The effect of tessellation on stiffness in the hyoid arch of elasmobranchs

Tessellated cartilage forms much of the skeleton of sharks and rays, in contrast to most other aquatic vertebrates who possess a skeleton of bone. Interestingly, many species of sharks and rays also regularly generate exceptionally high forces in the execution of day-to-day activities, such as when feeding on bony fish, mammals, and hard-shelled invertebrates. Tessellated cartilage differs from other types of cartilage in that they are covered by an outer layer of small mineralized tiles (tesserae) that are connected by fibrous connective tissue. Tesserae, therefore, are hypothesized to play a role in stiffening the cartilaginous skeleton for food capture and other activities that require the generation of high forces. In this study, the hyomandibula and ceratohyal cartilages, which support the jaw and throat regions of sharks and rays, were tested under compressive load in a material testing system to determine the contribution of tesserae to stiffness. Previous hypotheses suggest an abrupt upward shift in the slope of the stress-strain curve in tessellated materials due to collision of tesserae. Young's Modulus (E) was calculated and used to evaluate cartilage stiffness in a range of elasmobranch species. Our results revealed that there was an abrupt shift in Young's Modulus for elements loaded in compression. We postulate that this shift, characterized by an inflection point in the stress-strain curve, is the result of the tesserae approaching one another and compressing the intervening fibrous tissue, supporting the hypothesis that tesserae function to stiffen these cartilages under compressive loading regimes. Using published data for nontessellated cartilage for comparison, we show that this shift is, as expected, unique to tessellated cartilage.