Brachiopod Shell Proteins: Their Functions and Taxonomic Significance

The calcareous shell of the Brachiopoda is interspersed withorganic material, chiefly protein and polysaccharide. The aminoacid compositions of these proteins reflect their geneticallycoded biosynthesis and are phylogenetically and taxonomicallyinformative. The taxonomic scheme based on protein data agreeswith the scheme based on morphological and anatomical criteria.These findings indicate Crania occupies an anomolous position. Brachiopoda exhibit two main types of calcification, carbonateand phosphate. The hydroxyproline found in phosphatic inarticulateshell protein suggests an analogy with bone collagen, but theglycine content is too low to allow triple-helix formation. The number and nature of polypeptide chains making up the shellproteins have been determined by amino and carboxy end-groupanalysis as well as disc electrophoresis with SDS. In the nativestate the shell proteins are molecular aggregates and are dissolvedby 8 M urea, suggesting that the inter-chain links are largelyH–bonds. Articulate shell protein probably serves as a resilient cushioningbetween calcite fibers to protect against mechanical shock.This would be permitted by the amorphous flexible characterof the polypeptide chain. The shell proteins of the Inarticulataare different,their chitin-protein laminated shell is more sheet-likeand its structure requires less cushioning. Study of fossil protein can shed further light on shell proteinancestry and hence on brachiopod phylogeny.