Conchiolin (sometimes referre to as conchin) and perlucin are complex proteins which are secreted by a mollusc's outer epithelium (the mantle). These proteins are part of a matrix of organic macromolecules, mainly proteins and polysaccharides, that assembled together form the microenvironment where crystals nucleate and grow. This organic matrix also holds and binds to the crystals of aragonite which give such shells their stiffness. The ions necessary to form calcium carbonate are also secreted by the mantle, but it is the tailored environment created by the organic matrix which causes aragonite (rather than calcite) crystals to nucleate, in much the same way that collagen nucleates hydroxyapatite crystals. Conchiolin serves as a relatively flexible, crack-deflecting matrix for the mineral aggregate particles; its strength and the strong bonding of perlucin give the finished material (nacre) an impressive level of toughness.