Okano Toshiyuki, Fukada Yoshitaka
Department of Biophysics and Biochemistry, Graduate School of Science, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033.
J Biochem. 2003 Dec;134(6):791-7. doi: 10.1093/jb/mvg221.
Many tissues in non-mammalian vertebrates contain both photoreceptors and circadian clock systems. Among these photosensitive clock structures, the chick pineal gland has been characterized in detail from cellular and molecular aspects of the clock oscillation and entrainment. Analyses of the pineal photic-input pathway revealed a phase-shifting mechanism mediated by activation of G11, one of the Gq-type G-proteins. A major photoreceptive molecule, pinopsin, likely triggers this pathway by transmitting the light signal to the circadian oscillator. In the chick pineal oscillator, the transcription/translation-based autoregulatory feedback loop is composed of positive and negative elements (clock gene products) that are homologous to those identified in mammals. In the molecular cycling, a CACGTG E-box located in the promoter region of the negative element genes plays a central role in the transcriptional regulation. The phase of the molecular cycling is modulated by many regulatory components, among which E4BP4 and extracellular signal-regulated kinase (ERK) are closely associated with the photic entrainment. A light-responsive element was found in the promoter region of the Pinopsin gene, and the element included a CACGTG E-box, suggesting a novel role of the E-box as a point of convergence of light and circadian signals. These observations together point to general and unique features of the chick pineal circadian system among animal clocks.
非哺乳类脊椎动物的许多组织都同时含有光感受器和生物钟系统。在这些光敏生物钟结构中,鸡的松果体已从生物钟振荡和同步的细胞及分子层面进行了详细的特征描述。对松果体光输入途径的分析揭示了一种由Gq型G蛋白之一G11激活介导的相移机制。一种主要的光感受分子视锥蛋白,可能通过将光信号传递给生物钟振荡器来触发这一途径。在鸡的松果体振荡器中,基于转录/翻译的自动调节反馈环由与哺乳动物中鉴定出的那些同源的正、负元件(生物钟基因产物)组成。在分子循环中,位于负元件基因启动子区域的CACGTG E盒在转录调控中起核心作用。分子循环的相位受到许多调节成分的调控,其中E4BP4和细胞外信号调节激酶(ERK)与光同步密切相关。在视锥蛋白基因的启动子区域发现了一个光反应元件,该元件包含一个CACGTG E盒,这表明E盒作为光信号和生物钟信号的汇聚点具有新的作用。这些观察结果共同指出了鸡松果体生物钟系统在动物生物钟中的普遍和独特特征。