Microstructure and crystallography of the wall plates of the giant barnacle Austromegabalanus psittacus: a material organized by crystal growth.

In biomineralization, it is essential to know the microstructural and crystallographic organization of natural hard tissues. This knowledge is virtually absent in the case of barnacles. Here, we have examined the crystal morphology and orientation of the wall plates of the giant barnacle Austromegabalanus psittacus by means of optical and electron microscopy, and electron backscatter diffraction. The wall plates are made of calcite grains, which change in morphology from irregular to rhombohedral, except for the radii and alae, where fibrous calcite is produced. Both the grains and fibres arrange into bundles made of crystallographically co-oriented units, which grow onto each other epitaxially. We call these areas crystallographically coherent regions (CCRs). Each CCR elongates and disposes its c-axis perpendicularly or at a high angle to the growth surfaces, whereas the a-axes of adjacent CCRs differ in orientation. In the absence of obvious organic matrices, this pattern of organization is interpreted to be produced by purely crystallographic processes. In particular, due to crystal competition, CCRs orient their fastest growth axes perpendicular to the growth surface. Since each CCR is an aggregate of grains, the fastest growth axis is that along which crystals stack up more rapidly, that is, the crystallographic c-axis in granular calcite. In summary, the material forming the wall plates of the studied barnacles is under very little biological control and the main role of the mantle cells is to provide the construction materials to the growth front.