e-book Lithium Clockwork

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QuantumKraken July 27, , am 2. C,D, Period and amplitude analysis. The amplitude changes were analyzed based on the first peak after treatment in the baseline corrected data.

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E, Bright field and bioluminescence imaging of lung slices. Peak and trough of oscillations were determined based on the bioluminescent intensity of the CCD camera recording. White arrowheads indicate bronchiole and blood vessel. Cultured cells harbour self-sustained and cell-autonomous circadian clocks that resemble the molecular oscillations in the brain, therefore providing an amenable in vitro clock model to perform more detailed mechanistic studies [31] , [32].

Similar responses to lithium were also observed in isolated lung fibroblasts Fig. There were significant dose-dependent period lengthening effects 0.

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In contrast to ectopic SCN tissues, there were no significant effects on either period or amplitude of PERLUC oscillation in lung slices or fibroblasts treated with lower doses 1 or 2 mM of lithium, indicating likely tissue differences in sensitivity to lithium. Black arrow indicates time of treatment. F, Amplitude analysis for data in D and E. In contrast to the period lengthening effect of lithium, BIO significantly shortened circadian period in lung fibroblasts by 0.

No significant additive effects on amplitude of PER2 expression were observed compared to lithium treatment alone lithium alone, 2. These findings indicate that although GSK3 inhibition cannot explain the period lengthening effect of lithium, it may play a role in the increase of PERLUC amplitude upon lithium treatment. To investigate how lithium impacts on the dynamics of other clock genes, we next used Rat1 cells stably transfected with Bmalluc reporter [25].

Lithium lengthened period of Bmalluc transcriptional rhythms in a dose-dependent manner, but there was no significant elevation of Bmal1 -driven bioluminescence level or oscillation amplitude Fig. In cultured lung slices, a similar analysis revealed no significant change in half-life following 10 mM lithium treatment 2. If lithium alters PER2 stability, we would predict that cells with different decay rate of PER2 may respond differently to lithium.

Top panel, Raw traces; bottom panel, baseline corrected data. To calculate a relative change the average of the 3 replicate CT values for each sample was used. We next tested whether lithium may directly augment endogenous Per2 transcription, leading to elevated PER2 protein expression. These results therefore suggest that lithium elicits induction of Per2 transcription, but does not alter the degradation of PER2 protein in cells.

Lithium salt is considered to be one of the most effective drug treatments for long term mood stabilization in BPD patients. However, there is also large inter-individual variability in responses to lithium, and it can cause serious side effects. It is therefore important to understand the pharmacological actions and molecular targets.

There is compelling evidence that BPD is associated with disruptions in circadian rhythms [2] , [34] , [35] , and lithium has been shown to robustly lengthen behavioral circadian period in many species [4]. Consistent with earlier reports, our results demonstrate that lithium decelerates circadian behavioural rhythms and molecular pacemaking in the brain and peripheral tissues and cells. However, it is unclear whether lithium has other effects on the molecular circadian clockwork in addition to the altered circadian clock period.

In addition to the well-established period lengthening effect, we show here that the oscillation amplitude of the PER2 protein rhythms in both central SCN and peripheral clocks lung and lung fibroblasts is significantly augmented by lithium. There are at least three possible explanations. Finally, lithium may regulate Per2 at transcriptional or post-transcriptional level.

This hypothesis is supported by our data, and by earlier reports which showed that lithium treatment selectively increased Per2 mRNA levels in serum-shocked NIH 3T3 cells [19] , [36]. Further studies are required to understand whether and how increased transcription of Per2 leads to enhanced amplitude of PERLUC rhythms, as well as to explore the additional post-transcriptional or post-translational mechanisms.

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These observations in HEK T cells provide a potential functional explanation for the actions of lithium on the circadian clockwork. However, our results using a Bmalluc reporter cell line that has RORE elements in the Bmal1 promoter did not show significant alteration of Bmalluc bioluminescence upon lithium treatment. Consistent with our observations, studies using Northern blot analysis over a time course of 48 hours revealed no significant increase of endogenous Bmal1 mRNA in NIH 3T3 cells upon lithium treatment [19].

Use of different cell line models may account for the discrepancies observed.

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An increase in amplitude by lithium independent of GSK3 inhibition would predict that combined treatment of lithium and GSK3 inhibitor should have additive effects. This hypothesis is not supported by our data, since fibroblast cells treated with both BIO and lithium demonstrated no further significant additive effects on amplitude of PER2 expression, indicating that inhibition of GSK3 may be the causal mechanism for induction of PER2 protein rhythms by lithium. The period effect, on the other hand, may involve additional mechanisms, as also suggested by Hirota et al [20].

Thus, delayed nuclear entry upon lithium treatment has been proposed as one potential mechanism for lithium-induced period lengthening. Within the molecular oscillator, PER2 acts as the negative regulator which suppresses transactivation of E-box regulated genes, including its own transcription. This may consequently lead to the induction of Per2 and high amplitude oscillation of PER2 proteins.

Similar doses have been used in other studies to reveal the effects of lithium on circadian clocks [12] , [19] , [20] , [36]. In fact, lithium in cells may trigger multiple distinct pathways. Therefore, further investigation is warranted to identify additional mechanisms. Data were subjected to baseline correction which removed the first 12 hours data.

Black arrow indicates time of lithium or BIO treatment. Cells were first treated with the AKT inhibitor A as indicated by the arrow. After 1 hr, cells were treated with lithium 20 mM. A, raw data; B, baseline correction of raw data. Data are presented from a single experiment. We thank Professor J. Competing Interests: The authors have declared that no competing interests exist. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

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National Center for Biotechnology Information , U. PLoS One. Published online Mar Andrew S. Shin Yamazaki, Editor.

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Author information Article notes Copyright and License information Disclaimer. Received Oct 21; Accepted Feb Copyright Li et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are properly credited.

This article has been cited by other articles in PMC. Abstract Lithium salt has been widely used in treatment of Bipolar Disorder, a mental disturbance associated with circadian rhythm disruptions. Statistical analysis Statistical analysis was performed based on single experiments with several samples biological replicates. Results Lithium lengthens circadian period in behavioural rhythms Lithium is known to lengthen circadian free-running rhythms in a variety of species, including humans and rodents.

Open in a separate window. Figure 1. Lithium lengthens period for behavioral rhythms and alters molecular oscillations in the SCN. Lithium impacts on clock period and PERLUC oscillation amplitude in the central and peripheral clockwork Lithium treatment of dispersed SCN neurons lengthens the circadian period of firing rate rhythms in a dose-dependent manner [5]. Figure 2. Figure 3. Figure 4. Lithium lengthens period of Bmalluc oscillation in Rat1 cells.

Figure 5. Discussion Lithium salt is considered to be one of the most effective drug treatments for long term mood stabilization in BPD patients. TIF Click here for additional data file. Acknowledgments We thank Professor J. Footnotes Competing Interests: The authors have declared that no competing interests exist. References 1. New York: Oxford University Press; McClung CA.

Circadian genes, rhythms and the biology of mood disorders.