Kaneko M, Hamblen M J, Hall J C
Department of Biology, Brandeis University, Waltham, MA 02454 USA.
J Biol Rhythms. 2000 Feb;15(1):13-30. doi: 10.1177/074873040001500103.
Phases of circadian locomotor activity rhythms of adult Drosophila reared in constant darkness have been shown to be set by a light stimulus delivered as early as the first-instar larval stage. This implies that a circadian clock functions continuously throughout postembryonic development. The clock genes period (per) and timeless (tim) are expressed cyclically in the larval central nervous system of Drosophila, and daily oscillations of per expression persist throughout metamorphosis in a group of cells, which gives rise to the pacemaker cells underlying locomotor activity rhythms of adults. Therefore, PER and TIM cyclings in these neurons may be responsible for the phenomenon of "larval time-memory." In the absence of any evidence for the involvement of these genes in such a developmental clock, and because circadian-pacemaker functions are underanalyzed in terms of the functions during development, the authors tested the time-memory of a fast-clock period mutant. They show that dark-reared perS mutant individuals as well as wild-type flies can be entrained as larvae and that a brief light pulse given to such entrained larvae can induce phase shifts in animals of either genotype. However, the direction and magnitude of phase shifts were different between wild type and perS, suggesting that a clock under the control of period gene participates in the regulation of developmental time-memory. The authors show that the relevant clock can be entrained by two light input pathways, one involving the phospholipase C encoded by the norpA gene, the other mediated by the blue-light receptor cryptochrome. Phase shifts of molecular oscillations during the larval stage were smaller than those measured by adult behavior, suggesting molecularly transient responses during development.
在持续黑暗中饲养的成年果蝇的昼夜节律运动活动节律阶段已被证明是由早在一龄幼虫阶段给予的光刺激设定的。这意味着昼夜节律钟在胚胎后发育过程中持续发挥作用。生物钟基因周期(per)和无时间(tim)在果蝇幼虫中枢神经系统中周期性表达,并且per表达的每日振荡在变态过程中在一组细胞中持续存在,这些细胞产生了成年动物运动活动节律的起搏细胞。因此,这些神经元中的PER和TIM循环可能是“幼虫时间记忆”现象的原因。由于没有证据表明这些基因参与这样的发育时钟,并且由于昼夜节律起搏器功能在发育过程中的功能方面未得到充分分析,作者测试了快速时钟周期突变体的时间记忆。他们表明,在黑暗中饲养的perS突变体个体以及野生型果蝇在幼虫期都可以被同步,并且给予这样同步的幼虫一个短暂的光脉冲可以诱导两种基因型动物的相位变化。然而,野生型和perS之间的相位变化方向和幅度不同,这表明受周期基因控制的时钟参与了发育时间记忆的调节。作者表明,相关的时钟可以通过两种光输入途径被同步,一种涉及由norpA基因编码的磷脂酶C,另一种由蓝光受体隐花色素介导。幼虫期分子振荡的相位变化小于通过成年行为测量的相位变化,这表明发育过程中分子的短暂反应。