Regeneration and calcification in the Spirobranchus lamarcki operculum: development and comparative genetics of a novel appendage
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Regeneration, the replacement of lost or damaged body parts, and biomineralisation, the biologically controlled formation of minerals, are important and widespread abilities in the animal kingdom. Both phenomena have a complex evolutionary history; thus their study benefits from investigations in diverse animals. Spirobranchus (formerly Pomatoceros) lamarcki is a small tube-dwelling polychaete worm of the serpulid family. Serpulids have evolved a novel head appendage, the operculum, which functions as a defensive tube plug and regenerates readily when lost. In S. lamarcki, the end of the operculum is reinforced by a calcareous plate; thus, the operculum is a good system in which to study both regeneration and biomineralisation. This thesis explores several aspects of these important processes in the adult operculum. First, a time course of normal regeneration is established. Next, cell proliferation patterns are described, suggesting a combination of proliferation-dependent and proliferation-independent elements in opercular regeneration. The formation of the calcareous opercular plate is examined using both microscopic observations of whole opercular plates and X-ray diffraction analysis of isolated plate mineral, revealing a large shift in mineralogy over the course of regeneration. Histochemical study of alkaline phosphatase enzyme activity indicates the importance of these enzymes in the operculum, although their precise functions are as yet unclear. Finally, a preliminary survey of three opercular transcriptomic datasets is presented, with a broad sampling of gene families with regeneration- or biomineralisation-related roles in other animals. The opercular transcriptome constitutes the first biomineralisation transcriptome from any annelid, and one of the first transcriptomic datasets related to annelid regeneration. Many of the candidate genes examined here display interesting behaviour and suggest targets for further investigation. The work presented here establishes the S. lamarcki operculum as a promising model system in the field of evolutionary developmental biology.
Thesis, PhD Doctor of Philosophy