Biological Control of Ca-Carbonate Crystal Microstructure and Texture: Adapting Crystal Morphology, Orientation, and Arrangement to Biomaterial Function

The mineralized cover of chiton (Polyplacophora) soft tissue consists of aragonite, developed as shell-plates, girdle-scales, and girdle-spicules. This study characterizes crystallographic aspects of the girdle-spicules of the species Ischnochiton rissoi, Rhyssoplax olivacea, Acanthopleura vaillantii, and Acanthopleura spinosa. Spicule crystal arrangements and texture variations are described. Different misorientations between the spicule crystals are shown and are discussed with respect to the physical properties of the biomaterial. Characterization was performed with electron backscattered diffraction (EBSD), as well as with laser confocal and Field emission scanning electron microscopy (FE-SEM) imaging. All investigated species had porous spicules and distinct structural characteristics. Spicule crystal co-orientation strength was strongly increased for R. olivacea and I. rissoi, and it was almost random for A. vaillantii. R. olivacea, I. rissoi. A. spinosa spicule crystal texture was axial, whereas A. vaillantii spicule crystals were almost untextured. For all species investigated, spicule aragonite was twinned, as demonstrated with the strong 63°/64° peak in the misorientation angle distribution diagram, indicating a {110}-twin relationship. R. olivacea and I. rissoi spicules consisted of few twinned crystals and twin boundaries; A. vaillantii and A. spinosa spicules showed an abundance of twinned crystals and twin boundaries. We observed a difference in spicule dimension, morphology, arrangement on the girdle, and crystal organization for the investigated species, but always the generation of twinned aragonite.