Abstract. The size, morphology and species-specific texture of mollusc shell biominerals is one of the unresolved questions in nature. In search of molecular control principles, chitin has been identified by Weiner and Traub (FEBS Lett. 1980, 111 : 311-316) as one of the organic compounds with a defined co-organization with mineral phases. Chitin fibers can be aligned with certain mineralogical axes of crystalline calcium carbonate in a speciesspecific manner. These original observations motivated the functional characterization of chitin forming enzymes in molluscs. The full-length cDNA cloning of mollusc chitin synthases identified unique myosin domains as part of the biological control system. The potential impact of molecular motors and other conserved domains of these complex transmembrane enzymes on the evolution of shell biomineralization is investigated and discussed in this article.
Chitin in organisms Chitin in extracellular matrices of cellsIn the end of the 19th century, there was a hot debate regarding the chemical nature of chitin. It lasted until acetylated glucosamine was found in alkaline melts of the carapace of diverse arthropods in contrast to tunicate cellulose [1], and "mycosin" of fungi in contrast to "fungal cellulose" (reviewed by [2]). Today, fungi are well accepted as one of the most prominent groups of eukaryotic organisms which contain chitin as part of their extracellular matrix [3]. Since chitin is usually associated with proteins, complex carbohydrates and sometimes mineral phases, it remains a challenge until today to characterize the diversity of cell walls and integuments in both, uni-and multicellular organisms [4][5][6][7][8][9]. One of the best examples for naturally pure chitin are cell wall appendices of diatoms [10,11], and some enzymes involved in their formation have recently been identified [12,13]. As originally determined by fiber diffraction studies [14-17], three major modifications of chitin are distinguished: a-, b-, and g-chitin [18,19], which differ from each other by the arrangement and molar fractions of differently oriented poly-N-acetylated linear b(1!4) glucosamine (i.e. chitobiose homopolymer, Fig. 1) backbones. They assemble into fiber crystals which are stabilized by H-bonds in either two or three dimensions [4,20,21]. Squids such as the European Squid Loligo vulgaris or the Humboldt squid Dosidicus gigas, which are members of the molluscan class Cephalopoda, perform the biosynthesis of different chitin polymorphs in a tissuespecific manner. The chitinous organs are associated with different sets of proteins [22]. The N-acetyl-glucosamine (GlcNAc) chains are assembled into fibrils and hierarchical structures in order to accomodate specific functions such as mechanical stiffness and strength [23,24]. This is important for the functional design, e.g. of insect exoskeletons and additional functions such as structural colours and adhesive micro-pillar appendages [25,26].A major achievement is the abundant information regarding primary structures of enzyme...