Nature has evolved efficient strategies to synthesize complex mineralized structures that exhibit exceptional damage tolerance. One such example is found in the hypermineralized hammer-like dactyl clubs of the stomatopods, a group of highly aggressive marine crustaceans. The dactyl clubs from one species, Odontodactylus scyllarus, exhibit an impressive set of characteristics adapted for surviving high-velocity impacts on the heavily mineralized prey on which they feed. Consisting of a multiphase composite of oriented crystalline hydroxyapatite and amorphous calcium phosphate and carbonate, in conjunction with a highly expanded helicoidal organization of the fibrillar chitinous organic matrix, these structures display several effective lines of defense against catastrophic failure during repetitive high-energy loading events.

https://science.sciencemag.org/content/sci/336/6086/1275.full.pdf

The stomatopod dactyl club : a formidable damage-tolerant biological hammer

http://maeresearch.ucsd.edu/mckittrick/pubs/2008_Acta_Biomater_3_587_Crab.pdf

Structure and mechanical properties of crab exoskeletons.

Crustacea, the subphylum of Arthropoda which dominates the aquatic environment, is of major importance in ecology and fisheries. Here we report the genome sequence of the Pacific white shrimp Litopenaeus vannamei, covering ~1.66 Gb (scaffold N50 605.56 Kb) with 25,596 protein-coding genes and a high proportion of simple sequence repeats (>23.93%). The expansion of genes related to vision and locomotion is probably central to its benthic adaptation. Frequent molting of the shrimp may be explained by an intensified ecdysone signal pathway through gene expansion and positive selection. As an important aquaculture organism, L. vannamei has been subjected to high selection pressure during the past 30 years of breeding, and this has had a considerable impact on its genome. Decoding the L. vannamei genome not only provides an insight into the genetic underpinnings of specific biological processes, but also provides valuable information for enhancing crustacean aquaculture. The Pacific white shrimp Litopenaeus vannamei is an important aquaculture species and a promising model for crustacean biology. Here, the authors provide a reference genome assembly, and show that gene expansion is involved in the regulation of frequent molting as well as benthic adaptation of the shrimp.

Penaeid shrimp genome provides insights into benthic adaptation and frequent molting

Skeletal mineralogy in a high-CO2 world

Abstract A generalized physicochemical model of the response of marine organisms’ calcifying fluids to CO2-induced ocean acidification is proposed. The model is based upon the hypothesis that some marine calcifiers induce calcification by elevating pH, and thus ΩA, of their calcifying fluid by removing protons (H+). The model is explored through two end-member scenarios: one in which a fixed number of H+ is removed from the calcifying fluid, regardless of atmospheric pCO2, and another in which a fixed external–internal H+ ratio ([H+]E/[H+]I) is maintained. The model is able to generate the full range of calcification response patterns observed in prior ocean acidification experiments and is consistent with the assertion that organisms’ calcification response to ocean acidification is more negative for marine calcifiers that exert weaker control over their calcifying fluid pH. The model is empirically evaluated for the temperate scleractinian coral Astrangia poculata with in situ pH microelectrode measurements of the coral’s calcifying fluid under control and acidified conditions. These measurements reveal that (1) the pH of the coral’s calcifying fluid is substantially elevated relative to its external seawater under both control and acidified conditions, (2) the coral’s [H+]E/[H+]I is approximately the same under control and acidified conditions, and (3) the coral removes fewer H+ from its calcifying fluid under acidified conditions than under control conditions. Thus, the carbonate system dynamics of A. poculata’s calcifying fluid appear to be most consistent with the fixed [H+]E/[H+]I end-member scenario. Similar microelectrode experiments performed on additional taxa are required to assess the model’s general applicability.

A physicochemical framework for interpreting the biological calcification response to CO2-induced ocean acidification

The integument of decapod crustaceans consists of an outer epicuticle, an exocuticle, an endocuticle and an inner membranous layer underlain by the hypodermis. The outer three layers of the cuticle are calcified. The mineral is in the form of calcite crystals and amorphous calcium carbonate. In the epicuticle, mineral is in the form of spherulitic calcite islands surrounded by the lipid-protein matrix. In the exo- and endocuticles the calcite crystal aggregates are interspersed with chitin-protein fibers which are organized in lamellae. In some species, the organization of the mineral mirrors that of the organic fibers, but such is not the case in certain cuticular regions in the xanthid crabs. Thus, control of crystal organization is a complex phenomenon unrelated to the gross morphology of the matrix.

Since the cuticle is periodically molted to allow for growth, this necessitates a bidirectional movement of calcium into the cuticle during postmolt and out during premolt resorption of the cuticle. In two species of crabs studied to date, these movements are accomplished by active transport effected by a Ca-ATPase and Na/Ca exchange mechanism.

The epi- and exocuticular layers of the new cuticle are elaborated during premolt but do not calcify until the old cuticle is shed. This phenomenon also occurs in vitro in cuticle devoid of living tissue and implies an alteration of the nucleating sites of the cuticle in the course of the molt.

/pdf/the-structure-and-calcification-of-the-crustacean-cuticle-110ts1pea1.pdf

The Structure and Calcification of the Crustacean Cuticle

The exoskeletons of pancrustaceans, as typified by decapod crustaceans and insects, demonstrate a high degree of similarity with respect to histology, ultrastructure, function, and composition. The cuticular envelope in insects and the outer epicuticle in crustaceans both serve as the primary barrier to permeability of the exoskeleton, preventing loss of water and ions to the external medium. Prior to and following ecdysis, there is a sequence of expression and synthesis of different proteins by the cuticular epithelium for incorporation into the pre-exuvial and post-exuvial procuticle of insects and the exocuticle and endocuticle of crustaceans. Both exhibit regional differences in cuticular composition, e.g., the articular (intersegmental) membranes of insects and the arthrodial (joint) membranes of crustaceans. The primary difference between these cuticles is the ability to mineralize. Crustaceans' cuticles express a unique suite of proteins that provide for the nucleation and deposition of calcium carbonate. Orthologs of genes discussed in the present review were mined from a recently completed cuticular transcriptome of the crayfish, Cherax quadricarinatus, providing new insights into the nature of these proteins.

http://lifeserv.bgu.ac.il/wb/sagia/media/Roer-2015-Exoskeletons_across_the_Pancrustacea.pdf

Robert Roer

Papers

The Structure and Calcification of the Crustacean Cuticle

Exoskeletons across the Pancrustacea: Comparative Morphology, Physiology, Biochemistry and Genetics