Ultrastructural analysis of dissolution of shell of the bivalve Mytilus edulis by the accessory boring organ of the gastropod Urosalpinx cinerea

Abstract

The shell penetrating mechanism of the muricid gastropod Urosalpinx cinerea follyensis Baker was examined with reference to solubilizing effects of the secretion of the accessory boring organ (ABO) on the ultrastructural organic and mineral components of the shell of the bivalve Mytilus edulis Linne. The fine structure of shell etched by the secretion was contrasted with normal shell and shell solubilized artificially by acids, a chelating agent, and enzymes as an aid in interpreting the pattern etched by the secretion. A synoptic series of scanning electron micrographs of representative regions of the normal shell of M. edulis was prepared to serve as a standard for ultrastructural interpretation of the patterns of dissolution. Intercrystalline conchiolin of the mosaicostracal, prismatic, myostracal, and nacreous strata was dissolved as readily as the periostracum by the ABO secretion. In the prismatic region, maximum depth of dissolution of intercrystalline organic matrix occurred when long axes of prisms were approximately perpendicular to the surface being dissolved. Microscopic solubilization of organic matrix noticeably preceded dissolution of mineral crystals, revealing subsurface prisms and lamellae similar in size and form to the distinctively shaped normal shell units. After solubilization of intercrystalline conchiolin, further dissolution by the ABO secretion revealed a variety of what appeared like internal structures in prisms and lamellae. The form of these subunits varied from that of platelets to nodules in prisms and from laths to nodules in lamellae. These intracrystalline patterns of dissolution probably resulted from preferential etching of (a) soluble intracrystalline conchiolin membranes or other internal aggregates of nacrin, (b) heterogeneously distributed trace and minor elements, or (c) from both. Carriker and Williams (1978) hypothesized that a combination of HCl, chelating agents, and enzymes in a hypertonic mucoid secretion released by the ABO dissolve shell during hole boring. The similarity of patterns of dissolution etched by the ABO secretion and those produced artificially by HCl and EDTA in the present study support the hypothesis that these chemicals, or chemicals similar to them, are constituents of the ABO secretion. Lactic and succinic acids and a chitinase-like enzyme were also suggested by Carriker and Williams as possible agents in shell dissolution. Alteration of the shell surface by experimental application of these, and other, chemical agents was not sufficiently comparable to that etched by the ABO secretion to support the suggestion. Preferential dissolution of shell matrix by the ABO secretion is functionally advantageous to boring gastropods because it increases the surface area of mineral crystals exposed to solubilization and facilitates removal of shell units from the surface of the borehole by the radula.