Structure and function of the periostracum in the bivalve Perna viridis

The periostracum is an outermost coating of all shelled-molluscs such as bivalves, corresponding to the interfacial layer separating the calcareous shells from the environment. It therefore plays a key role in the growth and survival of bivalves. Nevertheless, the periostracum has attracted little attention. Here, using the optical microscope (OM) and field-emission scanning electron microscope (FE-SEM), we investigate the structure and variation of the periostracum in the green mussel Perna viridis. We find that this periostracum has a novel sandwich structure with an outer (ODL) and inner dense layer (IDL) interleaved with a middle fibrous layer (MFL). The latter consists of locally parallel fibers (57-112 nm wide) and exhibits rich iridescent colors with a reversible hydrochromic behavior. Moreover, we find that this periostracum shows a significant variation in individual shells. Impressively, its thickness varies continuously along the shell edge. In addition, the periostracum at the ventral region is not only the thickest in a shell but also reinforced with the inorganic phosphate. We assume that this unusual variation in a same shell probably originates form defensive adaptations to predation and abrasion. Although many questions remain unanswered, this work reveals a new structure model of the periostracum, which not only advances our understanding of the periostracal formation mechanism, but also provides a natural prototype for design and synthesis of biomimetic coating and photonic materials.