Structure and Mechanical Adaptability of a Modern Elasmoid Fish Scale from the Common Carp

The carp (Cyprinus carpio) has typical elasmoid scales commonly found on teleosts. They provide protection while retaining flexibility and maneuverability of the fish. The exterior surface of the scale consists of an ultrathin discontinuous mineral layer on top of mineralized woven collagen fibrils. The underlying foundation is composed of two collagenous components. The major one consists of a single-twisted Bouligand'' structure with a twisting angle of 36 degrees. A secondary sheet-like'' structure, formed by thinner collagen fibrils oriented along the thickness direction, acts to increase the integrity of the scale. Here, we identify the deformation and failure mechanisms of the carp scale, revealing slight tensile anisotropy. Using in situ small-angle X-ray scattering during tensile testing, the toughening mechanisms of the scale, including the adaptive structural reorientation of lamellae as well as fibrillar sliding and elastic deformation, are quantified and compared with those of other fish scales.