Lenticles: innervated secretory structures that are expressed at every other larval moult

Lenticles are dome-shaped circles or ovals of cuticle with a dark rim. They occur with a precise segmental arrangement in the larvae and pupae of lycaenid and hesperiid butterflies. In Calpodes ethlius (Lepidoptera, Hesperiidae) each lenticle is secreted by a pair of large polyploid epidermal cells. The dark rim or annulus is formed from a ring-shaped cell. The dome, which consists of an epicuticle with a perforate intermediate layer like a pepper-pot, is formed by a central goblet cell. Between the perforate intermediate layer and the cell surfaces there is a cavity that contains material presumed to be secretion. Both cells have elaborate basal plasma membrane reticular systems and the apical microvilli associated with an extensive smooth endoplasmic reticulum that is typical of lipid secreting cells. In addition, there is a plasma membrane reticular system in the ring cell and between it and the goblet cell that contains the endings of nerves having neurosecretory vesicles. Lenticles thus have a structure appropriate for an innervated organ of lipid secretion. However, in their development, lenticles arise from bristles that are presumed to be sensory. Lenticles or their precursors are segmentally arranged in the five larval instars and the pupa, but the pattern changes at each moult. The cells that form a lenticle at one moult have a rest period at the next one when they only secrete surface cuticle. Many lenticles are paired in their cycle of development, with only one of the pair making a lenticle at a particular moult. For example, the dorsal and lateral lenticles alternate in position between anterior and posterior. The second and fourth instar segments have anterior and the third and fifth instars have posterior lenticles. In the first instar the cells that will make lenticles for the second and third instars both make bristles. Lenticles are thus formed by cells that not only change their response to ecdysone qualitatively by switching from bristle to lenticle but also alternate in their later responses, switching back and forth at alternate moults between the formation of a lenticle and the secretion of surface cuticle.