Masking by light in hamsters with SCN lesions

Inhibition of wheel running by light (masking) was investigated in Syrian hamsters with suprachiasmatic nucleus or sham lesions. Approximately 90% of the wheel revolutions made by hamsters with complete suprachiasmatic nucleus lesions, as judged by histology and power spectrum analysis of their wheel running, occurred during the dark phases of an ultradian light-dark cycle (3.5 h light, 3.5 h dark). This was demonstrated for two illumination levels (380 lx and 6 lx). Similar results were obtained with sham-operated animals. In further tests, the hamsters with lesions also retained a strong preference for the dark side of a box divided into dark and light sides. These results demonstrate that the suprachiasmatic nucleus is not necessary for masking by light or the preference for a dark over a light compartment. Evidently the direct effects of light can substitute for the endogenous control by the suprachiasmatic nucleus to maintain appropriate behaviour in time and space. Accepted: 30 January 1999     

Enhanced masking response to light in hamsters with IGL lesions

Syrian hamsters with intergeniculate leaflet or sham lesions were given tests with a series of light pulses of gradually decreasing intensities. The light pulses were given early in the night, at zeitgeber time 14–15. The amount of wheel running during the pulses was compared to that in the same hour on a night with no light pulses. Hamsters with intergeniculate leaflet lesions showed a significantly greater suppression of their wheel running in response to light than the sham-lesioned animals. The lesioned animals also had larger negative phase angles of entrainment to the 14:10-h light-dark cycle than sham-operated controls. However, phase shifting in response to light pulses at either zeitgeber time 14 or 18 was not significantly altered by the lesions. Preferences for spending more time in a dark than a light area were not abolished by the lesions. It is concluded that the intergeniculate leaflet in the Syrian hamster cannot be of paramount importance for masking of locomotor activity by light but may play a modulating role. Accepted: 30 January 1999     

Complete suprachiasmatic lesions eliminate circadian rhythmicity of body temperature and locomotor activity in golden hamsters

The effects of suprachiasmatic and control lesions on the circadian rhythms of locomotor activity and body temperature were studied in golden hamsters (Mesocricetus auratus) maintained in constant light as well as constant darkness. Large suprachiasmatic lesions, but not control lesions, eliminated circadian rhythmicity in locomotor activity as well as in body temperature. Analysis of the robustness of the rhythms of locomotor activity and body temperature in unlesioned and lesioned animals suggests that, because body temperature rhythmicity is more robust than locomotor rhythmicity, lesions that spare a small number of suprachiasmatic cells might abolish the latter but not the former. Our results do not support the hypothesis that the body temperature rhythm is controlled by a circadian pacemaker distinct from the main pacemaker located in the suprachiasmatic nuclei.     

Seasonality of circadian locomotor activity in an insect

Daily periodic locomotor activity of Carabus auronitens was recorded in a climate-constant laboratory with the animals exposed to naturally changing photoperiods. Most actograms exhibit directed seasonal variations of duration and phase position of daily activity. Seasonal locomotor activity starts in early spring (following dormancy) on a low daily level, first being confined to a short time span around dusk (and even shorter around dawn). In the course of season, the daily onsets of activity become closely related with sunset and the duration of daily activity is steadily extended with both parts of the bimodal phase fusing to a common, unimodal activity band by late spring. Subsequently, it is further extended into forenoon, until in summer (shortly before aestivation), spontaneous phase inversion turns activity periodicity from nocturnality into diurnality within 1 day. Such seasonal variations are paralleled by changes in the precision of synchronization of the individuals' activity rhythms to the entraining light/dark cycle. No geographical differences were detected. The results support the idea of the circadian clock as a system of two dynamically coupled physiological oscillators that invert their phase relation as soon as the natural dark phase falls short of some minimum-tolerable night length.Abbreviations LD light/dark cycle - nLD natural light/dark cycle - DD constant darkness - endogenous period length - SS sunset - SR sunrise - PRC phase-response curve     

Feeding effects on daily locomotor activity in Carabus auronitens

Effects of food supply on daily periodic locomotor activity in the ground beetle Carabus auronitens were recorded in the laboratory with the animals exposed to natural daylight and photoperiod. Inspection of individual actograms exhibits that in most cases feeding leads to strongly reduced and more or less irregularly distributed activity for a period of up to 4 days. In terms of quantitative parameters, the animals respond to food supply by an immediate reduction of mean amounts of daily activity. This is achieved through the reduction of duration (as a consequence of reduced burstlengths instead of burstnumber) and, less distinctly, the reduction of intensity. The initial levels of parameters are reattained after 4 days. Direct comparison by means of standardized (i.e. level-independent) relative parameter functions demonstrates close concordance between the amount and duration of daily activity; however, distinct synchronous change of intensity is apparent as a consequence of food supply only. This means that any variation of activity amounts is due to changing duration rather than to intensity variation. In analogy to the feeding-dependent parameter dynamics, also the proportion of animals with strong synchronization to the natural light/dark cycle is considerably reduced by food supply. The feeding-dependent reduction of activity is paralleled by temporarily reduced activity of proteolytic enzymes in the beetles. It is argued that, for the purpose of energy saving, the animals will remain inactive as long as reduced enzyme activity inhibits further consumption and digestion of food.Abbreviations DD constant darkness - LD light/dark cycle - nLD natural light/dark cycle - endogenous periodlength     

Different circadian pacemakers control feeding and locomotor activity rhythms in European starlings

Summary In higher organisms, many physiological and behavioral functions exhibit daily variations, generated by endogenous circadian oscillators. It is not yet clear whether all the various rhythms that occur within an individual depend on one and the same pacemaker or whether different pacemakers are involved. To examine this question, the feeding and perch-hopping rhythms were measured in European starlings (Sturnus vulgaris) under light-dark cycles and continuous dim light. In dim light, the internal phase relationship between the feeding and perch-hopping rhythms changed systematically as a function of the circadian period, and the two rhythms could even dissociate and show different circadian periods in individuals with extremely long or extremely short circadian periods. Moreover, in some birds kept on lowamplitude light-dark cycles, the rhythm of feeding was synchronized 180° out of phase with the rhythm of locomotor activity. These results strongly suggest that in the European starling the feeding and locomotor activity rhythms are controlled by separate circadian pacemakers.     

Time-dependent effects of leptin on food intake and locomotor activity in goldfish

The present study investigates the possible circadian dependence of leptin effects on food intake, locomotor activity, glycemia and plasma cortisol levels in goldfish (Carassius auratus). Fish were maintained under 12L:12D photoperiod and subjected to two different feeding schedules, one group fed during photophase (10:00) and the other one during scotophase (22:00). Leptin or saline were intraperitoneally injected at two different times (10:00 or 22:00), coincident or not with the meal time. To eliminate the entraining effect of the light/dark cycle, goldfish maintained under 24 h light (LL) were fed and leptin-injected at 10:00. A reduction in food intake and locomotor activity and an increase in glycemia were found in goldfish fed and leptin-injected at 10:00. No significant changes in circulating cortisol were observed. Those effects were not observed when leptin was administered during the scotophase, regardless the feeding schedule; neither in fish maintained under LL, suggesting that a day/night cycle would be necessary to observe the actions of leptin administered during the photophase. Changes in locomotor activity and glycemia were only observed in goldfish when leptin was injected at daytime, coincident with the feeding schedule, suggesting that these leptin actions could be dependent on the feeding time as zeitgeber. In view of these results it appears that the circadian dependence of leptin actions in goldfish can be determined by the combination of both zeitgebers, light/dark cycle and food. Our results point out the relevance of the administration time when investigating regulatory functions of hormones.     

Pharmacological effects of vinorelbine on body temperature and locomotor activity circadian rhythms in mice

The effects of vinorelbine (VRL) on the circadian rhythms in body temperature and locomotor activity were investigated in unrestrained B6D2F₁ mice implanted with radio-telemetry transmitters. A single intravenous VRL dose (24 or 12 mg/kg) was given at 7 h after light onset (HALO), a time of high VRL toxicity, and resulted in transient suppression of temperature and activity circadian rhythms in mice kept in light-dark (LD) 12h:12h. Such suppression was dose-dependent. It occurred within 1-5 d after VRL dosing. Recovery of both rhythms was partially complete within 5 d following the high dose and within 2 or 3 d after the low dose and was not influenced by suppression of photoperiodic synchronization by housing in continuous darkness. Moreover, VRL induced a dose-dependent relative decrease in amplitude and phase shift of the temperature circadian rhythm. The mesor and amplitude of the activity rhythm were markedly reduced following the VRL administration. The relevance of VRL dosing time was studied in mice housed in LD 12h:12h. Vinorelbine was injected weekly (20 mg/kg/injection) for 3 wk at 6 or 18 HALO. Vinorelbine treatment ablated the rest-activity and temperature rhythms 3-6 d after each dose, with fewer alterations after VRL dosing at 18 HALO compared to 6 HALO, especially for the body temperature rhythm. There was at least partial recovery 1 wk after dosing, suggesting the weekly schedule of drug treatment is acceptable for therapeutic purposes. Our findings demonstrate that VRL can transiently, yet profoundly, alter circadian clock function. Vinorelbine-induced circadian dysfunction may contribute to the toxicokinetics of this and possibly other anticancer drugs.     

Feeding entrainment of locomotor activity rhythms in the goldfish is mediated by a feeding-entrainable circadian oscillator

Periodic food availability can act as a potent zeitgeber capable of synchronizing many biological rhythms in fishes, including locomotor activity rhythms. In the present paper we investigated entrainment of locomotor rhythms to scheduled feeding under different light and feeding regimes. In experiment 1, fish were exposed to a 12:12?h light/dark cycle and fed one single daily meal in the middle of the light phase. In experiment 2, we tested the effect of random versus scheduled feeding on the daily distribution of activity. During random feeding, meals were randomly scheduled with intervals ranging from 12 to 36?h, while scheduled feeding consisted of one single daily meal set in the middle of the light or dark phase. Finally, in experiment 3, we studied the synchronization of activity rhythms to feeding under constant darkness (DD) and after shifting the feeding cycle by either advancing or delaying the feeding cycle by 9?h. The results revealed that goldfish synchronized to feeding, overcame light entrainment and significantly changed their daily distribution of activity according to their feeding schedule. In addition, the daily activity pattern modulated by feeding differed between layers: a peak of activity being noticeable directly after feeding at the bottom, while an anticipatory behaviour was obvious at the surface of the tank. Under DD and no food, free-running rhythms averaging 25.5?± 1.9?h (mean?±?SD) were detected. In conclusion, some properties of feeding entrainment (e.g. anticipation of the feeding time, free-running rhythms following termination of periodic feeding, and the stability of ø after shifting the feeding cycle) suggested that goldfish have (a) separate but tightly coupled light- and food-entrainable oscillators, or (b) a single oscillator that is entrainable by both light and food (one synchronizer being eventually stronger than the other).     

Further characterization of the phenotype of mCry1/mCry2-deficient mice

Mice lacking cryptochromes (mCry1^-/- mCry2^-/-) were kept in a 16h light, 8h dark, light-dark (16:8 LD) cycle and were given additional pulses of light of different brightness, starting 2h after dark onset and lasting for 1h. The suppression of wheel running during these light pulses (i.e., masking) was compared to that of wild types. No evidence of any decrement in the masking response to light was detected. As well as studying masking, minor bouts of activity occurring in the main light portion of a light-dark cycle were quantified. One possible explanation of such predark activity is that some damped endogenous process is spared in mCry1/mCry2 double-knockout mice. (Chronobiology International, 18(4), 613-625, 2001)     

Circadian rhythm of spontaneous locomotor activity in the bed bug, Cimex lectularius L.

Bed bugs must avoid detection when finding hosts and returning to hidden harborages. Their stealthy habits include foraging when hosts are asleep. Characteristics of spontaneous locomotor activity rhythm of bed bugs with different feeding histories were studied. In the absence of host stimuli, adults and nymphs were much more active in the dark than in the light. The onset of activity in the scotophase commenced soon after lights-off. The free-running period (τ) for all stages was longer in continuous darkness (DD) than in continuous light (LL). The lengthening of τ in DD is an exception for the circadian rule that predicts the opposite in nocturnal animals. Activity in all stages was entrained to reverse L:D regimes within four cycles. Short-term starved adults moved more frequently than recently fed adults. While bed bugs can survive for a year or more without a blood meal, we observed a reduction in activity in insects held for five weeks without food. We suggest that bed bugs make a transition to host-stimulus dependent searching when host presence is not predictable. Such a strategy would enable bed bugs to maximize reproduction when resources are abundant and save energy when resources are scarce.     

Irradiance dependency of UV-A induced phase shifts in the locomotor activity rhythm of the field mouse Mus booduga

In the nocturnal field mouse Mus booduga, the responsiveness of the circadian system to UV-A light of 2.5 W/m² and 30 minutes duration is known to be phase dependent. The results of our experiments indicate that the phase shifts evoked by UV-A at the two phases, CT14 (circadian time 14) and CT20 increases nonlinearly with irradiance. (Chronobiology International, 17(6), 777-782, 2000)     

Dissociation between the circadian rhythm of locomotor activity and the pineal clock in the Japanese newt

The circadian locomotor activity rhythm of the Japanese newt has been thought to be driven by a putative brain oscillator(s) subordinate to the pineal clock. The existence of mutual coupling between the pineal clock and the brain oscillator(s) in vivo was examined. We covered the newt's skull with aluminum foil and simultaneously reversed the light-dark cycle, thereby allowing the pineal organ to be exposed to constant darkness while the rest of the animal was exposed to the reversed light-dark cycle. In control animals, whose heads were covered with transparent plastic, the rhythm of synaptic ribbon number in the pineal photoreceptor cells was entrained to the reversed light-dark cycle. Rhythms from newts whose heads were shielded, however, were similar to those observed in the unoperated newts kept under constant darkness. The locomotor activity rhythms of both head-covered animals and control animals were entrained to the reversed light-dark cycle. These data suggest that extrapineal photoreception can entrain the putative brain oscillator(s), but not the pineal clock. Thus, at least in an aspect of photic entrainment, there seems to be little or no mutual coupling between the pineal clock and the putative brain oscillator(s) in the circadian system of the Japanese newt.Abbreviations LD light-dark - DD constant darkness - SCN suprachiasmatic nucleus - SR synaptic ribbon     

Masking: History, Definitions, and Measurement

Masking and entrainment are two different processes that result in rhythms in physiology and behavior. Considering its functional importance, lamentably little study has been devoted to masking compared to that lavished on entrainment. This paper discusses the origin and definition of terminology (positive and negative masking, paradoxical masking) and methods of quantifying masking. How masking is scored can greatly influence the interpretation of results.     

Pineal and retinal melatonin is involved in the control of circadian locomotor activity and body temperature rhythms in the pigeon

Summary Although pinealectomy or blinding resulted in loss of the clarity of the free-running rhythm of locomotor activity and body temperature and reduced the peak level of circulating melatonin rhythms to approximately a half in intact pigeons, neither pinealectomy nor blinding abolished any of these rhythms. However, when pinealectomy and blinding were combined, the rhythms of locomotor activity and body temperature disappeared in prolonged constant dim light, and melatonin concentration was reduced to the minimum level of detection. In order to examine the role of melatonin in the pigeon's circadian system, it was administered either daily or continuously to PX + EX-pigeons in LLdim. Daily administration of melatonin restored circadian rhythms of locomotor activity which entrained to melatonin injections, but continuous administration did not induce any remarkable change of locomotor activity. These results suggest that melatonin synthesized in the pineal body and the eye contributes to circulating melatonin and its rhythmicity is important for the control of circadian rhythms of locomotor activity and body temperature in the pigeon.Abbreviations LD Light-dark - LLdim constant dim light - LLbright constant bright light - PX pinealectomy - EX blinding - SCN suprachiasmatic nucleus     

Phase-shifting effects of dusklike and dawnlike light pulses on the circadian activity rhythms of Syrian hamsters

This study tested whether light pulses with a dusklike offset or a dawnlike onset caused phase shifts of different sizes in the circadian wheel-running activity of Syrian hamsters, Mesocricetus auratus. Six experiments were conducted, each with 30 hamsters; the hamsters received first one type of pulse and then the other type a few weeks later, allowing a paired comparison. The six experiments represented the combination of two maximum light intensities (150 and 250 lux) and three zeitgeber times (ZTs) at which the pulses were given (ZT13.5, ZT14.5, and ZT20). Pulses were 30 minutes long, a relatively short duration to minimize circadian time effects. Aschoff's type II method of measuring phase shifts was used. In none of the six experiments did a two-tailed paired t test detect a significant difference in the size of phase shifts caused by dusklike versus dawnlike pulses. A three-way analysis of variance (ANOVA) on the combined data from all six experiments (with pulse type, pulse intensity, and ZT as factors) also failed to detect a significant effect of pulse type. Statistical power was calculated and found to be reasonably good. These negative results are in line with those of a previous study in which a different methodology was used. (Chronobiology International, 18(3), 413-421, 2001)     

Pineal-dependent locomotor activity of lamprey,Lampetra japonica,measured in relation to LD cycle and circadian rhythmicity

Summary Locomotor activity of the river lamprey, Lampetra japonica, was investigated under a light-dark (LD 1212) cycle and under continuous dark conditions. Intact lampreys were entrained to the light:dark cycle. They were active mainly in the early half of the dark period and inactive in light period. The light:dark entrainment continued in 72.7% of lampreys after the removal of bilateral eyes, but additional pinealectomy made the entrainment disappear in all lampreys. When lampreys were pinealectomized with their eyes intact, light: dark entrainment was abolished in most cases. The results indicate that the pineal organ of the lamprey is a photoreceptive organ responsible for synchronizing locomotor activity to LD cycle. Under continuous dark conditions, the locomotor activity began to free-run with a period of 21.3 ± 0.9 h (mean ± SD, n = 53). This circadian rhythmicity was not affected by the removal of lateral eyes but was abolished by pinealectomy. The pineal organ appears to function as an oscillator, or as one of the oscillators, for the circadian locomotor rhythm of lampreys.Abbreviations DD continuous dark - LD light:dark     

Circadian Rhythms and Masking: An Overview

Masking, as is well known, enables an organism to act immediately and in an appropriate way to changes of the environment, integrating with internally produced rhythmicity. It now appears that masking can be used to cover a far wider range of problems than was originally intended. To separate masking effects from the effects due to an internal oscillator, several techniques have been used. Such protocols, however, like the constant routine protocol, often replace one form of masking by another. The situation becomes even more complex when one realizes that the output of an internal oscillator modifies the input. The question might be asked whether it is possible to study the properties of the internal oscillator in vivo at all. This article attempts to produce a framework for future discussions.     

Diazepam affects both level and amplitude of rat locomotor activity rhythm but has no effect on core body temperature

The present study investigates the effects of a chronic administration of diazepam, a benzodiazepine widely used as an anxiolytic, on locomotor activity and body core temperature rhythms in male Wistar rats housed under 12∶12 light∶dark (LD) cycle conditions. Diazepam was administered subcutaneously for 3 wks in a dosage of 3 mg/kg body weight/day, 1 h before the onset of darkness. Diazepam increased the level of locomotor activity from the first day until the end of treatment, and also increased the amplitude of the activity circadian rhythm, but only on the third wk of treatment. Diazepam exerted no effects on the length of the period and did not affect the phase of the locomotor activity rhythm. The body temperature rhythm of rats was affected neither by short‐term (a single injection) nor by long‐term (every day for 3 wks) diazepam treatment. Diazepam lacked effect on body core temperature even on the first day of administration, thereby ruling out the possibility of drug tolerance development. The fact that diazepam affects locomotor activity, but not core body temperature, suggests that different mechanisms mediate the actions of diazepam on locomotor activity and on core body temperature.