page_title: Mollusc shell
section_title: Development
hierarchical_section_title: Mollusc shell / Formation / Development
caption_attribution_description: English: Mollusc Shells in Manchester Museum
context_page_description: The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes. Not all shelled molluscs live in the sea; many live on the land and in freshwater.
  The ancestral mollusc is thought to have had a shell, but this has subsequently been lost or reduced on some families, such as the squid, octopus, and some smaller groups such as the caudofoveata and solenogastres. Today, over 100,000 living species bear a shell; there is some dispute as to whether these shell-bearing molluscs form a monophyletic group or whether shell-less molluscs are interleaved into their family tree.
  Malacology, the scientific study of molluscs as living organisms, has a branch devoted to the study of shells, and this is called conchology—although these terms used to be, and to a minor extent still are, used interchangeably, even by scientists.
  Within some species of molluscs, there is often a wide degree of variation in the exact shape, pattern, ornamentation, and color of the shell.
context_section_description: The shell-secreting area is differentiated very early in embryonic development. An area of the ectoderm thickens, then invaginates to become a "shell gland". The shape of this gland is tied to the form of the adult shell; in gastropods, it is a simple pit, whereas in bivalves, it forms a groove which will eventually become the hinge line between the two shells, where they are connected by a ligament. The gland subsequently evaginates in molluscs that produce an external shell. Whilst invaginated, a periostracum - which will form a scaffold for the developing shell - is formed around the opening of the invagination, allowing the deposition of the shell when the gland is everted. A wide range of enzymes are expressed during the formation of the shell, including carbonic anhydrase, alkaline phosphatase, and DOPA-oxidase (tyrosinase)/peroxidase.
  The form of the molluscan shell is constrained by the organism's ecology. In molluscs whose ecology changes from the larval to adult form, the morphology of the shell also undergoes a pronounced modification at metamorphosis. The larval shell may have a completely different mineralogy to the adult conch, perhaps formed from amorphous calcite as opposed to an aragonite adult conch.
  In those shelled molluscs that have indeterminate growth, the shell grows steadily over the lifetime of the mollusc by the addition of calcium carbonate to the leading edge or opening. Thus the shell gradually becomes longer and wider, in an increasing spiral shape, to better accommodate the growing animal inside. The shell thickens as it grows, so that it stays proportionately strong for its size.
caption: Mollusc shells in Manchester Museum
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