Growth of protein-doped rhombohedra in the calcitic shell of craniid brachiopods

Abstract

The secondary shell of the living inarticulated brachiopod Neocrania consists of calcitic laminae interleaved with organic sheets, predominantly a 444 kDa protein with high levels of aspartic acid-asparagine and glutamic acid-glutamine. Laminae consist of tabular (01.4) rhombohedra that are composed of spherular or rhombohedral granules, ca. 30 nm in size. Rhombohedra increase by planar or spiral growth as granular, monolayered plates that commonly act as foundations for multilayered tablets up to 300 nm or so high. Rough (Ok.1) faces, kinked by cleavage, usually develop at either end of the long diagonals of rhombohedra, the edges of which may support ramparts that can accrete centripetally to enclose any organic material situated at tablet centres. Induced degradation by proteinase K shows that sectors subtended by (Ok.1) steps are doped by a fibrous polymer, identified as an exclusively intralaminar glycosylated 60 kDa protein. A 44 kDa protein has also been extracted from laminae and is presumably incorporated into tablets by centripetally growing ramparts.