Intestinal granular cells of a cartilaginous fish, thornback ray <i>Raja clavata</i>: morphological characterization and expression of different molecules

This investigation aims to fill gaps in our understanding of the intestinal immune cells of elasmobranchs. Whole digestive tracts of fifteen thornback ray Raja clavata were provided by a trawl fleet from the Gulf of Asinara (Sardinia, western Mediterranean Sea). Histochemical, immunohistochemical and ultrastructural observations were conducted on the spiral intestine. Three types of granular cells were identified; type I in epithelium, types II and III in lamina propria-submucosa, with each of them containing cytoplasmic granules with different ultrastructural characteristics. Data on size and density of each granular cell type are provided. Immunostaining of intestinal sections showed the reactivity of the granular cells: type I cells were positive for lysozyme, mast cell tryptase and tumor necrosis factor-ɑ based on antibody staining; type III cells were immune-reactive to anti-interleukin 6 antibody, whilst type II cells were negative to all the antibodies used. Comparison of each granular cell type with immune cells of teleosts or mammals and an hypothesis on their nature and function are reported. A potential role for granular cells in intestinal cellular immunity is also discussed with respect to type I and type III cells having similarities to Paneth cells and neutrophils, respectively