Morphogenesis of silkmoth chorion: sequential modification of an early helicoidal framework through expansion and densification.

The silkmoth chorion is a helicoidally layered, fibrous structure which is constructed in four sequential morphogenetic modes, beginning with the assembly of a thin, low density lamellar framework. Subsequently, the framework expands in height by the insertion of additional fiber sheets into the preexisting lamellae. This expansion mode begins farthest from the follicular secretory cells and progresses in reverse. Individual fibers then grow in thickness, presumably through accretion of newly synthesized proteins, and eventually fuse. This third mode, which also begins in the most distant lamellae and proceeds in reverse, is called densification, as it results in an approximately two fold increase in overall chorion density without further lamellar expansion. Finally, lamellogenesis is recapitulated in miniature in a region of the chorion's surface, where very-late-forming lamellae are molded into prominent surface structures, the aeropyle crowns. The densification and especially the expansion modes suggest considerable fluidity in the developing chorion, consistent with its proposed cholesteric liquid crystalline structure. Such a structure is also consistent with numerous deviations from the ideal helicoidal array. These distortions and defects are described and discussed in terms of their possible origin and function.