Ultrastructure of calcified cartilage in the endoskeletal tesserae of sharks

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Abstract
The tesserate pattern of endoskeletal calcification has been investigated in jaws, gill arches, vertebral arches and fins of the sharks Carcharhinus menisorrah, Triaenodon obesus and Negaprion brevirostris by techniques of light and electron microscopy. Individual tesserae develop peripherally at the boundary between cartilage and perichondrium. An inner zone, the body, is composed of calcified cartilage containing viable chondrocytes separated by basophilic contour lines which have been called Liesegang waves or rings. The outer zone of tesserae, the cap, is composed of calcified tissue which appears to be produced by perichondrial fibroblasts more directly, i.e., without first differentiating as chondroblasts. Furthermore, the cap zone is penetrated by acidophilic Sharpey fibers of collagen. It is suggested that scleroblasts of the cap zone could be classified as osteoblasts. If so, the cap could be considered a thin veneer of bone atop the calcified cartilage of the body of a tessera. By scanning electron microscopy it was observed that outer and inner surfaces of tesserae differ in appearance. Calcospherites and hydroxyapatite crystals similar to those commonly seen on the surface of bone are present on the outer surface of the tessera adjacent to the perichondrium. On the inner surface adjoining hyaline cartilage, however, calcospherites of variable size are the predominant surface feature. Transmission electron microscopy shows calcification in close association with coarse collagen fibrils on the outer side of a tessera, but such fibrils are absent from the cartilaginous matrix along the under side of tesserae. Calcified cartilage as a tissue type in the endoskeleton of sharks is a primitive vertebrate characteristic. Calcification in the tesserate pattern occurring in modern Chondrichthyes may be derived from an ancestral pattern of a continuous bed of calcified cartilage underlying a layer of perichondral bone, as theorized by Ørvig (1951); or the tesserate pattern in these fish may itself be primitive.