Structural diversity of crustacean exoskeletons and its implications for biomimetics

The crustacean cuticle is a biological composite material consisting of chitin–protein fibres in a mineralized matrix. Recent research has revealed a surprising range of fibre architectures and mineral compositions of crustacean skeletal structures adapted to various mechanical demands. It is becoming increasingly clear that the organic fibres in the cuticle may be organized in patterns differing from the standard twisted plywood model. Observed fibre architectures in protruding skeletal structures include longitudinal and circular parallel fibre arrays. Skeletal minerals often include calcium phosphates in addition to calcium carbonates. Furthermore, skeletal properties are affected by protein cross-linking, which replaces mineralization as a stiffening mechanism in some structures. Several common structural motifs, such as the stiffening of the outer skeletal layers, the incorporation of non-mineralized cuticle in exposed structures, and interchanging layers of parallel fibres and the twisted plywood structure, can be identified in skeletal elements with similar functions. These evolutionary solutions have the potential for biomimetic applications, particularly as manufacturing technologies advance. To make use of this potential, we need to understand the processes behind the formation of the crustacean exoskeleton and determine which features are truly adaptive and worth mimicking.