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.

A novel metallic silvery color caused by pointillistic mixing of disordered nano-to micro-pixels of iridescent colors.

Rich iridescent structural colors in nature, such as peacock feathers, butterfly wings, beetle scales, and mollusc nacre, have attracted extensive attention for a long time and they generally result from the interaction between light and periodic structures. However, non-iridescent structural colors, such as silvery structural colors, have received relatively little attention, and they usually result from non-periodic structures. Here, using optical microscopy, fiber-optic spectrometry, field emission scanning electron microscopy (FE-SEM), X-ray diffraction (XRD), Fourier-transform infrared spectroscopy (FTIR), and laser Raman spectroscopy, we investigate the origin of a novel structural color occurring at the edge of a bivalve shell (i.e., an otter shell). We find that: (1) the structural colors are observed to be uniform metallic silvery when viewed with the naked eye; (2) they are surprisingly multicolored with various colorful pixels juxtaposed together when viewed with an optical microscope; (3) each individual pixel shows a single color originating from a periodic, multilayered organic film with definite spacing (d); and (4) different pixels vary significantly in size, shape, and color with different d values (202-387 nm). Finally, we confirm that the macroscopic silvery color results from the pointillistic mixing of nano-to microscale iridescent pixels. We also discuss the special photonic structure responsible for the silvery color. We hope that this work can not only accelerate our comprehension of photonic materials, but also provide new inspiration for the synthesis of silvery white materials.