Truman James W, Riddiford Lynn M
Department of Biology, University of Washington, Seattle, WA 98195, USA.
Insect Biochem Mol Biol. 2007 Aug;37(8):761-70. doi: 10.1016/j.ibmb.2007.05.011. Epub 2007 May 29.
The maintenance of "status quo" in larvae by juvenile hormone (JH) involves both the programming of ecdysteroid-dependent synthesis during the molt and the suppression of morphogenetic growth during the intermolt. The latter morphostatic action does not require ecdysteroids, and has been studied in the formation of imaginal discs in Manduca sexta. Preultimate larval instars have both invaginated discs and imaginal primordia, both of which grow isomorphically with the larva. In the last instar, the young discs/primordia initiate the morphogenesis and patterning that results in a mature disc. JH suppresses both the initiation and progression of the signaling that transforms immature discs or primordia into a fully patterned imaginal disc. This transformation normally occurs in the context of the rapid growth of the last larval stage, and nutrient-dependent factors appear to be able to override the JH suppression. The morphostatic action of JH may have been important for the evolution of the larval stage. Studies on embryos of basal, hemimetabolous insects show that their premature exposure to JH can truncate patterning programs and cause precocious tissue maturation, factors essential for organizing a novel larval form. This suppression of embryonic patterning then results in embryonic fields that remain dormant as long as JH is present. These are the primordia that can transform into imaginal discs once JH disappears in preparation for metamorphosis.
保幼激素(JH)使幼虫维持“原状”,这既涉及蜕皮期间蜕皮甾类依赖性合成的编程,也涉及蜕皮间期形态发生生长的抑制。后一种形态静止作用不需要蜕皮甾类,并且已经在烟草天蛾的成虫盘形成过程中得到研究。倒数第二龄幼虫既有内陷盘又有成虫原基,二者均与幼虫同形生长。在最后一龄幼虫中,幼嫩的盘/原基启动形态发生和模式形成,从而形成成熟的盘。JH抑制将未成熟盘或原基转化为完全模式化成虫盘的信号传导的起始和进程。这种转化通常发生在最后一个幼虫阶段快速生长的背景下,并且营养依赖性因子似乎能够克服JH的抑制作用。JH的形态静止作用可能对幼虫阶段的进化很重要。对基础半变态昆虫胚胎的研究表明,它们过早暴露于JH会截断模式形成程序并导致早熟的组织成熟,这些都是构建新幼虫形态所必需的因素。这种对胚胎模式形成的抑制随后导致只要JH存在就保持休眠的胚胎区域。这些原基在JH消失以准备变态时可转化为成虫盘。