Corresponding attributes of neural development and function suggest arthropod and vertebrate brains may have an evolutionarily conserved organization. However, the underlying mechanisms have remained elusive. Here, we identify a gene regulatory and character identity network defining the deutocerebral–tritocerebral boundary (DTB) in Drosophila. This network comprises genes homologous to those directing midbrain-hindbrain boundary (MHB) formation in vertebrates and their closest chordate relatives. Genetic tracing reveals that the embryonic DTB gives rise to adult midbrain circuits that in flies control auditory and vestibular information processing and motor coordination, as do MHB-derived circuits in vertebrates. DTB-specific gene expression and function are directed by cis-regulatory elements of developmental control genes that include homologs of mammalian Zinc finger of the cerebellum and Purkinje cell protein 4. Drosophila DTB-specific cis-regulatory elements correspond to regulatory sequences of human ENGRAILED-2, PAX-2, and DACHSHUND-1 that direct MHB-specific expression in the embryonic mouse brain. We show that cis-regulatory elements and the gene networks they regulate direct the formation and function of midbrain circuits for balance and motor coordination in insects and mammals. Regulatory mechanisms mediating the genetic specification of cephalic neural circuits in arthropods correspond to those in chordates, thereby implying their origin before the divergence of deuterostomes and ecdysozoans.

神经发育和功能的相应属性表明节肢动物和脊椎动物的大脑可能具有进化上保守的组织。然而，潜在的机制仍然难以捉摸。在这里，我们确定了基因调控和字符标识网定义在deutocerebral-tritocerebral边界（DTB）果蝇。该网络包含与脊椎动物及其最近的脊索动物亲属中指导中脑-后脑边界 (MHB) 形成的基因同源的基因。遗传追踪表明，胚胎 DTB 产生了成人中脑回路，苍蝇控制听觉和前庭信息处理和运动协调，脊椎动物中的 MHB 衍生回路也是如此。DTB 特异性基因表达和功能由顺式指导-发育控制基因的调控元件，包括哺乳动物小脑锌指和浦肯野细胞蛋白 4 的同源物。果蝇DTB 特异性顺式调节元件对应于人类ENGRAILED-2、PAX-2和DACHSHUND-1 的调节序列，它们在胚胎小鼠脑中指导 MHB 特异性表达。我们证明顺式-调节元件和它们调节的基因网络指导昆虫和哺乳动物平衡和运动协调的中脑回路的形成和功能。调节节肢动物头神经回路的遗传特性的调节机制与脊索动物中的调节机制相对应，从而暗示它们在后口动物和蜕皮动物分化之前的起源。