Mollusc shell
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.
Formation
A mollusc shell is formed, repaired and maintained by a part of the anatomy called the mantle. Any injuries to or abnormal conditions of the mantle are usually reflected in the shape and form and even color of the shell. When the animal encounters harsh conditions that limit its food supply, or otherwise cause it to become dormant for a while, the mantle often ceases to produce the shell substance. When conditions improve again and the mantle resumes its task, a "growth line" is produced.The mantle edge secretes a shell which has two components. The organic constituent is mainly made up of polysaccharides and glycoproteins; its composition may vary widely: some molluscs employ a wide range of chitin-control genes to create their matrix, whereas others express just one, suggesting that the role of chitin in the shell framework is highly variable; it may even be absent in monoplacophora. This organic framework controls the formation of calcium carbonate crystals, and dictates when and where crystals start and stop growing, and how fast they expand; it even controls the polymorph of the crystal deposited, controlling positioning and elongation of crystals and preventing their growth where appropriate.
The shell formation requires certain biological machinery. The shell is deposited within a small compartment, the extrapallial space, which is sealed from the environment by the periostracum, a leathery outer layer around the rim of the shell, where growth occurs. This caps off the extrapallial space, which is bounded on its other surfaces by the existing shell and the mantle. The periostracum acts as a framework from which the outer layer of carbonate can be suspended, but also, in sealing the compartment, allows the accumulation of ions in concentrations sufficient for crystallization to occur. The accumulation of ions is driven by ion pumps packed within the calcifying epithelium. Calcium ions are obtained from the organism's environment through the gills, gut and epithelium, transported by the haemolymph to the calcifying epithelium, and stored as granules within or in-between cells ready to be dissolved and pumped into the extrapallial space when they are required. The organic matrix forms the scaffold that directs crystallization, and the deposition and rate of crystals is also controlled by hormones produced by the mollusc. Because the extrapallial space is supersaturated, the matrix could be thought of as impeding, rather than encouraging, carbonate deposition; although it does act as a nucleating point for the crystals and controls their shape, orientation and polymorph, it also terminates their growth once they reach the necessary size. Nucleation is endoepithelial in Neopilina and Nautilus, but exoepithelial in the bivalves and gastropods.
The formation of the shell involves a number of genes and transcription factors. On the whole, the transcription factors and signalling genes are deeply conserved, but the proteins in the secretome are highly derived and rapidly evolving. engrailed serves to demark the edge of the shell field; dpp controls the shape of the shell, and Hox1 and Hox4 have been implicated in the onset of mineralization. In gastropod embryos, Hox1 is expressed where the shell is being accreted; however no association has been observed between Hox genes and cephalopod shell formation. Perlucin increases the rate at which calcium carbonate precipitates to form a shell when in saturated seawater; this protein is from the same group of proteins as those responsible for the formation of eggshell and pancreatic stone crystals, but the role of C-type lectins in mineralization is unclear. Perlucin operates in association with Perlustrin, a smaller relative of lustrin A, a protein responsible for the elasticity of organic layers that makes nacre so resistant to cracking. Lustrin A bears remarkable structural similarity to the proteins involved in mineralization in diatoms - even though diatoms use silica, not calcite, to form their tests!
Development
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 /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.
Secondary loss
The loss of a shell in the adult form of some gastropods is achieved by the discarding of the larval shell; in other gastropods and in cephalopods, the shell is lost or demineralized by the resorption of its carbonate component by the mantle tissue.Shell proteins
Hundreds of soluble and insoluble proteins control shell formation. They are secreted into the extrapallial space by the mantle, which also secretes the glycoproteins, proteoglycans, polysaccharides and chitin that make up the organic shell matrix. Insoluble proteins tend to be thought of as playing a more important/major role in crystallization control. The organic matrix of shells tends to consist of β-chitin and silk fibroin. Perlucin encourages carbonate deposition, and is found at the interface of the chitinous and aragonitic layer in some shells. An acidic shell matrix appears to be essential to shell formation, in the cephalopods at least; the matrix in the non-mineralized squid gladius is basic.In oysters and potentially most molluscs, the nacreous layer has an organic framework of the protein MSI60, which has a structure a little like spider silk and forms sheets; the prismatic layer uses MSI31 to construct its framework. This too forms beta-pleated sheets. Since acidic amino acids, such as aspartic acid and glutamic acid, are important mediators of biomineralization, shell proteins tend to be rich in these amino acids. Aspartic acid, which can make up up to 50% of shell framework proteins, is most abundant in calcitic layers, and also heavily present in aragonitic layers. Proteins with high proportions of glutamic acid are usually associated with amorphous calcium carbonate.
The soluble component of the shell matrix acts to inhibit crystallization when in its soluble form, but when it attaches to an insoluble substrate, it permits the nucleation of crystals. By switching from a dissolved to an attached form and back again, the proteins can produce bursts of growth, producing the brick-wall structure of the shell.
It may be possible to use shell protein information in gastropod systematics, e.g. to discriminate species level diversity, but methods need further development.
Chemistry
The formation of a shell in molluscs appears to be related to the secretion of ammonia, which originates from urea. The presence of an ammonium ion raises the pH of the extrapallial fluid, favouring the deposition of calcium carbonate. This mechanism has been proposed not only for molluscs, but also for other unrelated mineralizing lineages.Structure
The calcium carbonate layers in a shell are generally of two types: an outer, chalk-like prismatic layer and an inner pearly, lamellar or nacreous layer. The layers usually incorporate a substance called conchiolin, often in order to help bind the calcium carbonate crystals together. Conchiolin is composed largely of quinone-tanned proteins.The periostracum and prismatic layer are secreted by a marginal band of cells, so that the shell grows at its outer edge. Conversely, the nacreous layer is derived from the main surface of the mantle.
Some shells contain pigments which are incorporated into the structure. This is what accounts for the striking colors and patterns that can be seen in some species of seashells, and the shells of some tropical land snails. These shell pigments sometimes include compounds such as pyrroles and porphyrins.
Shells are almost always composed of polymorphs of calcium carbonate - either calcite or aragonite. In many cases, such as the shells of many of the marine gastropods, different layers of the shell are composed of calcite and aragonite. In a few species which dwell near hydrothermal vents, iron sulfide is used to construct the shell. Phosphate is never utilised by molluscs, with the exception of Cobcrephora, whose molluscan affinity is uncertain.
Shells are composite materials of calcium carbonate and organic macromolecules. Shells can have numerous ultrastructural motifs, the most common being crossed-lamellar, prismatic, homogeneous, foliated and nacre. Although not the most common, nacre is the most studied type of layer.