Most hard tissues have as their primary purpose to be stiff. Outside the arthropods, mineralisation of a soft organic matrix is the almost universal method of producing high stiffness. However, stiffening brings with it the undesirable mechanical result of brittleness (lack of toughness). The mineralisation of some tissues, such as bone and dentine, can be modified rather easily, in evolutionary terms, to produce the optimum mix of stiffness with bending strength (which, except at the highest mineralisations, go together) on one hand and toughness on the other hand. However, in most other tissues, such as mollusc shell, echinoderm skeleton, brachiopod shell, barnacle shell and enamel, mineralisation is almost all-or-none, and no subtle gradations seem possible. In such cases, other features, such as architecture, must be modified to produce a useful skeleton. Not only the mechanical properties of the skeletal tissue, but its cost, mass and time taken for production will, biologists tend to assume, be balanced by natural selection to produce a satisfactory result. However, such complexity makes it difficult to be sure that we understand the extent to which mineralised skeletal materials are the best possible solution to the problems facing the animals and that we are not just telling ‘Just-So’ stories. Furthermore, there are some skeletal materials that do not seem to make much sense at the moment, although no doubt all will become clear eventually.

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