Due to the advantages of the hard, calcifying shell, the Mollusca are one of
the most successful animal phyla. The shell forms during embryonic and larval
development; however, many molluscan groups have a highly reduced shell or have
lost it completely as development and maturation proceeds. These major
developmental transitions in shell morphology frequently correlate with ecological
transitions (e.g. diet change/change from planktonic to bethic existence pre- and
post-metamorphosis, respectively). While shell loss may leave an organism
vulnerable to predation, many have evolved alternative means to deter predators.
Here we compare and contrast the post-hatching larval development and shell
growth through the use of the life cycle staging of Bursatella leachii and Aplysia
californica in laboratory settings. The larval developmental sequence of B. leachii is
indistinguishable to other previously described plankotrophic aplysiids. However, the
growth rate and size of B. leachii larvae differ from A. californica larvae substantially,
growing relatively faster and larger by an average of 10 μm. We also describe the
life cycle of B. leachii in context of the development of the larval shell and its
subsequent loss in post-metamorphic stages. Comparison of the Stage 6 shells,
both whole and cross-sections, of A. californica and B. leachii through the use of
SEM showed little difference in morphology. These data indicate that we have
established a reliable culturing technique for B. leachii in the laboratory which makes
this species can be easily amendable to experimentation at all developmental stages.
Metamorphosis and shell loss/reduction in A. californica and B. leachii highlight the
differences of the developmental program of both species, which reflects its
complexity at a molecular, cellular and organismal level. The comparison of sea
hares is an ideal evolutionary comparative model system for the loss of acquired
features.
Molluscan biomineralization has been of broad scientific interest ranging from
paleontological (molluscan shells provide one of the best fossil records for a
metazoan phylum), to material science (perl and nacre formation) research.
Although the properties (i.e. evolutionary origins, construction, patterning, physical) of
the molluscan shell have been studied for decades, the underlying molecular and
cellular mechanisms of how shell formation occurs are just recently surfacing with the
identification of a handful of shell forming proteins. It is now known that one of the
main components involved in the control of shell synthesis are the proteinaceous
constituents of the shell matrix with in different kinds of functions (i.e. cell signaling,
enzymatic activities), which are contributing to the diversity of different shell types in
gastropod, bivalve and scaphopod molluscs. However, the differential gene
expression and regulation within the mantle still remains unknown. Here we relate
the developmental expression of eleven genes present in the mantle, the organ
responsible for the secretion of the shell, in the sea hare Aplysia californica
(Opisthobranchia, Anaspidea). Six genes that show very little changes in expression
levels (Cluster 1). Three genes shows increased levels of expression during
trochophore and veliger stages which then decrease in metamorphic stages (Cluster
3). Two genes had peptide-like profiles, genes that low expression during early
development but have high expression levels late in development (Cluster 4). All
eleven genes display dynamic spatial and temporal expression profiles within the
larval shell field and mantle for the construction of the larval shell. The expression
data from these eleven genes reflect the regulatory complexity that underlies the
molluscan shell construction during larval stages. While the fabrication of the shell is
taking place, the incorporation of both ancient and novel genes during also suggest
that there is a core set of mantle-secreting genes for shell construction was provided
by a shared metazoan ancestor to produce the range of molluscan shell types we
see today.