Department of Biology, York University, Toronto, Ontario M3J 1P3, Canada.
J Comp Neurol. 2010 Apr 15;518(8):1264-82. doi: 10.1002/cne.22274.
The brain of larval Rhodnius prolixus releases neurohormones with a circadian rhythm, indicating that a clock system exists in the larval brain. Larvae also possess a circadian locomotor rhythm. The present paper is a detailed analysis of the distribution and axonal projections of circadian clock cells in the brain of the fifth larval instar. Clock cells are identified as neurons that exhibit circadian cycling of both PER and TIM proteins. A group of eight lateral clock neurons (LNs) in the proximal optic lobe also contain pigment-dispersing factor (PDF) throughout their axons, enabling their detailed projections to be traced. LNs project to the accessory medulla and thence laterally toward the compound eye and medially into a massive area of arborizations in the anterior protocerebrum. Fine branches radiate from this area to most of the protocerebrum. A second group of clock cells (dorsal neurons [DNs]), situated in the posterior dorsal protocerebrum, are devoid of PDF. The DNs receive two fine axons from the LNs, indicating that clock cells throughout the brain are integrated into a timing network. Two axons of the LNs cross the midline, presumably coordinating the clock networks of left and right sides. The neuroarchitecture of this timing system is much more elaborate than any previously described for a larval insect and is very similar to those described in adult insects. This is the first report that an insect timing system regulates rhythmicity in both the endocrine system and behavior, implying extensive functional parallels with the mammalian suprachiasmatic nucleus.
幼虫丽蝇的大脑会释放具有昼夜节律的神经激素,这表明在幼虫大脑中存在一个时钟系统。幼虫还具有昼夜节律的运动节律。本文详细分析了第五龄幼虫大脑中昼夜节律钟细胞的分布和轴突投射。钟细胞被鉴定为表现出 PER 和 TIM 蛋白昼夜循环的神经元。在近端视叶中,有一组 8 个侧时钟神经元 (LN),其轴突中始终含有色素分散因子 (PDF),从而能够追踪其详细的投射。LN 投射到 accessory medulla,然后向复眼的外侧和前脑的一个巨大的树突区域向内侧投射。从这个区域发出细分支到前脑的大部分区域。第二组时钟细胞 (背神经元 [DN]) 位于后脑背侧,不含 PDF。DN 接收来自 LN 的两个细轴突,表明整个大脑的时钟细胞都整合到一个计时网络中。LN 的两个轴突穿过中线,可能协调左右两侧的时钟网络。这个计时系统的神经结构比以前描述的任何幼虫昆虫都要复杂得多,与成年昆虫中描述的非常相似。这是第一个报告昆虫计时系统调节内分泌系统和行为的节律性的报告,暗示与哺乳动物视交叉上核有广泛的功能相似性。