Structure of the prostomial appendages and the central nervous system in the Protodrilida (Polychaeta)

Abstract

The prostomial appendages and the central nervous system have been investigated by electron microscopy in Protodriloides chaetifer, P. symbioticus, Protodrilus haurakiensis, P. oculifer, P. ciliatus, P. helgolandicus, P. adhaerens, Saccocirrus krusadensis and S. papillocereus. The tentacles are highly developed, mobile sensory structures and consist of cuticle, epidermis, a different number of intraepithelial nerves, a small blind-ending blood vessel and a bundle of longitudinal muscle fibres. An internal canal is only present in Protodrilus and Saccocirrus species. On the tentacles seven types of sensory cells have been found including different multiciliated and uniciliated sensory cells with cilia penetrating the cuticle, sensory cells with non-penetrative cilia, phaosomes and basal ciliated sensory cells. The latter are described for the first time in polychaetes. From the specific pattern of innervation by up to five nerves originating close to the brain from the dorsal and ventral roots of the circumoesophageal connectives it is evident that the prostomial appendages represent palps. In the palps the nerve fibres form neuroneuronal, myoneuronal and epithelioneuronal synapses. The brain also gives rise to the stomatogastric nerves and various dorsal nerves. The palp canals are separated from the surrounding tissue by a prominent extracellular matrix. The wall is formed by muscle cells. The centre is usually completely filled with the cell bodies of these muscle fibres and large coelenchyme-like cells. These cells move freely in the canals and they are very likely the structural basis for the hydroskeletal function of the canals. The canals are completely separated from other body cavities and fluid is probably driven into the canals from the blood vascular system via podocytes located in a specific zone in the prostomium. In particular, the structure of the central nervous system with its nerves, the pattern of innervation of the palps and the palp canal system are compared with those of other polychaetes with special emphasis to the Spionida, the taxon presumed to include the sister group of the Protodrilida.