Crystallography of Complex Forms: The Case of Octocoral Sclerites
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Octocorals sclerites are ideal structures to study the emergence of complex shapes
from particular arrangements of crystallites in biominerals. Sinularia polydactyla
sclerites have been studied by polarizing microscopy, scanning electron microscopy,
and electron backscattered diffraction. Small sclerites (<100 μm) are simple mesocrystalline
structures, with similarly oriented sub-micrometer crystallites arranged in elongated
fibers, with only a low degree of ordered misorientations between them. Large sclerites
(∼2 mm) are composite structures: (1) at their center, small proto-sclerites (≤20 μm)
act as nuclei around which growth proceeds. (2) The large sclerite axial frame is made
of crystallites with c axes arranged in three directions, at an inclination of about 22°
from the long sclerite axis. The axial frame displays a center of symmetry, and crystallites
are arranged in an opposite trihedral arrangement with respect to this center. (3) Inside
the large sclerite, deeply rooted tubercles develop with crystallite c axes close to perpendicular
to the long sclerite axis. The tubercles grow by a branching process and display crystallite
misorientations ordered around the three a axes of the hexagonal unit cell of calcite.
Crystallites in both the frame and the tubercles form trigonal inverse pyramids resulting
from sudden or progressive changes of crystallite orientations. A crystallographic model
emphasizes the importance of the trigonal inverse pyramid as a structural pattern.
Concerning the sclerite morphology, small crystallite sizes, ordered misorientations
and mesotwinning are important features to achieve concave shapes. The sclerite surface
morphology also results from the regulation of mesocrystalline growth and patterning
by cells or vacuoles. In this respect, the sclerite morphology is the product of internal
and external forces, among which crystallographic order and molding play important roles.
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