Fertilization in Callochiton castaneus (Mollusca)

A fine-structural study of fertilization in Callochiton castaneus has revealed that the mechanism of sperm penetration into the egg is intermediate between the primitive condition found in members of the order Lepidopleurida and the more derived condition found in the Chitonida. C. castaneus sperm have the long needlelike nuclear filament and reduced acrosome that characterizes all Chitonida, but they have retained several plesiomorphic features such as an unspecialized mid-piece and a lack of flagellar reinforcement. As in some Lepidopleurida but unlike any Chitonida, the egg hull in this species comprises a thick, smooth jelly coat permeated by pores that permit sperm rapid access to the vitelline layer. The jelly coat is delicate and quickly dissolves when a sperm concentrate is used, suggesting that excess acrosomal enzymes may be responsible. Once the sperm have penetrated the vitelline layer, the long nuclear filament bridges the gap to cups in the egg membrane. However, once the fertilization membrane is raised, the perivitelline space exceeds the length of the nuclear filament, preventing other sperm from penetrating the egg. A fertilization cone forms around the nuclear filament of the penetrating sperm, but it does not appear to engulf the body of the sperm. Rather, the nuclear chromatin is injected into the egg as a long thread. The remaining sperm organelles are apparently abandoned on the egg surface. If this is the case, it would be a significant departure from fertilization in other molluscs and many other metazoans, in which sperm organelles, such as centrioles and mitochondria, enter the egg. New sperm and egg characters, as well as significant differences in fertilization, indicate that Callochitonidae are basal to all other members of the order Chitonida and may warrant separation as the sister taxon to the suborders Chitonina and Acanthochitonina.