Formation of the Apical Flaps in Nematocysts of Sea Anemones (Cnidaria: Actiniaria)

Using scanning and transmission electron mi- croscopy, we studied formation of the structure at the apical end of sea anemone nematocysts through which the tubule everts at discharge. In anemones of the genus Metridium, we found that each of the three solid triangular apical flaps comprises two layers that are continuous with those of the capsule wall: the electron-lucent inner layer is bound to the electron-dense outer layer. The two-layer structure is obvi- ous in some discharged capsules in which, perhaps due to fixation, the layers part at the flap's periphery. Before the nematocyst discharges, a channel leads from a pore at the tip of the joined flaps into the lumen of the inverted tubule. The thin laminate layer that coats each flap lines the channel. The base of the nematocyst tubule adheres to the capsule wall near the capsule's apical end, and a branch of the tubule underlies part of the laminate layer that coats the flaps. Thus the tubule is not continuous with the capsule wall but structurally separate from it. This helps reconcile differences in understanding of the number of layers con- stituting the capsule wall, and makes clear that the tubule should be considered part of the capsule contents.