Time Of Lights-on Research Articles

Impact of Light at Night Is Phase Dependent: A Study on Migratory Redheaded Bunting (Emberiza bruniceps)

Artificial light at night (LAN) alters the physiology and behavior of an organism; however, very little is known about phase-dependent effects of LAN, particularly, in night migratory songbirds. Therefore, in this study, we investigated whether the effects of LAN on daily activity and photoperiodic responses in the Palearctic Indian migratory songbird, redheaded buntings (Emberiza bruniceps), is dependent on the different phases of the night. Male buntings maintained under short photoperiod (8L:16D; L = 100 lux, D < 0.1 lux) in individual activity cages were exposed to LAN (2 lux) for 6 weeks either in 4 h bin given at the different phases of 16 h night (early, mid, or late at ZT 08–12, ZT 14–18, or ZT 20–24, respectively; n = 9 each group) or throughout 16 h night (all night light, n = 6, ZT 08–24, the time of lights ON was considered as Zeitgeber time 0, ZT 0). A group (n = 6) with no LAN served as control. The results showed that LAN at the different phases of night induced differential effects as shown by an intense activity during the night, altered melatonin and temperature rhythms, and showed an increase in body mass and body fattening, food intake, and gonadal size. Midnight light exposure has a greater impact on migration and reproduction linked phenotypes, which is similar to the ones that received light throughout the night. The highlights of this study are that (i) LAN impacts day-night activity behavior, (ii) its continuity with the day alters the perception of day length, (iii) birds showed differential sensitivity to LAN in a phase-dependent manner, (iv) the direction of placing LAN affects the daily responses, e.g., LAN in the early night was “accepted” as extended dusk but the late night was considered as early dawn, and (v) midnight LAN was most effective and induced similar responses as continuous LAN. Overall, LAN induces long day responses in short days and shows differential sensitivity of the different phases of the night toward the light. This information may be valuable in adopting a part-night lighting approach to help reduce the physiological burden, such as early migration and reproduction, of artificial lighting on the nocturnal migrants.

The Food-entrainable Oscillator Is a Complex of Non-SCN Activity Bout Oscillators Uncoupled From the SCN Circadian Pacemaker

The food-entrainable oscillator, which underlies the prefeeding activity peak developed by restricted daily feeding (RF) in rodents, does not depend on the circadian pacemaker in the suprachiasmatic nucleus (SCN) or on the known clock genes. In the present study, to clarify the roles of SCN circadian pacemaker and nutrient conditions on the development of prefeeding activity peak, RF of 3-h daily feeding was imposed on four groups of adult male mice for 10 cycles at different circadian times, zeitgeber time (ZT)2, ZT8, ZT14, and ZT20, where ZT0 is the time of lights-on in LD12:12. Seven days after the termination of RF session with ad libitum feeding in between, total food deprivation (FD) for 72 h was imposed. Wheel-running activity and core body temperature were measured throughout the experiment. Immediately after the RF or FD session, the PER2::LUC rhythms were measured in the cultured SCN slices and peripheral tissues. Not only the buildup process and magnitude of the prefeeding activity peak, but also the percentages of nocturnal activity and hypothermia developed under RF were significantly different among the four groups, indicating the involvement of light entrained circadian pacemaker. The buildup of prefeeding activity peak was accomplished by either phase-advance or phase-delay shifts (or both) of activity bouts comprising a nocturnal band. Hypothermia under FD was less prominent in RF-exposed mice than in naïve counterparts, indicating that restricted feeding increases tolerance to caloric restriction as well as to the heat loss mechanism. RF phase-shifted the peripheral clocks but FD did not affect the clocks in any tissue examined. These findings are better understood by assuming multiple bout oscillators, which are located outside the SCN and directly drive activity bouts uncoupled from the circadian pacemaker by RF or hypothermia.

Discrepancies between self-reported and device-measured sleep parameters in adults with multiple sclerosis.

Sleep problems are a common consequence of multiple sclerosis; however, there is limited evidence regarding the agreement between device-measured and self-reported sleep parameters in adults with multiple sclerosis. The present study examined the agreement between self-reported and device-measured parameters of sleep quality in a sample of adults with multiple sclerosis. Participants (n = 49) completed a 7-day sleep diary and wore a wrist-worn ActiGraph GT3×+ (ActiGraph Corp., Pensecola, FL) for seven consecutive nights to quantify self-reported and device-measured sleep parameters, respectively. There was a significant discrepancy between self-reported and device-measured parameters of total time in bed (mean difference = 19.8 [51.3] min), sleep onset latency (mean difference = 22.2 [19.5] min), and frequency of awakenings during the night (mean difference = 12.8 [6.8]). Intraclass correlation estimates indicated poor agreement between methods on most parameters, except for total time in bed (intraclass correlation = 0.80). Bland-Altman plots suggested that total time in bed and total sleep time had acceptable levels of agreement and linear regression analyses indicated that sleep onset latency (F = 113.91, B = -1.34, P < .001), number of awakenings (F = 543.34, B = 1.85, P < .001), and sleep efficiency (F = 18.39, B = -0.77, P < .001) had significant proportional bias. Our results draw attention to the discrepancies between sleep parameter measurements and highlight the importance of including both self-report and device-measured outcomes for a complete and accurate representation of sleep in adults with multiple sclerosis. Cederberg KLJ, Mathison BG, Schuetz ML, Motl RW. Discrepancies between self-reported and device-measured sleep parameters in adults with multiple sclerosis. J Clin Sleep Med. 2022;18(2):415-421.

Intermittent Food Absence Motivates Reallocation of Locomotion and Feeding in Spotted Munia (Lonchura punctulata).

Background: Daily feeding and locomotion are interrelated behaviours. The time spent in feeding and rate of food intake depends on food availability. In low food condition, the birds would show intense movement (locomotion) for a longer time throughout the day however during abundant food supply they may chose higher activity and food intake in the morning and evening only. In the present study we hypothesized that in Spotted Munia (Lonchura punctulata), intermittent food availability during day would reallocate their interrelated behaviors, the feeding (food intake) and locomotor activity patterns.Methods: Two groups of birds (N = 6 each) were kept individually in activity cages under 12L:12D. Group 1 (Control; C) had ad libitum food but group 2 (Treatment; T) had food for 6 hours only (2 h presence followed by 2 h absence; 2P:2A) during 12 hour light period. In the first week, group 2 received food with ‘lights on’ (TI; ZT 0–2, 4–6 and 8–10; where ZT 0= zeitgeber time 0, time of lights ON). In the following week, the food was given 2 hours after ‘lights on’ (TII; ZT 2–4, 6–8, 10–12). The food intake and locomotor activity under each condition were observed.Results: The results showed that locomotor activity was induced during food deprivation and suppressed during food availability. Also the food deprivation led to increased food intake.Conclusion: Our results suggest that intermittent food availability/deprivation reallocates the locomotor activity and food intake in Spotted Munia.

Temporal expression profiles of ceramide and ceramide-related genes in wild-type and mPer1/mPer2 double knockout mice

Most living organisms exhibit circadian rhythms in physiology and behavior. These oscillations are generated by an endogenous circadian clock and control many biological processes. Ceramide has attracted attention as a signal mediator in diverse cell processes including cell death and differentiation. The relationships between ceramide expression levels and the circadian clock have not previously been investigated. To determine if there are circadian variations in the content of ceramide, we measured ceramide concentrations in the livers of wild-type (WT) and mPer1/mPer2 double knockout (DKO) mice. The ceramide concentration in WT mice was dramatically increased at Zeitgeber Time 9 (ZT9; 9h after lights-on time) and ZT21 but no rhythmicity in ceramide expression was seen in DKO mice. Because ceramide can be generated by the hydrolysis of sphingomyelin via sphingomyelinase (SMase), or by ceramide synthase (CerS)-mediated synthesis, we assayed the expression patterns of ceramide-related genes using real-time PCR. CerS2 expression levels showed a biphasic pattern of expression in WT mice but no rhythmicity in DKO mice. While the neutral SMase (nSMase) and acidic SMase (aSMase) mRNA in WT mice were expressed in a circadian manner, the correlation between the expression levels of these SMases with times of day was weak in DKO mice. Collectively, our findings suggest that both SMases and CerS2 mRNA expression are regulated by the presence of mPer1/mPer2 circadian clock genes in vivo, and imply that ceramide may play a vital role in circadian rhythms and physiology.

Cu/Zn superoxide dismutase is differentially regulated in period gene-mutant mice

The circadian clock in the brain coordinates the phase of peripheral oscillators that regulate tissue-specific physiological outputs. Here we report that circadian variations in the expression and activity of Cu/Zn superoxide dismutase (SOD1; EC 1.15.1.1) are present in liver homogenates from mice. The SOD1 mRNA expression from wild-type (WT) mice peaked at Zeitgeber Time 9 (ZT9; 9h after lights-on time). While there was no rhythmicity in that from period2 (per2) gene knockout (P2K) mice, the level of SOD1 from per1/per2 double knockout (DKO) mice was significantly elevated at ZT5. The enzyme activity of SOD1 was also rhythmic in the mouse liver. Moreover, the total amount of the SOD1 exhibited a rhythmic oscillation with a peak at ZT9 in the liver from WT mice. We also found that tert-butylhydroperoxide (t-BHP)-induced oxidative damage in both WT and P2K mouse embryonic fibroblast (MEF) cells resulted in the up-regulation of SOD1 levels. Our data suggest that the expression of an important antioxidant enzyme, SOD1, is under circadian clock control and that mice are more susceptible to oxidative stress depending on the time of day.

Brief daily suckling shifts locomotor behavior and induces PER1 protein in paraventricular and supraoptic nuclei, but not in the suprachiasmatic nucleus, of rabbit does

Nursing in the rabbit is a circadian event during which mother and pups interact for a period of < 5 min every day. Here we explored behavioral and neuronal changes in the mother by analyzing the suprachiasmatic nucleus (SCN), and oxytocinergic (OT) neurons in the paraventricular nucleus (PVN) and the supraoptic nucleus (SON). We maintained lactating does in a light-dark cycle (lights on at 07 : 00 hours; ZT0); they were scheduled to nurse during either the day (ZT03) or the night (ZT19). Groups of intact and nursing females was perfused, one at each 4-h point through a 24-h cycle. We explored, by immunohistochemistry, the PER1 expression and double-labeling, with OT antibody, of neurons in the PVN and SON at lactation on day 7. In the SCN, intact and lactating groups had peak PER1 expression at ZT11; however, there was a reduction in PER1 at peak time in the nursing groups. There was a locomotor activity rhythm with increased activity around the time of lights-on in intact subjects and around the time of suckling in lactating does. There was an induction of PER1 in OT cells in the PVN and SON that shifted in phase with timing of nursing. We further explored the maintenance of the PER1 expression in OT cells in nursing-deprived does and found a significant decrease at 24 and 48 h after the last nursing. We conclude that suckling induced PER1 in the PVN and SON, but not in the SCN, in nursing does, and also shifted their locomotor behavior.

The phasing of circadian rhythms in mice kept under normal or short photoperiods

Mammals represent a multi-oscillatory system. Not only different rhythmic function but also a certain rhythmic function may be controlled by several oscillators. Also, the LD cycle, though being the main zeitgeber, may have different effects on different rhythmic functions. The aim of the present study was to investigate putative changes of internal phase relationships between different rhythms and their phase relationship with respect to the light–dark cycle following a gradual lengthening of the dark time. The investigations were carried out on adult female mice (HaZ:ICR). Animals were initially kept in a 12:12 h LD cycle. The dark time was then lengthened once a week by delaying the time of lights-on by 1 h until an L/D = 6:18 h was reached. The motor activity was recorded continuously. The daily profiles of food intake, liver glycogen and melatonin in the serum and the pineal gland were estimated as transverse studies under L/D = 12:12 h and L/D = 6:18 h. Under short-day conditions (L/D = 6:18 h), the evening onset of the main maximum of motor activity and food intake was delayed in relation to lights-off and the onset of the secondary maximum in the morning was advanced in relation to lights-on. Similar phase changes were found with respect to the onset of glycogen synthesis and its breakdown. In the case of melatonin, the rise and the fall occurred earlier or later, respectively, and the phase differences with respect to lights-off and lights-on became smaller. If the mid-point between the evening and morning onsets of activity and food intake and between the rises and falls of glycogen and melatonin concentrations are considered, then the changes in all rhythmic functions were similar. Consequently, the phase relationships between different rhythmic functions and with respect to the light–dark cycle, and thus the internal and the external temporal order, were preserved under different light–dark ratios. Also, the results are in good accordance with a modified two-oscillator model where lights-off triggers advances of an evening oscillator and lights-on delays a morning oscillator.

The pattern of melatonin secretion is rhythmic in the domestic pig and responds rapidly to changes in daylength.

The aim of the study was to investigate the capability of pigs to respond to abrupt changes in lighting conditions by means of alterations in circadian melatonin profiles. Sixteen pre-pubertal crossbred male pigs weighing 40-45 kg were housed in individual pens in four temperature- and lighting-controlled climate rooms (four pigs per room). In two rooms there was a light-dark cycle of 16 L:8 D (Group A) and in two other rooms 8 L:16 D (Group B). Under both lighting regimens light intensity at pig eye-level was 220-240 lx during the light phase and less than 7 lx (red light) during the dark phase. The lighting regimens were changed after 2 wks to the opposite regimen and the change was repeated after a further 2 wks, so that animals ended up with the same light cycle with which they started. Blood was sampled at 2-hr intervals for 48 hr spanning each time of change in lighting. A further 24-hr sampling was performed at the end of the experiment (2 wks after the last change) in both groups and 1 wk after the change from short to long day lighting in Group A. On 83/86 occasions, pigs exhibited a clear circadian rhythm in plasma melatonin under both lighting regimens. Pigs responded immediately to the change from long to short day lighting by advancing melatonin secretion to the earlier lights-off time and some pigs were able to extend secretion to the delayed lights-on time. For short to long day changeover there was a small immediate response, with secretion pattern following the previously entrained endogenous rhythm to within 3 hr of the previous lights-on time. After 1 wk commencement of secretion was delayed by up to 2 hr, while after 2 wks some pigs were able to delay commencement of secretion until lights-off or to cease at lights-on. It is concluded that the domestic pig is able to commence adjustment to abrupt changes in photoperiod within a 1-wk acclimatization by altering circadian melatonin secretion. The present study suggests that it may be possible to use simplified lighting regimens instead of stepwise changing lighting programs in commercial piggeries to reduce the influence of season on production.

Juvenile circadian core temperature rhythm in Wistar and lean (Fa/-) Zucker rat pups

We compared the daily rhythms of cold defense in Wistar and lean (Fa/-) Zucker rat pups isolated from maternal care. Pups, born in a colony with lights-on from 7:00 to 19:00, were artificially reared from 10 to 13 days of age in continuous light at moderate cold loads, while we recorded core temperature (Tc) and metabolic rate (MR). Pups of both strains showed daily rhythms distinctly different from those of adult rats: on the first day of artificial rearing Tc steeply decreased shortly before the time of lights-on in the breeding colony and reached a daily minimum of 31.8 ± 0.4°C at 7:17 ± 17 min in Zucker pups (± SE, N = 24) and of 33.4 ± 0.2°C at 8:59 ± 9 min ( N = 25) in Wistar pups. On each day, the average daily minium Tc was significantly higher in Wistar than in Zucker rats, whereas the daily maximum Tc were close to 36°C in both strains. The daily rhythms of MR resembled those of Tc. We conclude that circadian differences of cold defense, which must be expected to influence a variety of behavioral and physiological parameters, are not a pecularity of Zucker rat pups but also occur, though with a smaller amplitude, in 10-day-old Wistar pups.

Time-of-day discrimination by pigeons,Columba livia

Pigeons were trained to discriminate between four keys. One provided food in the mornings, another provided food in the afternoons, and two never provided food. Three experiments were performed to determine whether pigeons could track food availability over a 24-h period. All the subjects appeared to demonstrate time-place associative learning. A fourth experiment was designed to investigate the mechanisms underlying the timing behavior. Lights-on time was shifted back by 6 h, and no decrease in performance was found during the first session following the phase shift. This suggests that a circadian type of timing mechanism with a self-sustaining oscillator mediates time-place learning over a period of 24 h. Further support for this notion was found in a fifth experiment, in which the subjects were tested in constant dim light. In that experiment, the subjects’ continued correct responding provides additional support for a self-sustaining circadian timer.

Daily profile of the extracellular concentration of glutamate in the suprachiasmatic region of the Siberian hamster.

Daily profiles of the extracellular concentrations of glutamate (GLU), aspartate, and glutamine in the region of the suprachiasmatic nuclei (SCN) of the Djungarian hamster were examined by in vivo microdialysis. Hamsters with a dialysis probe located in or near the SCN exhibited a diurnal variation in the extracellular concentration of GLU. Glutamate levels rose gradually during the latter half of the subjective day and through the night to reach peak levels near the time of lights-on. The extracellular concentration of GLU was significantly elevated (P < 0.05) during the latter half of the dark phase relative to the level at midday. The average peak in extracellular GLU was 217% of the midday value (P < 0.05; n = 5). Diurnal fluctuations were not consistently observed in the concentrations of aspartate in SCN dialysates. The nocturnal rise in extracellular GLU in the SCN was not attenuated by perfusion with 10 microM tetrodotoxin, although this concentration of tetrodotoxin completely blocked increases in extracellular GLU caused by simultaneous perfusion with 30 microM veratridine. Collectively, these results point to a diurnal rhythm in extracellular concentration of GLU in the SCN region which may not reflect diurnal variations in synaptic activity.

New sensitive serum melatonin radioimmunoassay employing the Kennaway G280 antibody: Syrian hamster morning adrenergic response

A new procedure with the G280 antibody of Kennaway provides an assay for circulating melatonin (aMT) with a sample volume (200 microliters), an analytic (0.33 pg/ml) and functional (0.62-0.80 pg/ml) detectability, a 50% displacement dose (6.4 pg/ml), a Kd (0.657 pM), and measured circulating daytime levels lower than reported for previous procedures, and 100% assay recovery. The normal daytime range in adult human and Syrian hamster serum was 0.4-4 pg/ml. The pattern of fall of the nocturnal surge of Syrian hamster serum aMT near the time of lights-on was unaltered by extended darkness. Isoproterenol (ISO) injection 1 hr after lights-on, when aMT had reached daytime levels, raised serum and pineal aMT dramatically 2 hr postinjection. The same dose of ISO injected 4 hr into light produced only a small detectable increase. Novel extension of nocturnal darkness did not affect the responses to ISO. Thus, when they are allowed to occur at the usual time on a 10-hr dark schedule, both the fall from the nocturnal aMT surge and the subsequent loss of pineal beta-adrenergic responsiveness in this species occur endogenously (probably entrained) rather than from gating by acute effects of morning light. Changes in daytime serum aMT consistent with concomitant changes in the pineal can be measured with a sufficiently sensitive radioimmunoassay.

Pineal melatonin and locomotor activity of rats under gradual illuminance transitions

The locomotor activity and pineal melatonin patterns of adult male rats were compared under two different lighting regimes. The animals were kept 8 days under 12/12 h light/dark cycles with abrupt or slowly decreasing and increasing transitions (twilight periods about 2 h). The onsets of high activity and melatonin rise were phase-locked in the two conditions and related to about half-maximal illuminance level of the gradual dusk. The high activity of the control rats stopped 30–60 min before the abrupt light onset and the rats under the gradual lighting transitions ceased the locomotor activity at about 1 hour before the half-maximal illuminance. The melatonin peak levels were found 4 h before the abrupt lights-on time. Under the slow illuminance transitions the average melatonin peak was related to the illuminance level between maximum and minimum in the morning. Thus, both the melatonin rhythm and the rest-activity rhythm under the gradual dawn and dusk were adjusted according to about half-maximal illuminances in the present conditions.

Rat circadian rhythms entrain to a descending saw-tooth light intensity cycle

We reexamined whether rat circadian rhythms entrained to the light intensity cycle of a descending saw-tooth (ST-d) form, in which illuminance decreased rectilinearly from 300 lx to 0 lx in 24 h, and abruptly returned to 300 lx (lights-on time). Ambulation, drinking and subcutaneous body temperature were simultaneously monitored in 5 intact, 5 pinealectomized and 5 orchiectomized rats. Additionally, sleep was monitored in the intact rats. In all the rats, entrainment was confirmed during 65 days' exposure to the ST-d cycle. The waveforms of the entrained rhythms were much modified compared with those during LD 12:12. The estimated activity periods of the entrained rhythms straddled the lights-on time of the ST-d cycle. In all the groups of rats, administration of a single ST-d cycle in constant dim red light produced only delay shifts irrespective of its circadian phase, and there was no significant circadian variation in the magnitude of phase shifts. The results indicate that rat circadian rhythms entrain to the ST-d cycle with an unexpected phase position, which cannot be explained by the phase-response curve.

Effects of Photoperiod on Spawning Rhythm in the Tobinumeri-Dragonet Repomucenus beniteguri.

In order to record the daily spawning time of individual fish, one female and two male tobinu-meri-dragonet were reared in separate indoor tanks (1×2×0.8 m) from September 30. The photo-period in each tank was regulated by a timer and water temperature was maintained at 22-24°C during the experiment and the spawning time was determined by checking the existence of spawned eggs on the net placed at the overflow outlet of each tank every 15 minutes. In experiment 1, two females continued daily spawning. Spawning occurred during 1700-1900 hr when the fish were kept under a photophase of 0500-1900h (14L10D). When the photo-phase was shifted from 0500-1900h to 0100-1500h (14L10D), the spawning time was advanced by an average of 3.2 hours. Thereafter, we delayed the time of lights-on by 4 hours (10L14D); how-ever, spawning time was delayed only 0.7 hours. By delaying the time of lights-off by 4 hours (14L10D), spawning time was delayed 2.6 hours. In experiment 2, four females continued daily spawning. When the time of lights-off was advanced 4 hours (10L14D), spawning time advanced 3.1 hours. These results indicate that the spawning time is determined photoperiodically. The time of lights-off seems more significant than that of lights-on in the regulation of spawning time, indicating that “lights-off” is working as a synchronizer.

Sparrow circadian rhythm responses to rotated light-dark schedules

House sparrows, Passer domesticus, were individually subjected to light-dark regimens while their perching activity was continuosly monitored. The sparrows resynchronized in 5 days when LD8:16 (8 hr of light alternating with 16 hr of dark) was advanced by 8 hr; however, the sparrows were 1.7 hr from resynchronization after 5 days when the schedule was delayed 8 hr. Sparrows subjected to two simultaneously presented light-dark schedules (rotated LD8:16 provided by one light source imposed together with LD12:12 provided by a second light source) did not ignore either cycle; instead the sparrows responded as to a sequence of photoperiods; they resynchronzied the onsets of their activity with the first time of lights-on of the collective photoperiod provided by the two light sources.