Ultrastructure and functional morphology of the appendages in the reef-building sedentary polychaete Sabellaria alveolata (Annelida, Sedentaria, Sabellida)

Abstract

Background The sedentary polychaete Sabellaria alveolata, the sandcastle or honeycomb worm, possesses four different kinds of appendages besides the parapodia: opercular papillae, tentacular filaments, palps, and branchiae. It exhibits a highly specialized anterior end, the operculum, formed by the prostomium, peristomium, and two anterior segments. The operculum comprises opercular papillae, tentacular filaments, and palps. Paired branchiae are present from the second thoracic chaetiger onwards on the posteriorly following segments except for the last ones. Ultrastructural data on these appendages are either scanty, incomplete, or even lacking in Sabellariidae. In order to analyze their functional morphology, to bridge the data gap, and providing data for future phylogenetic and evolutionary analyses, we investigated the appendages of S. alveolata by applying light microscopy, confocal laser scanning microscopy, scanning, and transmission electron microscopy. Results In S. alveolata the entire body is covered by a thin cuticle characterized by the absence of layers of parallel collagen fibers with no differentiation between the various body regions including the branchiae. The opercular papillae bear numerous tufts of receptor cells and lack motile cilia. The tentacular filaments show a distinctive pattern of motile cilia. Their most conspicuous morphological feature is a cell-free cartilaginous endoskeletal structure enclosed by ECM. Besides musculature the filaments include a single coelomic cavity but blood vessels are absent. The palps are ciliated and possess two coelomic cavities and a single blind-ending internal blood vessel. Besides external ciliation and receptor cells, the coelomate branchiae are highly vascularized and equipped with numerous blood spaces extending deep between the epidermal cells resulting in low diffusion distances. Conclusions All appendages, including the branchiae, bear receptor cells and, as such, are sensory. The opercular papillae resemble typical parapodial cirri. In contrast, the tentacular filaments have a triple function: sensing, collecting and transporting particles. A similarity to branchiae can be excluded. The palps are typical grooved palps. A revised classification of polychaete branchiae is suggested; thereby, the branchiae of S. alveolata belong to the most common type comprising coelom, musculature, and blood vessels. The results indicate that diffusion distances between blood and environment have been underestimated in many cases.