred during the light phase. A change from LD 8:16 to LD 16:8 resulted in a decrease in the duration of elevated melatonin secretion (melatonin peak) and an increase in the duration of activity corresponding to the changes in the ratio of light to darkness. In all rams, a significant circadian rhythm of activity persisted over the first 2 weeks following transfer from an entraining photoperiod to DD, with a mean period of 23.77 hr. However, the activity rhythms subsequently became disorganized, as did the 24-hr melatonin rhythms. The introduction of a 1-hr light pulse every 24 hr (LD 1:23) for 2 weeks after 8 weeks under DD reinduced a rhythm in both melatonin secretion and activity: the end of the 1-hr light period acted as the dusk signal, producing a normal temporal association of the two rhythms. Under LL, the 24-hr melatonin rhythms were disrupted, though several rams still showed periods of elevated melatonin secretion. Significant activity rhythms were either absent or a weak component occurred with a period of 24 hr. The introduction of a 1-hr dark period every 24 hr for 2 weeks after 8 weeks under LL (LD 23:1) failed to induce or entrain rhythms in either of the parameters. The occurrence of 24-hr activity rhythm in some rams under LL may indicate nonphotoperiodic entrainment signals in our experimental facility. Reproductive responses to the changes in photoperiod were also monitored. After pretreatment with LD 8:16, the rams were sexually active; exposure to LD 16:8, DD, or LL resulted in a decline in all measures of reproductive function. The decline was slower under DD than LD 16:8 or LL.(ABSTRACT TRUNCATED AT 400 WORDS)     

模拟空间节律（L：D=0.75h：0.75h）对树qu...

It is known that contemporary space station revolves at the altitude of 400-500 km in the outer space. The present study was undertaken to investigate the effects of simulated space rhythm (L:D = 0.75 h:0.75 h) at this altitude on circadian rhythm in tree shrews (Tupaia belangeri chinensis) and the effects of endogenous sleep inducing neuropeptide Asp5-alpha-DSIP on the space-rhythm-entrained circadian rhythm. This primitive stock serves as one species of Tupaiidae and is a unique native of South China. Our previous studies have shown that this species showed striking differences in natural circadian rhythm between day and night (e.g. 3.03 degrees C of rectal temperature). Results showed that the above mentioned space rhythm (L:D = 0.75h:0.75h) could drastically disturb the inherent circadian rhythm of Tupaia belangeri chinensis. The maximal peak of motor activity dropped significantly in the morning (0600-1200) and a new enhanced peak (more than 20 times greater than that of the control) was found between 1800-2400, whereas the maximal trough of motor activity (2400-0600) remained basically unchanged. Concurrently, the total amount of 24-h motor activity was significantly decreased and the recovery after the cessation of space rhythm was slow. Experimental results also demonstrated that consecutive administration of Asp5-alpha-DSIP (30 micrograms/kg, i.p.) for 5 days (2 days before and 3 days during space rhythm) did not prevent the basic disturbance of circadian rhythm of Tupaia belangeri chinensis caused by the space rhythm (L:D = 0.75h:0.75h). Nevertheless, no decrease or even some enhancement of the total amount of 24th motor activity was observed during space rhythm or after its cessation.     

Seasonal changes in the circadian activity rhythm of light-dark (LD) and completely dark (DD) regimen in the mouse submandibular gland in the presence of light-dark (LD) and completely dark (DD) regimen

Influence of seasons on circadian activity changes and the influence of one and six weeks of DD upon these changes of acid phosphatase (AP) and beta-acetylglucosaminidase (AM) was studied in the submandibular gland of sexually mature male mice. Total enzyme activity was determined in tissue homogenates at four-hour intervals in March, June, October, and February under standard LD12/12 conditions and after one and six weeks of the DD regime. The rhythms were analysed according to cosinor method. Under constant lighting conditions the seasonal differences in the AM circadian activity rhythm were found. AP activity was considerably less influenced by seasonal changes. Both enzyme activity changes were independent of each other and each rhythm was differently influenced by DD. In the case of AM the most pronounced circadian activity changes had the highest amplitude and mesor occurred in summer. The strongest influence of DD upon this enzyme activity rhythm was observed in spring and summer especially after the first week, after six weeks the acrophase returned to the LD group value (spring). In autumn and winter the reaction to DD was different to that of summer and spring. For AP the circadian changes of activity were non-rhythmic in spring, whereas in all other seasons the acrophases occurred almost at the same time in the afternoon. In DD the activity rhythm significantly changed after six weeks. In all seasons, except spring the circadian rhythm of activity was not observed after six weeks of DD. An attempt was made to explain the observed results by the certain kind of genetic memory present in laboratory animals the neurohormonal system of which is influenced by seasonal changes.     

Splitting of the circadian rhythm of activity in hamsters: Effects of exposure to constant darkness and subsequent re-exposure to constant light

Summary The circadian rhythm of wheel running behavior was observed to dissociate into two distinct components (i.e. split) within 30 to 110 days in 56% of male hamsters exposed to constant light (Figs. 1–2). Splitting was abolished in all 16 animals that were transferred from constant light (LL) to constant darkness (DD) within 1–4 days of DD, and the components of the re-fused activity rhythm assumed a phase relationship that is characteristic of hamsters maintained in DD (Figs. 3–5). Re-fusion of the split activity rhythm was accompanied by a change in period (); in 14 animals increased while in the other 2 animals decreased after transfer to DD.After 10–30 days in DD, the hamsters were transferred back into LL at various time points throughout the circadian cycle. A few of these animals went through two or three LL to DD to LL transitions. The effect of re-exposure to LL was dependent on the phase relationship between the transition into LL and the activity rhythm. A rapid (i.e. 1–4 days) induction of splitting was observed in 7 of 9 cases when hamsters were transferred into LL 4–5 h after the onset of activity (Fig. 5). In the other 2 animals, the activity pattern was ultradian or aperiodic for 20 to 50 days before eventually coalescing into a split activity pattern. In contrast, transfer of animals (n = 13) from DD to LL at other circadian times did not result in the rapid induction of splitting and the activity rhythm continued to free-run with a single bout of activity (Fig. 5). Importantly, a transfer from DD to LL 4–5 h after the onset of activity did not induce splitting if the hamsters had not shown a split activity rhythm during a previous exposure to LL (n=10; Fig. 6).These studies indicate that transfer of split hamsters from LL to DD results in the rapid re-establishment of the normal phase relationship between the two circadian oscillators which underlie the two components of activity during splitting. In addition, there appears to be a history-dependent effect of splitting which renders the circadian system susceptible to becoming split again. The rapid re-initiation of the split condition upon transfer from DD to LL at only a specific circadian time is discussed in terms of the phase response curve for this species.Abbreviation PRC phase response curve This investigation was supported by NIH grants HD-09885 and HD-12622 from the National Institute of Child Health and Human Development and by a grant from the Whitehall FoundationRecipient of Research Career Development Award K04 HD-00249 from the National Institute of Child Health and Human Development     

Serotonin increases the phase shift of the circadian locomotor activity rhythm in mice after dark pulses in constant light conditions

Investigations on the effects of the 5-HT agonists and antagonists on the phase of the circadian locomotor activity rhythm of animals kept in constant light conditions (LL) are rare. Therefore the influence of R-(+)-OH-DPAT (5-HT1A receptors agonist) and metergoline (5-HT1/2/7 receptors antagonist) on the phase shift of the locomotor-activity rhythm alone and when combined with dark pulses in mice kept in LL are examined. The results indicate that 8-OH-DPAT administered independently at 12.00CT (Circadian Time) shifted the phase of the circadian rhythm and reinforced the effect of dark pulses on this parameter. 12.00CT was defined arbitrarily as the onset of locomotor activity in constant conditions. Metergoline diminished the phase shifts after dark pulses compared to 8-OH-DPAT. The influence of the serotonin agonist showed that serotonin can reinforce the phase shifting effect of the locomotor activity rhythm after dark pulses in LL condition.     

Entrainment of the circatidal swimming activity rhythm in the cumacean Dimorphostylis asiatica (Crustacea) to 12.5-hour hydrostatic pressure cycles

The cumacean Dimorphostylis asiatica (Crustacea) exhibits a circatidal swimming activity rhythm. The animals were exposed to a 12.5 hr sinusoidal change of hydrostatic pressure of 0.3 atm amplitude in the laboratory. Under constant dark conditions, most of the specimens were entrained to a daily bimodal swimming activity rhythm by the hydrostatic pressure cycle. A small number of individuals exhibited a unimodal daily rhythm, with no apparent entraining from the administered cycles. A marked feature was a flexible phase relationship between the entrained daily bimodal rhythm and the hydrostatic pressure cycles: the swimming activity of most of the specimens occurred around the pressure-decreasing phase, but for a small number of individuals it coincided with the pressure-increasing phase. Such flexibility suggests a weak entraining effect of hydrostatic pressure on the circatidal rhythm of this species. When exposed to 24 hr light-dark cycles and a hydrostatic pressure cycle simultaneously, the specimens exhibited a rhythmic activity entrained by the hydrostatic pressure cycle during the dark period, which closely resembles the temporal activity pattern of this species in the field. The light cycles entrained the swimming activity via direct inhibition and induction of activity (i.e., masking). Under light-dark conditions, the specimens exhibited activity on the pressure-increasing phase more frequently compared with specimens kept in constant darkness.     

Pineal N-acetyltransferase in chickens: Rhythm persists in constant darkness

Summary The daily rhythm in an enzyme, pineal serotonin N-acetyltransferase activity was studied in chickens kept in conditions of constant dark. The rhythm persisted and had a period length of approximately 24 hours which are characteristics of truly circadian rhythms. The detailed documentation of the rhythm (Fig. 1) shows the shape of the oscillation and clear anticipation of the time of lights on. Both the rise and fall of the rhythm occur without stimulation by light or dark.We thank Robert Weiner, William Finlay, Jay Freedman, Kathy Hall, Sylvia Goddes, Dr. S. K. Roberts, and the Temple University Computer Activity. Supported by NSF grant GB-43215, NSF Institutional Award, and Temple Grant-in-Aid to S. Binkley.     

Effect of constant and fluctuating temperature on the circadian foraging rhythm of the red imported fire ant,Solenopsis invicta Buren (Hymenoptera: Formicidae)

Understanding circadian foraging rhythms activity of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae) foragers at different temperatures is an important step towards developing control measures in Integrated Pest Management (IPM) programs. In this study, the circadian foraging rhythm activities of S. invicta foragersat different temperature were investigated under laboratory and field conditions. Results indicated that the foraging activity increased after sunrise, and maximum foraging occurred at 14:00 (foraging rate was 69.22 ± 0.57 and 72.58 ± 1.15 foragers/min in the first and second year, respectively) in the tea fields of Guangzhou during autumn. Furthermore, foragers demonstrated circadian rhythms and exhibited a unimodal after 24 h. A significant correlation was found between foraging activity and temperature. S. invicta colonies were active at moderate soil temperatures (approximately 26.65 °C to 29.24 °C). The preferred temperature of the colonies was 26 °C, followed by 22 °C and 18 °C in the laboratory. The individual S. invicta activity was maximum at 17:00 (18.67 ± 1.66 times /10 min) and minimum at 5:00 (8.33 ± 2.51 times/10 min) at 26 °C. The fluctuating temperature had a significant impact on individual locomotor activity (r = 0.8979, P < 0.01) but did not alter the rhythm activity. Our results demonstrated that temperature might play an important role in circadian foraging rhythms activity of S. invicta. These results may have implications for the development of more effective fire ant management strategies.     

Internal desynchronization of the circadian activity and feeding rhythm in an owl monkey (Aotus lemurinus griseimembra): a case study

In the free-running circadian locomotor activity rhythm of a 7-year-old male owl monkey (Aotus lemurinus griseimembra) kept under constant light and climatic conditions (LL 0.2 lux, 25°C ± 1°C, 60 ± 5% relative humidity [RH]), a second rhythm component developed that showed strong relative coordination with the free-running activity rhythm of 24.4h and a 24h rhythm. The simultaneously recorded feeding activity rhythm strongly resembled this rhythm component. Therefore, it seems justified to infer that there was an internal desynchronization between the two behavioral rhythms or their circadian pacemakers, that is, between the light-entrainable oscillator located in the suprachiasmatic nuclei (SCN) and a food-entrainable oscillator located outside the SCN. This internal desynchronization may have been induced and/or maintained by a zeitgeber effect of the (irregular) 24h feeding schedule on the food-entrainable oscillator. The weak relative coordination shown by the activity rhythm indicates a much weaker coupling of the light-entrainable oscillator to the food-entrainable oscillator than vice versa. (Chronobiology International, 17(2), 147-153, 2000)     

Towards a characterization of the locomotor activity rhythm of the desertic species Porcellio olivieri (Crustacea,Isopoda) from Gabes (South of Tunisia)

The locomotor activity rhythm of the isopod, Porcellio olivieri, was investigated in Gannouch site in the south of Tunisia. The rhythm was monitored under constant temperature in individual animals in winter under two simultaneous regimens: the light–dark (LD) cycle and the continuous darkness (DD). Results revealed that whatever regimens, actograms, and mean activity curves showed that specimens of P. olivieri concentrated their activity during the experimental and subjective night. The species exhibited a locomotor rhythm period significantly shorter under LD (T = 23h13 ± 0h44) than DD (τ = 24h28 ± 0h58). However, the locomotor activity rhythm was less stable and the individuals were significantly more active under entraining conditions than constant darkness. The activity pattern of this species will be discussed as an adaptative strategy to respond to environmental conditions.     

Chronobiological background to cathemerality: circadian rhythms in Eulemur fulvus albifrons (Prosimii) and Aotus azarai boliviensis (Anthropoidea)

Cathemeral activity, in which the animals' motor activity is almost evenly distributed throughout the dark and the light portion of the day, has been described in various lemur genera (Eulemur, Hapalemur) and in the owl monkey Aotus azarai of the Argentinean Chaco. Proximate and ultimate factors responsible for this behaviour are still being debated. However, the chronobiological background of the behaviour has largely been ignored. We studied E. fulvus albifrons and A. a. boliviensis under controlled laboratory conditions to assess whether their activity rhythm is endogenously regulated by a circadian timing system that obeys general rules found in other mammals, or whether there are characteristic differences. To this end, we carried out long-term activity recordings on individuals of both subspecies kept under constant light and various light-dark cycles (LDs) using a PC-controlled electro-acoustic device in combination with telemetric body temperature measurements. Both subspecies developed free-running circadian activity and body temperature rhythms with periods deviating from 24 h in constant light, and LDs turned out to be the most efficient Zeitgeber synchronizing this endogenous rhythmicity to the external 24-hour day. The luminosity prevailing during the dark time of the LD had a decisive effect on levels of activity in the lemurs and induced strong masking effects on their circadian activity pattern. The results indicate that, from a chronobiological viewpoint, both species should be considered as dark active primates. Their diel activity rhythm is regulated by a normally responding circadian timing system and strong activity inhibiting or enhancing direct effects of light intensity. Thus, hypotheses on proximate and/or ultimate factors of cathemerality in primates must also consider its circadian background.     

Evidence for an endogenous circadian rhythm of feeding in the trout (Oncorhynchus mykiss)

Abstract

The possible endogenous circadian rhythm in the feeding activity of rainbow trout (Oncorhynchus mykiss) was investigated using individual fish previously trained for self‐feeding. Under LD 12:12 conditions, the fish showed a diurnal behaviour, in many cases with a feeding rhythm with two main peaks of food demand at dawn and dusk, with an 8h interval of low feeding activity, and the actograms showed an expected 24 h rhythm. Fish kept under constant conditions (L : L, 15°±0.5°C), showed free‐running feeding activity for about 12 days. Food demands were concentrated at dawn, with a periodogram of 25.3 hour, under continuous environmental conditions. Results showed evidence for the endogenous origin of the circadian rhythm of feeding in this species.     

INTERNAL DESYNCHRONIZATION OF THE CIRCADIAN ACTIVITY AND FEEDING RHYTHM IN AN OWL MONKEY (AOTUS LEMURINUS GRISEIMEMBRA): A CASE STUDY

In the free-running circadian locomotor activity rhythm of a 7-year-old male owl monkey (Aotus lemurinus griseimembra) kept under constant light and climatic conditions (LL 0.2 lux, 25°C ± 1°C, 60 ± 5% relative humidity [RH]), a second rhythm component developed that showed strong relative coordination with the free-running activity rhythm of 24.4h and a 24h rhythm. The simultaneously recorded feeding activity rhythm strongly resembled this rhythm component. Therefore, it seems justified to infer that there was an internal desynchronization between the two behavioral rhythms or their circadian pacemakers, that is, between the light-entrainable oscillator located in the suprachiasmatic nuclei (SCN) and a food-entrainable oscillator located outside the SCN. This internal desynchronization may have been induced and/or maintained by a zeitgeber effect of the (irregular) 24h feeding schedule on the food-entrainable oscillator. The weak relative coordination shown by the activity rhythm indicates a much weaker coupling of the light-entrainable oscillator to the food-entrainable oscillator than vice versa. (Chronobiology International, 17(2), 147–153, 2000)     

Functional diversities of two activity components of circadian rhythm in genetical splitting mice (CS strain)

CS mice, an inbred strain, showed two distinctive characteristics in the circadian rhythm of locomotor activity: (1) large variation in the freerunning period, and (2) spontaneous rhythm splitting under continuous darkness. In the splitting rhythm there was a positive correlation between the freerunning period of the evening component and the activity time of the morning component. The phase-shifting effect of a 15-min light pulse was examined on the two activity components of the splitting rhythm. There were significant differences in the amount of light-induced phase response between the two components. A light pulse during the late subjective night induced a phase advance shift only in the morning component, while a light pulse during the early subjective night induced a phase delay shift only in the evening component. These results indicate functional diversities of the two activity components in the circadian locomotor rhythm of CS mice, and suggest that the circadian system in CS mice consists of two mutually coupled oscillators which have different circadian periods and different responsiveness to light. The CS mouse is a useful model to explore a genetic background of oscillator coupling in the circadian system of nocturnal rodents. Accepted: 19 November 1998     

A comparison of the activity patterns of the sand goby Pomatoschistus minutus (Pallas) from areas of different tidal range

Sand gobies ( Pomatoschistus minutus ) collected from beaches with a large tidal range (Scotland) exhibit a circatidal rhythm of activity in constant conditions in the laboratory. There is no endogenous circadian component to the rhythm. The phasing of the rhythm is such that peak activity occurs at the predicted time of ebb tide. Light-dark (LD) cycles applied in the laboratory have a marked effect on activity greatly enhancing it at night so that the original tidal rhythm becomes nocturnal. Some evidence was obtained that LD cycles can entrain a weak nocturnal circadian rhythm in fish removed from tidal conditions. Contrary to expectations, some fish from beaches with a small and unpredictable tidal range (Oslofjord, Norway) also show a weakly persistent circatidal thythm without an endogenous circadian component, but great variability was noticed between individual fish. Laboratory LD cycles did not entrain a persistent circadian rhythm in the fish from the Oslofjord.     

Regulation of the pineal melatonin concentration in the rat (Rattus norvegicus) and in the Djungarian hamster (Phodopus sungorus)

1. Rapid changes of melatonin concentration in pineals of rats (Rattus norvegicus) after isoproterenol administration or in the course of a day were accompanied by parallel changes of N-acetyltransferase but not of hydroxyindole-O-methyltransferase activity. 2. In pineals of Djungarian hamsters (Phodopus sungorus) the evening and morning melatonin changes were also parallel with N-acetyltransferase changes. 3. In both species, melatonin concentration during its rise had approached the maximum earlier than N-acetyltransferase reached its highest activity. 4. It is proposed that the N-acetyltransferase rhythm drives the melatonin rhythm both in rats and in Djungarian hamsters; the maximum melatonin production may, however, depend also on hydroxyindole-O-methyltransferase activity and substrate concentration.     

Characteristics of the circadian activity rhythm in common marmosets (Callithrix j. jacchus)

The circadian activity rhythm of the common marmoset, Callithrix j. jacchus was investigated by long-term recording of the locomotor activity of 15 individuals (5 males, 10 females) from 1.5 to 8 years old, both under constant illumination and under LD 12:12. The mean period of the spontaneous circadian rhythm was 23.2 ± 0.3 h. Neither sex-specific differences nor a systematic influence of light intensity on the spontaneous period were observed, but the period was dependent on the duration of the trial and on the age of the individual. Due to the short spontaneous period, in LD 12:12 there was a distinct advance of the activity phase with respect to the light time and a masking of the true onset of activity by the inhibitory direct effect of low light intensity during the dark time. After an 8 h delay shift of the LD 12:12, re-entrainment of the circadian activity rhythm required an average of 6.8 ± 0.7 days; the average re-entrainment time after an 8 h phase advance of the LD cycle was 8.6 ± 1.3 day. This directional effect is ascribed to characteristics of the phase-response curve. No ultradian components were observed, either in the LD-entrained or the free-running circadian activity rhythm.     

Effect of constant light on the circadian rhythm in locomotor activity in intact or eyelids-removed rats

According to the Aschoff's role, exposure to continuous light (LL) results in the elongation of the free-running period of the rat circadian rhythm. However, the LL may not always mean the constant intensity of the light for the suprachiasmatic nucleus, since the rat may regulate the contrast of the illumination by their eyelids which are closed during the sleep phase. In this study, the surgical removal of the eyelids under the LL caused arrhythmicity of the locomotor activity in 7 of 10 rats. The remaining 3 rats maintained the free-running rhythm after the removal of the eyelids. These results suggest that constant light may affect the free-running rhythm of the rat with or without eyelids in the different manner.     

Temperature and light effects on the circadian rhythm and locomotory activity of the plains garter snake (Thamnophis radix hayendi)

Abstract

The Locomotory activity of the Plains Garter snake was determined under L/D: 12/12 conditions at five constant temperatures and three light intensities during the light period. The snakes were diurnal at low temperatures with nocturnal activity increasing in various amounts at higher temperatures. The different light intensities had relatively small effects on the activity rhythm.

Activity was recorded under four constant light conditions at five constant temperatures and the characteristics of the free‐running rhythm measured. A comparison of the characteristics of the free‐running rhythm to Aschoff's circadian rule indicates that this snake is an exception to this rule.

Increase light intensity decreased total activity under all conditions. Under a L/D: 12/12 cycle the decrease in activity was more pronounced during the dark period than the light period.

It is suggested that crepuscular or nocturnal activity shown by snakes at high temperatures may be an effect the temperature level has on the biological clock and activity controlling mechanisms rather than temperature selection by the snake.