Doi M, Nakajima Y, Okano T, Fukada Y
Department of Biophysics and Biochemistry, Graduate School of Science, University of Tokyo, Japan.
Proc Natl Acad Sci U S A. 2001 Jul 3;98(14):8089-94. doi: 10.1073/pnas.141090998. Epub 2001 Jun 26.
The chicken pineal gland contains the autonomous circadian oscillator together with the photic-input pathway. We searched for chicken pineal genes that are induced by light in a time-of-day-dependent manner, and isolated chicken homolog of bZIP transcription factor E4bp4 (cE4bp4) showing high similarity to vrille, one of the Drosophila clock genes. cE4bp4 was expressed rhythmically in the pineal gland with a peak at very early (subjective) night under both 12-h light/12-h dark cycle and constant dark conditions, and the phase was nearly opposite to the expression rhythm of cPer2, a chicken pineal clock gene. Luciferase reporter gene assays showed that cE4BP4 represses cPer2 promoter through a E4BP4-recognition sequence present in the 5'-flanking region, indicating that cE4BP4 can down-regulate the chick pineal cPer2 expression. In vivo light-perturbation studies showed that the prolongation of the light period to early subjective night maintained the high level expression of the pineal cE4bp4, and presumably as a consequence delayed the onset of the induction of the pineal cPer2 expression in the next morning. These light-dependent changes in the mRNA levels of the pineal cE4bp4 and cPer2 were followed by a phase-delay of the subsequent cycles of cE4bp4/cPer2 expression, suggesting that cE4BP4 plays an important role in the phase-delaying process as a light-dependent suppressor of cPer2 gene.
鸡松果体包含自主昼夜节律振荡器以及光输入通路。我们寻找以昼夜时间依赖方式被光诱导的鸡松果体基因,并分离出bZIP转录因子E4bp4的鸡同源物(cE4bp4),它与果蝇生物钟基因之一的vrille高度相似。在12小时光照/12小时黑暗周期和持续黑暗条件下,cE4bp4在松果体中呈节律性表达,在非常早的(主观)夜间达到峰值,且该相位与鸡松果体生物钟基因cPer2的表达节律几乎相反。荧光素酶报告基因检测表明,cE4BP4通过存在于5'-侧翼区域的E4BP4识别序列抑制cPer2启动子,这表明cE4BP4可以下调鸡松果体中cPer2的表达。体内光扰动研究表明,将光照期延长至早期主观夜间可维持松果体cE4bp4的高水平表达,并且推测这一结果延迟了次日早晨松果体cPer2表达诱导的起始。松果体cE4bp4和cPer2 mRNA水平的这些光依赖性变化之后是cE4bp4/cPer2表达后续周期的相位延迟,这表明cE4BP4作为cPer2基因的光依赖性抑制因子在相位延迟过程中起重要作用。