Variation of visual detection over the 24-hour period in humans

A circadian rhythm for visual sensitivity has been intensively assessed in animals. This rhythm may be due to the existence of a circadian clock in the mammalian eye, which could account for fluctuating sensitivity to light over the day in certain species. However, very few studies have been devoted to the human visual system. The present experiment was designed to assess a possible rhythm of visual sensitivity using a psychophysical method over the whole 24h period. Twelve subjects underwent visual detection threshold measures in a protocol that allowed one point every 2h. The results show that the visual detection threshold changes over the 24h period, with high thresholds in the morning, a progressive decrease over the day and the early night, and an increase during the last part of the night. These data suggest that a circadian rhythm influences visual sensitivity to mesopic luminance in humans. (Chronobiology International, 17(6), 795-805, 2000)     

The possible role of prolactin in the circadian rhythm of leptin secretion in male rats

In humans there is a circadian rhythm of leptin concentrations in plasma with a minimum in the early morning and a maximum in the middle of the night. By taking blood samples from adult male rats every 3 hr for 24 hr, we determined that a circadian rhythm of plasma leptin concentrations also occurs in the rat with a peak at 0130h and a minimum at 0730h. To determine if this rhythm is controlled by nocturnally released hormones, we evaluated the effect of hormones known to be released at night in humans, some of which are also known to be released at night in rats. In humans, prolactin (PRL), growth hormone (GH), and melatonin are known to be released at night, and adrenocorticotropic hormone (ACTH) release is inhibited. In these experiments, conscious rats were injected intravenously with 0.5 ml diluent or the substance to be evaluated just after removal of the first blood sample (0.3 ml), and additional blood samples (0.3 ml) were drawn every 10 min thereafter for 2 hr. The injection of highly purified sheep PRL (500 microg) produced a rapid increase in plasma leptin that persisted for the duration of the experiment. Lower doses were ineffective. To determine the effect of blockade of PRL secretion on leptin secretion, alpha bromoergocryptine (1.5 mg), a dopamine-2-receptor agonist that rapidly inhibits PRL release, was injected. It produced a rapid decline in plasma leptin within 10 min, and the decline persisted for 120 min. The minimal effective dose of GH to lower plasma leptin was 1 mg/rat. Insulin-like growth factor (IGF-1) (10 microg), but not IGF-2 (10 microg), also significantly decreased plasma leptin. Melatonin, known to be nocturnally released in humans and rats, was injected at a dose of 1 mg/rat during daytime (1100h) or nighttime (2300h). It did not alter leptin release significantly. Dexamethasone (DEX), a potent glucocorticoid, was ineffective at a 0. 1-mg dose but produced a delayed, significant increase in leptin, manifest 100-120 min after injection of a 1 mg dose. Since glucocorticoids decrease at night in humans at the time of the maximum plasma concentrations of leptin, we hypothesize that this increase in leptin from a relatively high dose of DEX would mimic the response to the release of corticosterone following stress in the rat and that glucocorticoids are not responsible for the circadian rhythm of leptin concentration. Therefore, we conclude that an increase in PRL secretion during the night may be responsible, at least in part, for the nocturnal elevation of leptin concentrations observed in rats and humans.     

Circadian activation of bullfrog retinal mitogen-activated protein kinase associates with oscillator function

The vertebrate retina retains a circadian oscillator, and its oscillation is self-sustained with a period close to 24 h under constant environmental conditions. Here we show that bullfrog retinal mitogen-activated protein kinase (MAPK) exhibits an in vivo circadian rhythm in phosphorylation with a peak at night in a light/dark cycle. The phosphorylation rhythm of MAPK persists in constant darkness with a peak at subjective night, and this self-sustained rhythm is also observed in cultured retinas, indicating its close interaction with the retinal oscillator. The rhythmically phosphorylated MAPK is detected only in a discrete subset of amacrine cells despite ubiquitous distribution of MAPK throughout the retinal layers. Treatment of the cultured retinas with MAPK kinase (MEK) inhibitor PD98059 suppresses MAPK phosphorylation during the subjective night, and this pulse perturbation of MEK activity induces a significant phase delay (4-8 h) of the retinal circadian rhythm in MAPK and MEK phosphorylation. These observations strongly suggest that the site-specific and time-of-day-specific activation of MAPK contributes to the circadian time-keeping mechanism of the retinal clock system.     

Extremely low frequency magnetic field exposure modulates the diurnal rhythm of the pain threshold in mice

The aim of this study was to determine whether exposure to extremely low frequency magnetic field (ELF-MF) affects the normal diurnal rhythm of the pain threshold in mice. Pain thresholds were evaluated in mice using the hot plate test. A significant increase of pain threshold during night was observed compared to that during day. This rhythm was attenuated by both constant exposure to light (LL) and constant exposure to darkness (DD) for 5 days. Under DD exposure, the diurnal rhythm in pain threshold was restored when mice were exposed to ELF-MF (60 Hz, 1.5 mT for 12 h daily, from 08:00 to 20:00 h) for 5 days. The diurnal rhythm was not reversed under dark with reversed ELF-MF cycle (exposure to 1.5 mT from 20:00 to 08:00 h, next day) for 5 days, although pain threshold in the ELF-MF exposed period of night was slightly decreased. The diurnal rhythm of melatonin analgesic effect related to pain threshold was also observed under DD by the exposure of ELF-MF for 5 days, but not for 5 nights. The present results suggest that ELF-MF may participate in the diurnal rhythm of pain threshold by acting on the system that is associated with environmental light-dark cycle.     

Evidence that photoperiodic, dark time measurement in Pharbitis nil involves a circadian rather than a semidian rhythm

Experiments were carried out to determine whether a semidian (12 h) rhythm in flowering response operates in Pharbitis nil as the basis for photoperiodic time measurement. The effect of 5 min far-red light followed by 85 min dark (FRD) given 4, 8,14 and 22 h before the end of a 48 h photoperiod on night-break timing and critical night length was determined. When given 4 h before the end of a 48 h photoperiod, an interruption with FRD advanced the phase of the circadian rhythm in the night-break inhibition of flowering. In contrast, earlier interruptions of the photoperiod had no effect on the phase of the rhythm. The critical night length was modified by FRD given 4 h (shortened) or 8 h (lengthened) before the end of the photo-period; when given at other times FRD did not alter the critical night length. The results are discussed in relation to the basis for photoperiodic timekeeping, with particular reference to suggestions for the involvement of a semidian rhythm. A circadian model based on the concept of limit cycles is described.     

Shifts in the semidian rhythm of flowering do determine night-break timing and critical night length in Pharbitis nil

There is a semidian (≈12 h) rhythm in the flowering response of the short-day plant Pharbitis nil Choisy following 90 min exposure to either far-red light/darkness or a temperature drop (27 °C to 12 °C) given at various times in constant conditions before an inductive dark period. This semidian rhythmic response to the temperature-drop pretreatments in the light is also evident through the inductive dark period without change of phase. Furthermore, those pretreatments which increase flowering also advance the time of maximum sensitivity to red light (R) interruptions of the dark period by up to 1.5 h and shorten the critical night length. Conversely, pretreatments which reduce flowering delay the time of maximum R inhibition by up to 1.5 h and increase the critical night length by the same amount. However the phase of a circadian rhythm of flowering response had no effect on either the time of maximum R inhibition or the critical night length. Thus, the semidian rhythm determines both the time of maximum R inhibition and the critical night length in Pharbitis. Received: 8 November 1997 / Accepted: 7 January 1998     

Temporal variation in the specific dynamic action of juvenile New Zealand rock lobsters, Jasus edwardsii

To enhance the on-growing of Jasus edwardsii in culture, it is important to understand the feeding physiology of juveniles. In crustaceans, there is a loss of energy and an increase in oxygen consumption (specific dynamic action or SDA) associated with feeding. The present research measured the SDA of juvenile J. edwardsii fed either in the morning or at night held at 15 degrees C. Closed box respirometry was used to measure oxygen consumption (MO(2)) and ammonia excretion in juvenile lobsters. Juveniles exhibited a nocturnal rhythm in both MO(2) and ammonia excretion. The factorial rise in MO(2) (1.58+/-0.03 times) for lobsters fed in the morning was significantly less than lobsters fed at night (1.80+/-0.01 times). Lobsters fed in the morning had a significantly shorter SDA (30+/-1.2 h) response compared to lobsters fed at night (36+/-1 h). Energy loss as a result of digestion was less for lobsters fed in the morning. Therefore, if juvenile J. edwardsii are fed in the morning, they could optimise the energy content of the meal and this could result in increased growth.     

中缅树鼩体温、代谢率和蒸发失水日节律

为研究中缅树鼩(Tupaia belangeri)体温、代谢率和蒸发失水的日节律变化,采用植入式体温计测定了中缅树鼩24 h的体温,以及24 h中4个时间段(05:00～07:00时、11:00～13:00时、17:00～19:00时和23:00～01:00时)热中性区(30℃)的静止代谢率(RMR)、非颤抖性产热(NST)和蒸发失水(EWL)。结果显示,中缅树鼩的体温具有日节律变化,最高值和最低值分别出现在11:00时和03:00时,各为(39.45±0.26)℃和(36.34±0.24)℃;静止代谢率、非颤抖性产热和蒸发失水在4个时间段都有显著差异,表现出显著的日节律变化,代谢率在23:00～01:00时最大,O2含量为(2.58±0.04)ml/(g.h),在11:00～13:00时最小,O2含量为(2.28±0.09)ml/(g.h);非颤抖性产热在05:00～07:00时最大,O2含量为(3.08±0.14)ml/(g.h),在11:00～13:00最小,O2含量为(2.69±0.63)ml/(g.h);蒸发失水在17:00～19:00时最大,失水量为(3.60±0.31)mg/(g.h)。结果表明,体温的日节律变化主要与环境温度的日节律变化和下午出窝取食活动性增强有关;当夜晚环境温度相对较低的时候,通过增强静止代谢率和非颤抖性产热来增加产热,而白天环境温度相对较高的时候,通过增强蒸发失水散热来调节体温。     

Angiotensin receptor blockers shift the circadian rhythm of blood pressure by suppressing tubular sodium reabsorption

Recently, we found that an angiotensin II receptor blocker (ARB) restored the circadian rhythm of the blood pressure (BP) from a nondipper to a dipper pattern, similar to that achieved with sodium intake restriction and diuretics (Fukuda M, Yamanaka T, Mizuno M, Motokawa M, Shirasawa Y, Miyagi S, Nishio T, Yoshida A, Kimura G. J Hypertens 26: 583-588, 2008). ARB enhanced natriuresis during the day, while BP was markedly lower during the night, resulting in the dipper pattern. In the present study, we examined whether the suppression of tubular sodium reabsorption, similar to the action of diuretics, was the mechanism by which ARB normalized the circadian BP rhythm. BP and glomerulotubular balance were compared in 41 patients with chronic kidney disease before and during ARB treatment with olmesartan once a day in the morning for 8 wk. ARB increased natriuresis (sodium excretion rate; U(Na)V) during the day (4.5 ± 2.2 to 5.5 ± 2.1 mmol/h, P = 0.002), while it had no effect during the night (4.3 ± 2.0 to 3.8 ± 1.6 mmol/h, P = 0.1). The night/day ratios of both BP and U(Na)V were decreased. The decrease in the night/day ratio of BP correlated with the increase in the daytime U(Na)V (r = 0.42, P = 0.006). Throughout the whole day, the glomerular filtration rate (P = 0.0006) and tubular sodium reabsorption (P = 0.0005) were both reduced significantly by ARB, although U(Na)V remained constant (107 ± 45 vs. 118 ± 36 mmol/day, P = 0.07). These findings indicate that the suppression of tubular sodium reabsorption, showing a resemblance to the action of diuretics, is the primary mechanism by which ARB can shift the circadian BP rhythm into a dipper pattern.     

Study of autacoids role in the regulation of circadian rhythm of the human renal function

The circadian rhythm of urine formation was studied in younger (27 +/- 8 yr) and aged (76 +/- 3 yr) males. In 11 younger healthy examined persons a decrease of diuresis during the night as compared with the day time was due to a rise of solute free water reabsorption. In 31 aged males the change of the urine formation rhythm with increase of the nocturnal diuresis is based on a rise of osmolal clearance combined with an increase of the solute free water reabsorption. It is shown that blockade of autacoid secretion leads to normalization of the diuresis circadian rhythm. In 10 aged men, nocturia was due to a decrease of vasopressin secretion which resulted in a decrease of the solute free water reabsorption and an increase of diuresis. The obtained data are considered as an evidence for the role of renal autacoids, alongside with vasopressin, in regulation of the circadian rhythm of the kidney function.     

VARIATION OF VISUAL DETECTION OVER THE 24-HOUR PERIOD IN HUMANS

A circadian rhythm for visual sensitivity has been intensively assessed in animals. This rhythm may be due to the existence of a circadian clock in the mammalian eye, which could account for fluctuating sensitivity to light over the day in certain species. However, very few studies have been devoted to the human visual system. The present experiment was designed to assess a possible rhythm of visual sensitivity using a psychophysical method over the whole 24h period. Twelve subjects underwent visual detection threshold measures in a protocol that allowed one point every 2h. The results show that the visual detection threshold changes over the 24h period, with high thresholds in the morning, a progressive decrease over the day and the early night, and an increase during the last part of the night. These data suggest that a circadian rhythm influences visual sensitivity to mesopic luminance in humans. (Chronobiology International, 17(6), 795–805, 2000)     

Melatonin accelerates reentrainment of the circadian rhythm of its own production after an 8-h advance of the light-dark cycle

Summary The rhythm in melatonin production in the rat is driven by a circadian rhythm in the pineal N-acetyltransferase (NAT) activity. Rats adapted to an artificial lighting regime of 12 h of light and 12 h of darkness per day were exposed to an 8-h advance of the light-dark regime accomplished by the shortening of one dark period; the effect of melatonin, triazolam and fluoxetine, together with 5-hydroxytryptophan, on the reentrainment of the NAT rhythm was studied.In control rats, the NAT rhythm was abolished during the first 3 cycles following the advance shift. It reappeared during the 4th cycle; however, the phase relationship between the evening rise in activity and the morning decline was still compressed.Melatonin accelerated the NAT rhythm reentrainment. In rats treated chronically with melatonin at the new dark onset, the rhythm had already reappeared during the 3rd cycle, in the middle of the advanced night, and during the 4th cycle, the phase relationship between the evening onset and the morning decline of the NAT activity was the same as before the advance shift. In rats treated chronically with melatonin at the old dark onset or in those treated with melatonin 8 h, 5 h and 2 h after the new dark onset during the 1st, 2nd and 3rd cycle, respectively, following the advance shift, the NAT rhythm reappeared during the 3rd cycle as well but in the last third of the advanced night only.Neither triazolam nor fluoxetine together with 5-hydroxytryptophan administered around the new dark onset facilitated NAT rhythm reentrainment after the 8-h advance of the light-dark cycle.Abbreviations NAT N-acetyltransferase - LD cycle light-dark cycle - CT circadian time - LD xy light dark cycle comprising x h of light and y h of darkness     

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.     

Circadian expression of clock genes in the rat eye and brain

The light sensing system in the eye directly affects the circadian oscillator in the mammalian suprachiasmatic nucleus (SCN). To investigate this relationship in the rat, we examined the circadian expression of clock genes in the SCN and eye tissue during a 24 h day/night cycle. In the SCN, rPer1 and rPer2 mRNAs were expressed in a clear circadian rhythm like rCry1 and rCry2 mRNAs, whereas the level of BMAL1 and CLOCK mRNAs decreased during the day and increased during the night with a relatively low amplitude. It seems that the clock genes of the SCN may function in response to a master clock oscillation in the rat. In the eye, the rCry1 and rCry2 were expressed in a circadian rhythm with an increase during subjective day and a decrease during subjective night. However, the expression of Opn4 mRNA did not exhibit a clear circadian pattern, although its expression was higher in daytime than at night. This suggests that cryptochromes located in the eye, rather than melanopsin, are the major photoreceptive system for synchronizing the circadian rhythm of the SCN in the rat.     

Altered cardiovascular responses to chronic angiotensin II infusion in aged rats

In this work we determined by telemetry the cardiovascular effects produced by Ang II infusion on blood pressure (BP) and heart rate (HR) in aged rats. Male Wistar aged (48-52 weeks) and young (12 weeks) rats were used. Ang II (6 microg/h, young, n=6; aged, n=6) or vehicle (0.9% NaCl 1 microl/h, young, n=4; aged, n=5) were infused subcutaneously for 7 days, using osmotic mini-pump. The basal diurnal and nocturnal BP values were higher in aged rats (day: 98+/-0.3 mm Hg, night: 104+/-0.4 mm Hg) than in the young rats (day: 92+/-0.2 mm Hg, night: 99+/-0.2 mm Hg). In contrast, the basal diurnal and nocturnal HR values were significantly smaller in the aged rats. Ang II infusion produced a greater increase in the diurnal BP in the aged rats (Delta MAP=37+/-1.8 mm Hg) compared to the young ones (Delta MAP=30+/-3.5 mm Hg). In contrast, the nocturnal MAP increase was similar in both groups (young rats; Delta MAP=22+/-3.0 mm Hg, aged rats; Delta MAP=24+/-2.6 mm Hg). During Ang II infusion HR decreased transiently in the young rats. An opposite trend was observed in the aged rats. Ang II infusion also inverted the BP circadian rhythm, in both groups. No changes in HR circadian rhythm were observed. These differences suggest that the aging process alters in a different way Ang II-sensitive neural pathways involved in the control of autonomic activity.     

Entrainment of the circadian rhythm in the rat pineal N-acetyltransferase activity by prolonged periods of light

Summary Entrainment of the circadian rhythm in the pineal N-acetyltranferase activity by prolonged periods of light was studied in rats synchronized with a light:dark regime of 1212 h by observing phase-shifts in rhythm after delays in switching off the light in the evening or after bringing forward of the morning onset of light. When rats were subjected to delays in switching off the light of up to 10 h and then were released into darkness, phase-delays of the evening N-acetyltransferase rise during the same night corresponded roughly to delays in the light switch off. However, phasedelays of the morning decline were much smaller. After a delay in the evening switch off of 11 h, no N-acetyltransferase rhythm was found in the subsequent darkness. The evening N-acetyltransferase rise was phase-delayed by 6.2 h at most 1 day after delays. Phase-delays of the morning Nacetyltransferase decline were shorter than phasedelays of the N-acetyltransferase rise by only 0.7 h to 0.9 h at most. Hence, 1 day after delays in the evening switch off, the period of the high night N-acetyltransferase activity may be shortened only slightly. The N-acetyltransferase rhythm was abolished only after a 12 h delay in switching off the light.Rats were subjected to a bringing forward of the morning light onset and then were released into darkness 4 h before the usual switch off of light. In the following night, the morning N-acetyltransferase decline, but not the evening rise, was phase advanced considerably. Moreover, when the onset of light was brought forward to before midnight, the N-acetyltransferase rise was even phase-delayed. Hence, 1 day after bringing forward the morning onset of light, the period of the high night N-acetyltransferase activity may be drastically reduced. When rats were subjected to a 4 h light pulse around midnight and then released into darkness, the N-acetyltransferase rhythm in the next night was abolished.The data are discussed in terms of a two-component pacemaker controlling the N-acetyltransferase rhythm. It is suggested that delays in the evening switch off of light may disturb the N-acetyltransferase rhythm the next day only a little, as the morning component may adjust to phasedelays of the evening component almost within one cycle. On the other hand, bringing forward the morning onset of light may disturb the N-acetyltransferase rhythm heavily the next day, as the evening component not only does not adjust to phase-advances of the morning component, but it may even be phase-delayed when the light onset occurs before midnight.Abbreviations NAT N-acetyltransferase - PRC phase response curve - E evening component of the N-acetyltransferase rhythm or of its pacemaker - M morning component of the N-acetyltransferase rhythm or of its pacemaker - LD xy light dark cycle comprising x h of light and y h of darkness     

The semidian rhythm in flowering response of Pharbitis nil in relation to dark period time measurement and to a circadian rhythm

When seedlings of Pharbitis nil Choisy, cv. Violet, are exposed to a single inductive dark period at 27°C, brief interruptions with red light (R) can be promotive after 2–3 h of darkness but increasingly inhibitory to flowering up to the 8–9th h of darkness. This rhythmic response to R interruptions can be advanced in phase by > 1 h when the preceding light period is interrupted with far-red (FR) 2 h before darkness (FR -2 h) or with FR – 15 h, whereas FR –8 h or FR–22 h retard the rhythm. These shifts in the R interruption rhythm are paralleled by equal shifts in the length of the dark period required for flowering. Brief FR interruptions of darkness displayed a similar rhythm which was also advanced by FR –2 h and retarded by FR –8 h. We conclude therefore that the semidian rhythm in the light, which we have previously described, continues through at least the first 12 h of darkness, is manifested in the R interruption rhythm, and determines the critical night length. A circadian rhythm with a marked effect on flowering was also identified, but several lines of evidence suggest that the circadian and semidian rhythms have independent additive effects on flowering and do not appear to show phase interaction.     

Phase relations between the circadian leaf movement rhythm and its zeitgeber within the range of entrainment in the cotton plant,Gossypium hirsutum L.

Abstract

The leaf movement rhythm of Gossypium hirsutum L. (cv. Lakshmi) could be entrained to 24 h LD cycles with different photofractions varying from 4 to 20 h such that the night peak position of the rhythm occurred during darkness. The phase angle (ψ) of the rhythm varied in a regular manner with different photoperiods of a 24 h LD cycle. Under 24 h LD cycles with different photoperiods, the leaf movement shows probable evidences for the concurrent participation of a ‘light‐on’ and ‘light‐off rhythm.     

Protein synthesis is a required process for the optic lobe circadian clock in the cricket Gryllus bimaculatus

The effects of a translation inhibitor, cycloheximide (CHX), on the circadian neuronal activity rhythm of the optic lamina-medulla compound eye complex cultured in vitro were investigated in the cricket Gryllus bimaculatus. When the complex was treated with 10(-5) M CHX for 6 h, the rhythm exhibited a marked phase shift. The magnitude and direction of the phase shift were dependent on the phase at which the complex was treated with CHX; phase delays occurred during the late subjective day to early subjective night, whereas phase advances occurred around the late subjective night. Continuous application of CHX abolished circadian rhythms of both the spontaneous neuronal activity and the visually evoked response. However, it abolished neither the spontaneous activity nor the visually evoked response. As washed with fresh medium after CHX treatment, the rhythm soon reappeared and the subsequent phase was clearly correlated to the termination time of the treatment. These results suggest that protein synthesis is also involved in the cricket optic lobe circadian clock, and that the clock-related protein synthesis may be active during the late subjective day to subjective night.     

Constant light disrupts the circadian but not the circatidal rhythm in mangrove crickets

Mangrove crickets have a circatidal activity rhythm (~12.6 h cycles) with a circadian modulation under constant darkness (DD), whereby activity levels are higher during subjective night low tides than subjective day low tides. This study explored the locomotor activity rhythm of mangrove crickets under constant light (LL). Under LL, the crickets also exhibited a clear circatidal activity rhythm with a free-running period of 12.6 ± 0.26 h (mean ± SD, n = 6), which was not significantly different from that observed under DD. In contrast, activity levels were almost the same between subjective day and night, unlike those under DD, which were greater during subjective night. The loss of circadian modulation under LL may be explained by the suspension of the circadian clock in these conditions. These results strongly suggest that the circatidal activity rhythm is driven by its own clock system, distinct from the circadian clock.