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PubMed2005

Regulation of prokineticin 2 expression by light and the circadian clock.

BACKGROUND: The suprachiasmatic nucleus (SCN) contains the master circadian clock that regulates daily rhythms of many physiological and behavioural processes in mammals. Previously we have shown that prokineticin 2 (PK2) is a clock-controlled gene that may function as a critical SCN output molecule responsible for circadian locomotor rhythms. As light is the principal zeitgeber that entrains the circadian oscillator, and PK2 expression is responsive to nocturnal light pulses, we further investigated the effects of light on the molecular rhythm of PK2 in the SCN. In particular, we examined how PK2 responds to shifts of light/dark cycles and changes in photoperiod. We also investigated which photoreceptors are responsible for the light-induced PK2 expression in the SCN. To determine whether light requires an intact functional circadian pacemaker to regulate PK2, we examined PK2 expression in cryptochrome1,2-deficient (Cry1-/-Cry2-/-) mice that lack functional circadian clock under normal light/dark cycles and constant darkness. RESULTS: Upon abrupt shifts of the light/dark cycle, PK2 expression exhibits transients in response to phase advances but rapidly entrains to phase delays. Photoperiod studies indicate that PK2 responds differentially to changes in light period. Although the phase of PK2 expression expands as the light period increases, decreasing light period does not further condense the phase of PK2 expression. Genetic knockout studies revealed that functional melanopsin and rod-cone photoreceptive systems are required for the light-inducibility of PK2. In Cry1-/-Cry2-/- mice that lack a functional circadian clock, a low amplitude PK2 rhythm is detected under light/dark conditions, but not in constant darkness. This suggests that light can directly regulate PK2 expression in the SCN. CONCLUSION: These data demonstrate that the molecular rhythm of PK2 in the SCN is regulated by both the circadian clock and light. PK2 is predominantly controlled by the endogenous circadian clock, while light plays a modulatory role. The Cry1-/-Cry2-/- mice studies reveal a light-driven PK2 rhythm, indicating that light can induce PK2 expression independent of the circadian oscillator. The light inducibility of PK2 suggests that in addition to its role in clock-driven rhythms of locomotor behaviour, PK2 may also participate in the photic entrainment of circadian locomotor rhythms.

Animals↗
PubMed2006

Constant light disrupts the developing mouse biological clock.

The central biological clock of the brain, contained within the suprachiasmatic nuclei (SCN) of mammals, orchestrates an orderly "internal day" of physiology and behavior. The developing biological clock begins to respond to light at an early stage and a particular concern in humans is whether light exposure has disruptive effects on the developing biological clock of infants exposed to constant lighting conditions in neonatal intensive care units (NICUs). Worldwide, eighteen million, or 14%, of newborns estimated to be of low birth weight, are exposed to artificial lighting environments in hospital nurseries annually. Here, we have tested whether constant light (LL) exposure disrupts the developing biological clock of mice, using a circadian reporter transgenic mouse model in which the organization of the central biological clock can be assayed by real-time gene expression imaging. We now find that LL has both acute and long-term disruptive effects on developing biological clocks and that cyclic lighting conditions are critical for developing circadian clocks to coordinate their molecular circadian mechanisms. This suggests that, from the perspective of developing circadian organization in humans, cyclic light conditions in NICUs are likely to be most appropriate for infants.

Animals↗
PubMed2005

Rhythmic messenger ribonucleic acid expression of clock genes and adipocytokines in mouse visceral adipose tissue.

Various peripheral tissues show circadian rhythmicity, which is generated at the cellular level by their own core oscillators that are composed of transcriptional/translational feedback loops involving a set of clock genes. Although the circulating levels of some adipocytokines, i.e. bioactive substances secreted by adipocytes, are on a 24-h rhythmic cycle, it remains to be elucidated whether the clock gene system works in adipose tissue. To address this issue, we investigated the daily mRNA expression profiles of the clock genes and adipocytokines in mouse perigonadal adipose tissues. In C57BL/6J mice, all transcript levels of the clock genes (Bmal1, Per1, Per2, Cry1, Cry2, and Dbp) and adipocytokines (adiponectin, resistin, and visfatin) clearly showed 24-h rhythms. On the other hand, the rhythmic expression of these genes was mildly attenuated in obese KK mice and greatly attenuated in more obese, diabetic KK-A(y) mice. Obese diabetes also diminished the rhythmic expression of the clock genes in the liver. Interestingly, a 2-wk treatment of KK and KK-A(y) mice with pioglitazone impaired the 24-h rhythmicity of the mRNA expression of the clock genes and adipocytokines despite the antidiabetic effect of the drug. In contrast, pioglitazone improved the attenuated rhythmicity in the liver. These findings suggest that the intracellular clock gene system acts in visceral adipose tissues as well as liver and is influenced by the conditions of obesity/type 2 diabetes and pioglitazone treatment.

Adipocytes↗
PubMed2006

Circadian Clock mutation in dams disrupts nursing behavior and growth of pups.

To understand the role of the circadian molecular clock in mouse reproduction, we investigated the daily rhythms associated with nursing and pup growth in Clock-mutant mice maintained under light-dark housing conditions. The daily rhythm associated with the maternal behavior of crouching had a strong diurnal peak and two weak nocturnal peaks in wild-type dams, whereas homozygotes (Clock/Clock) exhibited no significant peaks in activity. Wild-type, but not Clock-mutant, dams showed high, rhythmic levels of prolactin content in serum that corresponded with crouching. Pup body weight increased at a significantly slower rate in Clock-mutant dams compared with wild-type dams under all experimental conditions when the pups ranged from 10-15 in number. Heterozygote dams equally bred wild-type, heterozygote, or homozygote pups. The amount of milk secreted from dams, as calculated by the increase in pup body weight through suckling, was lower in Clock-mutant mothers vs. wild-type mice. When Clock-mutant dams gave birth to more than 10 pups, survival was poor for offspring until the time of weaning. The present results demonstrate that Clock mutation disrupts daily maternal behavior and the growth and survival rate of pups, especially with the breeding of more than 10 pups.

Animals↗
PubMed1998

[Physiological, pharmacological and molecular aspects of mammalian biological clocks].

Circadian rhythm is an endogenous rhythm that persists in constant conditions with a period of nearly but not identical to 24 hr. Under natural conditions, the circadian clock is precisely entrained to the daily (24 hr) cycle, because environmental stimulus (especially light) induces a phase shift of the clock. In mammals, the suprachiasmatic nucleus (SCN) of the hypothalamus has been shown to be the primary pacemaker that drives daily rhythms of behavioral and physiological activity. Photic information is conveyed from the retina to the SCN directly by the retinohypothalamic tract (RHT) and indirectly by the geniculo-hypothalamic tract (GHT). The transmitter of the RHT is glutamate, while the GHT is GABA and neuropeptide Y. Serotonergic innervation from the median raphe and melatonin from the pineal body are likely to provide non-photic information to the SCN. Single gene mutations that dramatically alter circadian phenotype were found in the hamster (tau) and mouse (clock). Moreover, the homologous genes of the Drosophila clock gene, per, were found in mammals and the homologue of the mammalian clock was found in Drosophila. These data suggest that the some constitutes of the biological clock may be conserved between Drosophila and mammals, and a transcription-translation feedback loop involving some clock gene products may be a oscillator itself.

Animals↗
PubMed2005

[Circadian rhythms and light responses of clock gene and arylalkylamine N-acetyltransferase gene expressions in the pineal gland of rats].

This study was to investigate the circadian rhythms and light responses of Clock gene and arylalkylamine N-acetyltransferase (NAT) gene expressions in the rat pineal gland under the 12 h-light : 12 h-dark cycle condition (LD) and constant darkness (DD). Sprague-Dawley rats housed under the light regime of LD (n=36) for 4 weeks and of DD (n=36) for 8 weeks were sampled for the pineal gland once a group (n=6) every 4 h in a circadian day. The total RNA was extracted from each sample and the semiquantitative reverse transcription polymerase chain reaction (RT-PCR) was used to determine the temporal changes in mRNA levels of Clock and NAT genes during different circadian times or zeitgeber times. The data were analysed by the cosine function software, Clock Lab software and the amplitude F test was used to reveal the circadian rhythm. The main results obtained are as follows. (1) In DD or LD condition, both of Clock and NAT genes mRNA levels in the pineal gland showed robust circadian oscillation (P< 0.05) with the peak at the subjective night or at night-time. (2) In comparison with DD regime, the amplitudes and the mRNA levels at peaks of Clock and NAT genes expressions in LD in the pineal gland were significantly reduced (P< 0.05). (3) In DD or LD condition, the circadian expressions of NAT gene were similar in pattern to those of Clock gene in the pineal gland (P> 0.05). These findings suggest that the expressions of Clock and NAT genes in the pineal gland not only show remarkably synchronous endogenous circadian rhythmic changes, but also response to the ambient light signal in a reduced manner.

Animals↗
PubMed2002

Comparing clock tests for dementia screening: naïve judgments vs formal systems--what is optimal?

BACKGROUND: Clock drawing tests (CDTs) vary in format, scoring, and complexity. Herein, we compared the dementia screening performance of seven CDT scoring systems and the judgements of untrained raters. METHODS: 80 clock drawings by subjects of known dementia status were selected, 20 from each of four categories (Consortium to Establish a Registry for Alzheimer's disease [CERAD] defined normal, mild, moderate, and severe abnormality). An expert rater scored all clocks using published criteria for seven systems. Additionally, 20 naïve raters judged clocks as either normal or abnormal, without formal instructions. Clocks were then classified by drawers' dementia status for comparison of dementia detection across systems. RESULTS: Naïve and formal CDT systems showed 90-100% agreement in CERAD normal, moderate and severe categories, but poor agreement (mean = 39%) for mildly impaired clocks. When CDT systems were compared for accurate dementia classification, the Mendez and CERAD systems correctly identified the greatest proportion of subjects (84-85%), and Wolf-Klein the smallest (58%). The better systems correctly identified> 70% of mildly demented individuals (CDR = 1). In contrast, medical records from patients' personal physicians correctly identified only 24% of the mildly demented. Strikingly, naïve raters' CDT judgements were as effective as five of the seven CDT systems in dementia identification. CONCLUSIONS: While the Mendez system was the most accurate overall, it was not significantly better than CERAD, which had simpler scoring rules. Untrained raters discriminated normal from abnormal clocks with acceptable accuracy for community screening purposes. Results suggest that, if used, most CDT systems would improve personal physicians' dementia recognition in difficult to detect mildly demented subjects.

Aged↗
PubMed2002

Clock drawing test in mild and moderate dementia of the Alzheimer's type: a comparative and correlation study.

OBJECTIVES: (a) To compare two different clock drawing tests (CDTs) in mild and moderate dementia of the Alzheimer's type (DAT); (b) To examine presumed correlation between these CDTs and some demographic, cognitive and activities of daily living (ADL) variables in mild and moderate DAT. METHODS: Cross-sectional study. Psychogeriatric outpatient clinic. 49 DAT patients, total; 26-mild, 23-moderate, mean age 77.8 and 80.6, respectively. Evaluations included the Mini-Mental State Examination (MMSE), the Cambridge Cognitive Examination (CAMCOG), the Instrumental Activities of Daily Living Scale (IADL), and a Basic Activities of Daily Living (BADL)-dressing subscale. Severity of dementia was determined with the Clinical Dementia Rating (CDR). Each clock was blindly scored by the same investigator, according to Shulman's and Freedman's methods. RESULTS: Mild and moderate DAT groups were similar in age, gender and education. Performance on Shulman's clock was similar between groups while moderate DAT subjects performed significantly worse on Freedman's clock compared to mild DAT patients. Both clocks correlated highly in mild and moderate DAT. CDT scores correlated significantly with age and education only in mild DAT. Neither clock correlated with ADLs in either stage of dementia severity. CDTs correlated with the MMSE score, and the CAMCOG score in mild DAT, and only with the CAMCOG score in moderate DAT. These correlations were still significant after controlling for age and education. CONCLUSIONS: Different aspects of cognition and dementia severity are reflected depending on how a clock drawing is scored. Some scoring systems may have greater sensitivity than others in monitoring progression of cognitive deterioration. Correlation between different CDTs and the variables studied (demographic, cognitive, ADLs), when present, is not ubiqitous and changes with the dementia severity.

Activities of Daily Living↗
PubMed2005

Atypical patterns of circadian clock gene expression in human peripheral blood mononuclear cells.

Circadian ( approximately 24 h) rhythms in physiology and behaviour are observed in all mammals, including humans. These rhythms are generated by circadian clocks located in the hypothalamus and also in most peripheral tissues. Clock genes are essential components of circadian clocks, and mutations or polymorphisms within several of them have been associated with circadian disorders in humans. However, information about human clock gene expression has remained very limited. Peripheral blood mononuclear cells (PBMCs) represent an ideal material to investigate non-invasively the human clock at the molecular level. In the present study, we analysed the expression of three key clock genes, PER2, BMAL1 and REV-ERBalpha in PBMCs from ten healthy humans over a 24-h cycle. PER2 and BMAL1 were found to oscillate throughout the light-dark cycle in all subjects. Interestingly, despite normal melatonin and cortisol secretion patterns, two groups of subjects could be distinguished with significantly different mean PER2 and BMAL1 acrophases. BMAL1 oscillated with approximately the same phase as PER2, instead of being anti-phasic as anticipated from data previously obtained in other peripheral tissues. Furthermore, this unusual phase relationship of PER2 and BMAL1 in human PBMCs was associated with a constant expression of REV-ERBalpha, a crucial regulator of BMAL1, which is highly rhythmic in many other systems. These results reveal the existence of different chronotypes of clock gene expression patterns and suggest specific regulatory mechanisms in human PBMCs.

ARNTL Transcription Factors↗
PubMed2004

Flies by night: Effects of changing day length on Drosophila's circadian clock.

In Drosophila, two intersecting molecular loops constitute an autoregulatory mechanism that oscillates with a period close to 24 hr. These loops touch when proteins from one loop, PERIOD (PER) and TIMELESS (TIM), repress the transcription of their parent genes, period (per) and timeless (tim), by blocking positive transcription factors from the other loop. The arrival of PER and TIM into the nucleus of a clock cell marks the timing of this interaction between the two loops; thus, control of PER:TIM nuclear accumulation is a central component of the molecular model of clock function. If a light pulse occurs early in the night as the heterodimer accumulates in the nucleus of clock cells, TIM is degraded, PER is destabilized, and clock time is delayed. Alternatively, if TIM is degraded during the later part of the night, after peak accumulation, clock time advances. Current models state that the effect of a light pulse depends on the state of the PER:TIM oscillation, which turns on the changing levels of TIM. However, previous studies have shown that light:dark (LD) regimes mimicking seasonal changes cause behavioral adjustments while altering clock gene expression. This should be reflected in the adjustment of PER and TIM dynamics. We manipulated LD cycles to assess the effects of altered day length on PER and TIM dynamics in clock cells within the central brain as well as light-induced resetting of locomotor rhythms.

Animals↗
PubMed2006

Serotonergic pre-treatments block in vitro serotonergic phase shifts of the mouse suprachiasmatic nucleus circadian clock.

The suprachiasmatic nucleus (SCN) contains a circadian clock that maintains its time-generating and phase-modulating capacities in vitro. Previous studies report clear differences in the ability of serotonergic stimuli to phase-shift the SCN clock when applied directly to the SCN either in vivo or in vitro: while mice and rat circadian clocks are readily phase-shifted by serotonin (5-HT) or 5-HT agonists applied in vitro, hamster and mice circadian clocks respond inconsistently to 5-HT agonists injected directly into the SCN in vivo. Here we have investigated one possible explanation for these differences: that the SCN isolated in vitro experiences reduced endogenous 5-HT signaling, which increases clock sensitivity to subsequent 5-HT stimulation. For these experiments we treated mouse SCN brain slices with low concentrations of compounds that increase serotonin signaling: 5-HT, a 5-HT agonist (8-OH-DPAT), the 5-HT precursor, l-tryptophan, or the 5-HT re-uptake inhibitor, fluoxetine. Pretreatment with each of these substances completely blocked subsequent phase-shifts induced by mid-subjective day treatment with either 5-HT or 8-OH-DPAT, while they did not block phase-shifts induced by the adenylate cyclase activator, forskolin. Time-course data on l-tryptophan-induced inhibition are consistent with this treatment inducing receptor internalization, while timing of the recovery from inhibition is consistent with receptor reinsertion. Together these data support the hypothesis that SCN clock sensitivity to serotonergic phase modulation is affected by the amount of prior serotonin signaling present in the SCN, and that this signaling alters the density of surface 5-HT receptors on SCN clock neurons.

8-Hydroxy-2-(di-n-propylamino)tetralin↗
PubMed1999

Insect circadian clocks: is it all in their heads?

Circadian rhythms are ubiquitous in living organisms, synchronizing life functions at the biochemical, physiological, and behavioral levels. The rhythm-generating mechanisms, collectively known as circadian clocks, are not fully understood in any organism. Research in the fruit fly Drosophila has led to the identification of several clock genes that are involved in the function of the brain-centered clock, which controls behavioral rhythms of adult flies. With the use of clock genes as markers, putative circadian clocks were mapped in the fly peripheral organs and shown to be independent from clocks located in the brain. A homologue of fruit fly period gene has been identified in moths and other insects, allowing investigations of this gene's role in known insect rhythms. This approach may increase our understanding of how circadian clocks are organized into the circadian system that orchestrates temporal integration of life processess in insects.

Journal Article↗
PubMed2005

An optical lattice clock.

The precision measurement of time and frequency is a prerequisite not only for fundamental science but also for technologies that support broadband communication networks and navigation with global positioning systems (GPS). The SI second is currently realized by the microwave transition of Cs atoms with a fractional uncertainty of 10(-15) (ref. 1). Thanks to the optical frequency comb technique, which established a coherent link between optical and radio frequencies, optical clocks have attracted increasing interest as regards future atomic clocks with superior precision. To date, single trapped ions and ultracold neutral atoms in free fall have shown record high performance that is approaching that of the best Cs fountain clocks. Here we report a different approach, in which atoms trapped in an optical lattice serve as quantum references. The 'optical lattice clock' demonstrates a linewidth one order of magnitude narrower than that observed for neutral-atom optical clocks, and its stability is better than that of single-ion clocks. The transition frequency for the Sr lattice clock is 429,228,004,229,952(15) Hz, as determined by an optical frequency comb referenced to the SI second.

Journal Article↗
PubMed2003

Nucleocytoplasmic shuttling and phosphorylation of BMAL1 are regulated by circadian clock in cultured fibroblasts.

BACKGROUND: Recent discoveries of clock proteins have unveiled an important part of the mammalian circadian clock mechanism. However, the molecular clockwork that cause these fundamental feedback loops to stably oscillate with a approximately 24 h-periodicity remain unclear. RESULTS: Serum-shocked fibroblasts were used as a cellular clock model. Circadian changes in the subcellular localization and phosphorylation of BMAL1 protein in these cells were assessed by immunocytochemistry and immunoblotting. A significant time lag between Bmal1 transcription and the cytoplasmic/nuclear accumulation of BMAL1 was observed. After its nuclear accumulation, BMAL1 accumulated in the cytoplasm again, mainly by nucleoexport, before the increase of Bmal1 transcripts. Nuclear accumulation of BMAL1 matched nuclear accumulation of CLOCK and the peak of Per1 transcription. Nuclear BMAL1 was gradually phosphorylated and then dephosphorylated in a temporally regulated manner, although cytoplasmic BMAL1 was not. In serum-shocked mCry1/mCry2 (CRY)-deficient fibroblasts, which lack a functional clock, both the cytoplasmic and nuclear BMAL1 were only present as hyperphosphorylated forms and their circadian nucleocytoplasmic shuttling was absent. CONCLUSIONS: We propose that the nucleocytoplasmic shuttling and phosphorylation states of BMAL1 are regulated by circadian clock, and that this temporally regulated and time-delayed nuclear entry of BMAL1 is important in the maintenance of a stably oscillating clock.

ARNTL Transcription Factors↗
PubMed2002

Distinct regulation of CAB and PHYB gene expression by similar circadian clocks.

Phytochrome B (phyB) is a major phytochrome active in light-grown plants. The circadian clock controls the expression of the PHYB gene. We have used the luciferase reporter gene (LUC) to monitor the rhythmic expression of PHYB in photoreceptor and clock-associated mutant backgrounds. Surprisingly, we found that PHYB and CAB expression have different free-running periods, indicating that separate circadian clocks control these genes. The effects of mutations show that the clocks share common components. This suggests that they are copies of the same clock mechanism in different locations, most likely in different cell layers. Furthermore, we show that phyB is required for a negative feedback loop that strongly antagonises the expression of PHYB. Compared to a system with only one clock, this regulatory complexity might allow the phase of peak expression for one clock-controlled gene to alter, relative to other genes or to changing environmental conditions.

Arabidopsis↗
PubMed2002

Loss of circadian clock function decreases reproductive fitness in males of Drosophila melanogaster.

Circadian coordination of life functions is believed to contribute to an organism's fitness; however, such contributions have not been convincingly demonstrated in any animal. The most significant measure of fitness is the reproductive output of the individual and species. Here we examined the consequences of loss of clock function on reproductive fitness in Drosophila melanogaster with mutated period (per(0)), timeless (tim(0)), cycle (cyc(0)), and Clock (Clk(Jrk)) genes. Single mating among couples with clock-deficient phenotypes resulted in approximately 40% fewer progeny compared with wild-type flies, because of a decreased number of eggs laid and a greater rate of unfertilized eggs. Male contribution to this phenotype was demonstrated by a decrease in reproductive capacity among per(0) and tim(0) males mated with wild-type females. The important role of clock genes for reproductive fitness was confirmed by reversal of the low-fertility phenotype in flies with rescued per or tim function. Males lacking a functional clock showed a significant decline in the quantity of sperm released from the testes to seminal vesicles, and these tissues displayed rhythmic and autonomous expression of clock genes. By combining molecular and physiological approaches, we identified a circadian clock in the reproductive system and defined its role in the sperm release that promotes reproductive fitness in D. melanogaster.

Animals↗
PubMed1996

Circadian clock-controlled genes isolated from Neurospora crassa are late night- to early morning-specific.

An endogenous circadian biological clock controls the temporal aspects of life in most organisms, including rhythmic control of genes involved in clock output pathways. In the fungus Neurospora crassa, one pathway known to be under control of the clock is asexual spore (conidia) development. To understand more fully the processes that are regulated by the N. crassa circadian clock, systematic screens were carried out for genes that oscillate at the transcriptional level. Time-of-day-specific cDNA libraries were generated and used in differential screens to identify six new clock-controlled genes (ccgs). Transcripts specific for each of the ccgs preferentially accumulate during the late night to early morning, although they vary with respect to steady-state mRNA levels and amplitude of the rhythm. Sequencing of the ends of the new ccg cDNAs revealed that ccg-12 is identical to N. crassa cmt encoding copper metallothionein, providing the suggestion that not all clock-regulated genes in N. crassa are specifically involved in the development of conidia. This was supported by finding that half of the new ccgs, including cmt(ccg-12), are not transcriptionally induced by developmental or light signals. These data suggest a major role for the clock in the regulation of biological processes distinct from development.

Chromosome Mapping↗
PubMed2006

The circadian clock within the cardiomyocyte is essential for responsiveness of the heart to fatty acids.

Cells/organs must respond both rapidly and appropriately to increased fatty acid availability; failure to do so is associated with the development of skeletal muscle and hepatic insulin resistance, pancreatic beta-cell dysfunction, and myocardial contractile dysfunction. Here we tested the hypothesis that the intrinsic circadian clock within the cardiomyocytes of the heart allows rapid and appropriate adaptation of this organ to fatty acids by investigating the following: 1) whether circadian rhythms in fatty acid responsiveness persist in isolated adult rat cardiomyocytes, and 2) whether manipulation of the circadian clock within the heart, either through light/dark (L/D) cycle or genetic disruptions, impairs responsiveness of the heart to fasting in vivo. We report that both the intramyocellular circadian clock and diurnal variations in fatty acid responsiveness observed in the intact rat heart in vivo persist in adult rat cardiomyocytes. Reversal of the 12-h/12-h L/D cycle was associated with a re-entrainment of the circadian clock within the rat heart, which required 5-8 days for completion. Fasting rats resulted in the induction of fatty acid-responsive genes, an effect that was dramatically attenuated 2 days after L/D cycle reversal. Similarly, a targeted disruption of the circadian clock within the heart, through overexpression of a dominant negative CLOCK mutant, severely attenuated induction of myocardial fatty acid-responsive genes during fasting. These studies expose a causal relationship between the circadian clock within the cardiomyocyte with responsiveness of the heart to fatty acids and myocardial triglyceride metabolism.

Animals↗