Willis J H, Rezaur R, Sehnal F
J Embryol Exp Morphol. 1982 Oct;71:25-40.
Metamorphosing insects treated with juvenoids may secrete composite cuticles which combine morphological features of two metamorphic stages within the area secreted by an individual epidermal cell. Characters found combined were pigmentation, tanning, surface sculpturing, and microtrichiae. Neighbouring cells frequently form different types of cuticle. Composite cuticles should not be confused with the more common mosaic cuticles, which are composed of discrete areas with different stage-specific morphology (e.g. some larval patches set in an otherwise normal adult cuticle). Treatment of last instar nymphs of Pyrrhocoris apterus (Hemiptera) with a juvenoid induced secretion of composite cuticle which combined larval morphological features with imaginal pigmentation. Cells remained susceptible to composite cuticle induction over an extended period prior to the actual initiation of cuticle deposition. Composite cuticle which contained larval microtrichiae and pupal tanning were induced with juvenoids in the lepidopterans Hyponomeuta malinella, Hyphantria cunea, Spodoptera littoralis, Mamestra brassicae, Pieris brassicae, Aglais urticae, but not in Galleria mellonella. Composite cuticle possessing both pupal and imaginal morphological features was produced with juvenoids in Leptinotarsa decemlineata (Coleoptera). It is suggested that this type of cuticle reveals the secretory capacities of a single epidermal cell when the cellular reprogramming from one developmental stage to the next has been stabilized at an intermediate point by juvenoids.
用保幼激素处理的变态昆虫可能会分泌复合表皮，这种复合表皮在单个表皮细胞分泌的区域内合并了两个变态阶段的形态特征。发现合并存在的特征有色素沉着、鞣化、表面纹饰和微刚毛。相邻细胞经常形成不同类型的表皮。复合表皮不应与更常见的镶嵌表皮相混淆，镶嵌表皮由具有不同阶段特异性形态的离散区域组成（例如，一些幼虫斑块镶嵌在其他部分正常的成虫表皮中）。用保幼激素处理无翅红蝽（半翅目）的末龄若虫会诱导复合表皮的分泌，这种复合表皮将幼虫形态特征与成虫色素沉着结合在一起。在实际开始表皮沉积之前的很长一段时间内，细胞仍然对复合表皮的诱导敏感。在鳞翅目的苹果巢蛾、美国白蛾、斜纹夜蛾、甘蓝夜蛾、粉蝶、荨麻蛱蝶中，用保幼激素诱导产生了含有幼虫微刚毛和蛹鞣化的复合表皮，但在大蜡螟中未诱导成功。在马铃薯甲虫（鞘翅目）中，用保幼激素产生了同时具有蛹和成虫形态特征的复合表皮。有人认为，当细胞从一个发育阶段到下一个发育阶段的重编程被保幼激素稳定在中间点时，这种类型的表皮揭示了单个表皮细胞的分泌能力。
