Merkel, Casjen; Griesshaber, Erika; Kelm, Klemens; Neuser, Rolf D.; Jordan, Guntram; Logan, A.; Mader, Werner; Schmahl, Wolfgang W.
Micromechanical properties and structural characterization of modern inarticulated brachiopod shells.
In: Journal of Geophysical Research G: Biogeosciences, Vol. 112, Nr. G2
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 We investigated micromechanical properties and ultrastructure ofthe shells of the modern brachiopod species Lingula anatina, Disciniscalaevis, and Discradisca stella with scanning electron microscopy (SEM,EDX), transmission electron microscopy (TEM) and Vickers microhardnessindentation analyses. The shells are composed of two distinct layers, anouter primary layer and an inner secondary layer. Except for the primarylayer in Lingula anatina, which is composed entirely of organic matter,all other shell layers are laminated organic/inorganic composites. Theorganic matter is built of chitin fibers, which provide the matrix forthe incorporation of calcium phosphate. Amorphous calcium phosphate inthe outer, primary layer and crystalline apatite is deposited into theinner, secondary layer of the shell. Apatite crystallite sizes in theumbonal region of the shell are about 50 x 50 nm, while within thevalves crystallite sizes are significantly smaller, averanging 10 x 25nm. There is great variation in hardness values between shell layers andbetween the investigated brachiopod species. The microhardness of theinvestigated shells is significantly lower than that of inorganichydroxyapatite. This is caused by the predominantly organic materialcomponent that in these shells is either developed as purely organiclayers or as an organic fibrous matrix reinforced by crystallites. Ourresults show that this particular fiber composite material is veryefficient for the protection and the support of the soft animal tissue.It lowers the probability of crack formation and effectively impedescrack propagation perpendicular to the shell by crack-deviationmechanisms. The high degree of mechanical stability and toughness isachieved by two design features. First, there is the fiber compositematerial which overcomes some detrimental and enhances some advantageousproperties of the single constituents, that is the softness andflexibility of chitin and the hardness and brittleness of apatite.Second, there is a hierarchical structuring from the nanometer to amicrometer level. We could identify at least seven levels of hierarchywithin the shells.