The crustacean cuticle is a composite material that covers the whole animal and forms the continuous exoskeleton. Nano-fibers composed of chitin and protein molecules form most of the organic matrix of the cuticle that, at the macroscale, is organized in up to eight hierarchical levels. At least two of them, the exo- and endocuticle, contain a mineral phase of mainly Mg-calcite, amorphous calcium carbonate and phosphate. The high number of hierarchical levels and the compositional diversity provide a high degree of freedom for varying the physical, in particular mechanical, properties of the material. This makes the cuticle a versatile material ideally suited to form a variety of skeletal elements that are adapted to different functions and the eco-physiological strains of individual species. This review presents our recent analytical, experimental and theoretical studies on the cuticle, summarising at which hierarchical levels structure and composition are modified to achieve the required physical properties. We describe our multi-scale hierarchical modeling approach based on the results from these studies, aiming at systematically predicting the structure-composition-property relations of cuticle composites from the molecular level to the macro-scale. This modeling approach provides a tool to facilitate the development of optimized biomimetic materials within a knowledge-based design approach.

Functional adaptation of crustacean exoskeletal elements through structural and compositional diversity: a combined experimental and theoretical study

We employ ab initio calculations and investigate the single-crystalline elastic properties of (Ca,Mg)CO3 crystals covering the whole range of concentrations from pure calcite CaCO3 to pure magnesite MgCO3. Studying different distributions of Ca and Mg atoms within 30-atom supercells, our theoretical results show that the energetically most favorable configurations are characterized by elastic constants that nearly monotonously increase with the Mg content. Based on the first principles-derived single-crystalline elastic anisotropy, the integral elastic response of (Ca,Mg)CO3 polycrystals is determined employing a mean-field self-consistent homogenization method. As in case of single-crystalline elastic properties, the computed polycrystalline elastic parameters sensitively depend on the chemical composition and show a significant stiffening impact of Mg atoms on calcite crystals in agreement with the experimental findings. Our analysis also shows that it is not advantageous to use a higher-scale two-phase mix of stoichiometric calcite and magnesite instead of substituting Ca atoms by Mg ones on the atomic scale. Such two-phase composites are not significantly thermodynamically favorable and do not provide any strong additional stiffening effect.

Ab initio study of single-crystalline and polycrystalline elastic properties of Mg-substituted calcite crystals.

Function-related adaptations of ultrastructure, mineral phase distribution and mechanical properties in the incisive cuticle of mandibles of Porcellio scaber Latreille, 1804

Ultrastructure and mineral composition of the cornea cuticle in the compound eyes of a supralittoral and a marine isopod.

The architecture of the joint head cuticle and its transition to the arthrodial membrane in the terrestrial crustacean Porcellio scaber

Like most terrestrial isopods, Porcellio scaber stores cuticular calcium in sternal deposits before moulting. The deposits consist of spherules that are formed within a confined ecdysial space containing a fluid of known cationic composition. The spherules are composed of granular particles containing mainly amorphous calcium carbonate (ACC), little amorphous calcium phosphate, and an organic matrix. We precipitated calcium carbonate using near physiological, but phosphate-free salines and matrix proteins extracted from native deposits, and analyzed structure, mineral phase, and composition of the precipitates. Within the test solutions, the total soluble fraction leads to precipitation of ACC granules. Agglomerations of ACC granules and the dried soluble fraction of the organic matrix are virtually devoid of organic phosphates. The agglomerations mimic several aspects in the architecture of native sternal deposits and were stable for at least one month. The saline alone has no effect on the crystal phase...

Influence of Near-Physiological Salines and Organic Matrix Proteins from Amorphous CaCO3 Deposits of Porcellio scaber on in Vitro CaCO3 Precipitation

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Influence of Near-Physiological Salines and Organic Matrix Proteins from Amorphous CaCO3 Deposits of Porcellio scaber on in Vitro CaCO3 Precipitation