Sulphated glycosaminoglycans support an assortment of planarian rhabdite structures

Planaria are soft-bodied, bilateral flatworms of the phylum Platyhelminthes. They are covered in cilia and use ciliary-gliding to traverse the substratum while hunting. Their body surface is covered in a layer of viscous slime primarily derived from specialised secretory granules known as rhabdites. The slime must somehow stay associated with the surface of the animal in aqueous environments whilst also lubricating the interface of the animal and the surfaces over which the animal moves. The slime prevents damage to the animal's soft body and also contributes to adhesion to the substratum. In order to gain insight into how it might achieve these diverse functions, we performed electron microscopic examination of the slime's structure. Analysis of two freshwater flatworms from the UK Schmidtea polychroa (Schmidt, 1861) and Polycelis tenuis (Ijima, 1884) revealed a high level of organisation of the slime layer and a variety of ejected slime structures. We show that these structures are rich in sulphated glycosaminoglycans (sGAGs). Most of these (269 of 285 examined) appear to be topologically closed spheroids that we name ball-GAGs. Another class appears to burst to release flower- and star-like clusters which adhere to motile cilia. We also observe fibrous nets that are associated with entrapped bacteria. Examination of the structure of rhabdites ejected onto a porous surface suggests a mechanism by which their structure allows them to both bind to the porous surface and provide a smooth layer over which the animal could glide. Such sGAG-based structures might provide models for the design of artificial biomimetic replacements for tears, saliva, bio-compatible lubricants or drug-delivery vehicles.