Graduate School of Natural Science and Technology, Kanazawa University, Kakuma-machi, Kanazawa, Japan.
PLoS Comput Biol. 2019 Feb 19;15(2):e1006787. doi: 10.1371/journal.pcbi.1006787. eCollection 2019 Feb.
Negative feedback loops (NFLs) for circadian clocks include light-responsive reactions that allow the clocks to shift their phase depending on the timing of light signals. Phase response curves (PRCs) for light signals in various organisms include a time interval called a dead zone where light signals cause no phase shift during daytime. Although the importance of the dead zone for robust light entrainment is known, how the dead zone arises from the biochemical reactions in an NFL underlying circadian gene expression rhythms remains unclear. In addition, the observation that the light-responsive reactions in the NFL vary between organisms raises the question as to whether the mechanism for dead zone formation is common or distinct between different organisms. Here we reveal by mathematical modeling that the saturation of a biochemical reaction in repressor synthesis in an NFL is a common mechanism of daytime dead zone generation. If light signals increase the degradation of a repressor protein, as in Drosophila, the saturation of repressor mRNA transcription nullifies the effect of light signals, generating a dead zone. In contrast, if light signals induce the transcription of repressor mRNA, as in mammals, the saturation of repressor translation can generate a dead zone by cancelling the influence of excess amount of mRNA induced by light signals. Each of these saturated reactions is located next to the light-responsive reaction in the NFL, suggesting a design principle for daytime dead zone generation.
负反馈环 (NFL) 是生物钟的一部分,包括光响应反应,使生物钟能够根据光信号的时间调整相位。各种生物的光信号相位反应曲线 (PRC) 包括一个称为盲区的时间间隔,在此期间,光信号在白天不会引起相位移动。尽管已经知道盲区对于稳健的光适应很重要,但 NFL 中生物钟基因表达节律的生化反应如何产生盲区仍然不清楚。此外,观察到 NFL 中的光响应反应在不同生物体之间存在差异,这就提出了一个问题,即盲区形成的机制在不同生物体之间是否普遍存在或独特。在这里,我们通过数学建模揭示了 NFL 中抑制物合成的生化反应饱和是白天盲区产生的常见机制。如果光信号增加了抑制蛋白的降解,如在果蝇中,那么抑制物 mRNA 转录的饱和会抵消光信号的影响,产生盲区。相比之下,如果光信号诱导抑制物 mRNA 的转录,如在哺乳动物中,那么抑制物翻译的饱和可以通过抵消光信号诱导的过量 mRNA 的影响来产生盲区。这些饱和反应中的每一个都位于 NFL 中的光响应反应旁边,这表明了产生白天盲区的设计原则。