Nacre

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The iridescent nacre inside a nautilus shell NautilusCutawayLogarithmicSpiral.jpg
The iridescent nacre inside a nautilus shell

Nacre ( /ˈnkər/ NAY-kər also /ˈnækrə/ NAK-rə), [1] also known as mother of pearl, is an organicinorganic composite material produced by some molluscs as an inner shell layer; it is also the material of which pearls are composed. It is strong, resilient, and iridescent.

Contents

Nacre is found in some of the most ancient lineages of bivalves, gastropods, and cephalopods. However, the inner layer in the great majority of mollusc shells is porcellaneous, not nacreous, and this usually results in a non-iridescent shine, or more rarely in non-nacreous iridescence such as flame structure as is found in conch pearls.

The outer layer of cultured pearls and the inside layer of pearl oyster and freshwater pearl mussel shells are made of nacre. Other mollusc families that have a nacreous inner shell layer include marine gastropods such as the Haliotidae, the Trochidae and the Turbinidae.

Physical characteristics

Structure and appearance

Schematic of the microscopic structure of nacre layers Nacre microscopic structure.png
Schematic of the microscopic structure of nacre layers
Electron microscopy image of a fractured surface of nacre Bruchflache eines Perlmuttstucks.JPG
Electron microscopy image of a fractured surface of nacre

Nacre is composed of hexagonal platelets of aragonite (a form of calcium carbonate) 10–20  µm wide and 0.5 µm thick arranged in a continuous parallel lamina. [2] Depending on the species, the shape of the tablets differ; in Pinna , the tablets are rectangular, with symmetric sectors more or less soluble. Whatever the shape of the tablets, the smallest units they contain are irregular rounded granules. [3] These layers are separated by sheets of organic matrix (interfaces) composed of elastic biopolymers (such as chitin, lustrin and silk-like proteins). This mixture of brittle platelets and the thin layers of elastic biopolymers makes the material strong and resilient, with a Young's modulus of 70  GPa and a yield stress of roughly 70 MPa (when dry). [4] Strength and resilience are also likely to be due to adhesion by the "brickwork" arrangement of the platelets, which inhibits transverse crack propagation. This structure, spanning multiple length sizes, greatly increases its toughness, making it almost as strong as silicon. [5]

The statistical variation of the platelets has a negative effect on the mechanical performance (stiffness, strength, and energy absorption) because statistical variation precipitates localization of deformation. [6] However, the negative effects of statistical variations can be offset by interfaces with large strain at failure accompanied by strain hardening. [6] On the other hand, the fracture toughness of nacre increases with moderate statistical variations which creates tough regions where the crack gets pinned. [7] But, higher statistical variations generates very weak regions which allows the crack to propagate without much resistance causing the fracture toughness decreases. [7] Studies have shown that this weak structural defects act as dissipative topological defects coupled by an elastic distortion. [8]

Nacre appears iridescent because the thickness of the aragonite platelets is close to the wavelength of visible light. These structures interfere constructively and destructively with different wavelengths of light at different viewing angles, creating structural colours.

The crystallographic c-axis points approximately perpendicular to the shell wall, but the direction of the other axes varies between groups. Adjacent tablets have been shown to have dramatically different c-axis orientation, generally randomly oriented within ~20° of vertical. [9] [10] In bivalves and cephalopods, the b-axis points in the direction of shell growth, whereas in the monoplacophora it is the a-axis that is this way inclined. [11] The interlocking of bricks of nacre has large impact on both the deformation mechanism as well as its toughness. [12] In addition, the mineral–organic interface results in enhanced resilience and strength of the organic interlayers. [13] [14] [15]

Formation

Nacre formation is not fully understood. The initial onset assembly, as observed in Pinna nobilis , is driven by the aggregation of nanoparticles (~50–80 nm) within an organic matrix that arrange in fibre-like polycrystalline configurations. [16] The particle number increases successively and, when critical packing is reached, they merge into early-nacre platelets. Nacre growth is mediated by organics, controlling the onset, duration and form of crystal growth. [17] Individual aragonite "bricks" are believed to quickly grow to the full height of the nacreous layer, and expand until they abut adjacent bricks. [11] This produces the hexagonal close-packing characteristic of nacre. [11] Bricks may nucleate on randomly dispersed elements within the organic layer, [18] well-defined arrangements of proteins, [2] or may grow epitaxially from mineral bridges extending from the underlying tablet. [19] [20] Nacre differs from fibrous aragonite – a brittle mineral of the same form – in that the growth in the c-axis (i.e., approximately perpendicular to the shell, in nacre) is slow in nacre, and fast in fibrous aragonite. [21]

A 2021 paper in Nature Physics examined nacre from various sponges and molluscs, noting that in each case the initial layers of nacre laid down by the organism contained spiral defects. Defects that spiralled in opposite directions created distortions in the material that drew them towards each other as the layers built up until they merged and cancelled each other out. Later layers of nacre were found to be uniform and ordered in structure. [22]

Function

Fossil nautiloid shell with original iridescent nacre in fossiliferous asphaltic limestone, Oklahoma. Dated to the late Middle Pennsylvanian, which makes it by far the oldest deposit in the world with aragonitic nacreous shelly fossils. Fossil nautiloid shell with original iridescent nacre in fossiliferous asphaltic limestone.jpg
Fossil nautiloid shell with original iridescent nacre in fossiliferous asphaltic limestone, Oklahoma. Dated to the late Middle Pennsylvanian, which makes it by far the oldest deposit in the world with aragonitic nacreous shelly fossils.

Nacre is secreted by the epithelial cells of the mantle tissue of various molluscs. The nacre is continuously deposited onto the inner surface of the shell, the iridescent nacreous layer, commonly known as mother of pearl. The layers of nacre smooth the shell surface and help defend the soft tissues against parasites and damaging debris by entombing them in successive layers of nacre, forming either a blister pearl attached to the interior of the shell, or a free pearl within the mantle tissues. The process is called encystation and it continues as long as the mollusc lives.

In different mollusc groups

The form of nacre varies from group to group. In bivalves, the nacre layer is formed of single crystals in a hexagonal close packing. In gastropods, crystals are twinned, and in cephalopods, they are pseudohexagonal monocrystals, which are often twinned. [11]

Commercial sources

The main commercial sources of mother of pearl have been the pearl oyster, freshwater pearl mussels, and to a lesser extent the abalone, popular for their sturdiness and beauty in the latter half of the 19th century.

Widely used for pearl buttons especially during the 1900s, were the shells of the great green turban snail Turbo marmoratus and the large top snail, Tectus niloticus . The international trade in mother of pearl is governed by the Convention on International Trade in Endangered Species of Wild Fauna and Flora, an agreement signed by more than 170 countries. [24]

Decorative uses

Architecture

White nacre mosaic tiles in the ceiling of the Criterion Restaurant in London Mother of pearl ceiling.jpg
White nacre mosaic tiles in the ceiling of the Criterion Restaurant in London

Both black and white nacre are used for architectural purposes. The natural nacre may be artificially tinted to almost any color. Nacre tesserae may be cut into shapes and laminated to a ceramic tile or marble base. The tesserae are hand-placed and closely sandwiched together, creating an irregular mosaic or pattern (such as a weave). The laminated material is typically about 2 millimetres (0.079 in) thick. The tesserae are then lacquered and polished creating a durable and glossy surface.

Instead of using a marble or tile base, the nacre tesserae can be glued to fiberglass. The result is a lightweight material that offers a seamless installation and there is no limit to the sheet size. Nacre sheets may be used on interior floors, exterior and interior walls, countertops, doors and ceilings. Insertion into architectural elements, such as columns or furniture is easily accomplished.[ citation needed ]

Fashion

Nacre bracelet Nacre sticks.JPG
Nacre bracelet

Mother of pearl buttons are used in clothing either for functional or decorative purposes. The Pearly Kings and Queens are an elaborate example of this.

Nacre is also used to decorate watches, knives, guns and jewellery.

Musical instruments

Nacre inlay is often used for music keys and other decorative motifs on musical instruments. Many accordion and concertina bodies are completely covered in nacre, and some guitars have fingerboard or headstock inlays made of nacre (as well as some guitars having plastic inlays designed to imitate the appearance of nacre). The bouzouki and baglamas (Greek plucked string instruments of the lute family) typically feature nacre decorations, as does the related Middle Eastern oud (typically around the sound holes and on the back of the instrument). Bows of stringed instruments such as the violin and cello often have mother of pearl inlay at the frog. It is traditionally used on saxophone keytouches, as well as the valve buttons of trumpets and other brass instruments. The Middle Eastern goblet drum (darbuka) is commonly decorated by mother of pearl.

Firearms

Mother of pearl is often used in the decorative grips of firearms, and in other gun furniture.

Other

Mother of pearl is sometimes used to make spoon-like utensils for caviar (i.e. caviar servers [25] [26] ) so as to not spoil the taste with metallic spoons.

Manufactured nacre

In 2012, researchers created calcium-based nacre in the laboratory by mimicking its natural growth process. [27]

In 2014, researchers used lasers to create an analogue of nacre by engraving networks of wavy 3D "micro-cracks" in glass. When the slides were subjected to an impact, the micro-cracks absorbed and dispersed the energy, keeping the glass from shattering. Altogether, treated glass was reportedly 200 times tougher than untreated glass. [28]

See also

Related Research Articles

Pearl Hard object produced within a living shelled mollusc

A pearl is a hard, glistening object produced within the soft tissue of a living shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pearl is composed of calcium carbonate in minute crystalline form, which has deposited in concentric layers. The ideal pearl is perfectly round and smooth, but many other shapes, known as baroque pearls, can occur. The finest quality of natural pearls have been highly valued as gemstones and objects of beauty for many centuries. Because of this, pearl has become a metaphor for something rare, fine, admirable and valuable.

Exoskeleton External skeleton of an organism

An exoskeleton is the external skeleton that supports and protects an animal's body, in contrast to the internal skeleton (endoskeleton) of, for example, a human. In usage, some of the larger kinds of exoskeletons are known as "shells". Examples of animals with exoskeletons include insects such as grasshoppers and cockroaches, and crustaceans such as crabs and lobsters, as well as the shells of certain sponges and the various groups of shelled molluscs, including those of snails, clams, tusk shells, chitons and nautilus. Some animals, such as the tortoise and turtle, have both an endoskeleton and an exoskeleton.

Aragonite Calcium carbonate polymorph

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3 (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Bivalvia Class of molluscs

Bivalvia, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. Bivalves as a group have no head and they lack some usual molluscan organs like the radula and the odontophore. They include the clams, oysters, cockles, mussels, scallops, and numerous other families that live in saltwater, as well as a number of families that live in freshwater. The majority are filter feeders. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment where they are relatively safe from predation. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and file shells, can swim. The shipworms bore into wood, clay, or stone and live inside these substances.

Conchiolin

Conchiolins are complex proteins which are secreted by a mollusc's outer epithelium.

Cultured pearl Pearl created under human-controlled conditions

A cultured pearl is a pearl initiated by a mussel or oyster farmer under controlled conditions using two very different groups of bivalve mollusks - the freshwater river mussels and saltwater pearl oysters.

Biomineralization Process by which living organisms produce minerals

Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often to harden or stiffen existing tissues. Such tissues are called mineralized tissues. It is an extremely widespread phenomenon; all six taxonomic kingdoms contain members that are able to form minerals, and over 60 different minerals have been identified in organisms. Examples include silicates in algae and diatoms, carbonates in invertebrates, and calcium phosphates and carbonates in vertebrates. These minerals often form structural features such as sea shells and the bone in mammals and birds. Organisms have been producing mineralized skeletons for the past 550 million years. Ca carbonates and Ca phosphates are usually crystalline, but silica organisms (sponges, diatoms...) are always non crystalline minerals. Other examples include copper, iron and gold deposits involving bacteria. Biologically formed minerals often have special uses such as magnetic sensors in magnetotactic bacteria (Fe3O4), gravity-sensing devices (CaCO3, CaSO4, BaSO4) and iron storage and mobilization (Fe2O3•H2O in the protein ferritin).

Sea snail Common name for snails that normally live in saltwater

Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the absence of a visible shell.

Biomaterial Any substance that has been engineered to interact with biological systems for a medical purpose

A biomaterial is a substance that has been engineered to interact with biological systems for a medical purpose, either a therapeutic or a diagnostic one. As a science, biomaterials is about fifty years old. The study of biomaterials is called biomaterials science or biomaterials engineering. It has experienced steady and strong growth over its history, with many companies investing large amounts of money into the development of new products. Biomaterials science encompasses elements of medicine, biology, chemistry, tissue engineering and materials science.

<i>Neopilina</i> Genus of molluscs

Neopilina is a highly derived genus of modern monoplacophoran.

Bivalve shell

A bivalve shell is part of the body, the exoskeleton or shell, of a bivalve mollusk. In life, the shell of this class of mollusks is composed of two hinged parts or valves. Bivalves are very common in essentially all aquatic locales, including saltwater, brackish water, and freshwater. The shells of bivalves commonly wash up on beaches and along the edges of lakes, rivers, and streams. Bivalves by definition possess two shells or valves, a "right valve" and a "left valve", that are joined by a ligament. The two valves usually articulate with one another using structures known as "teeth" which are situated along the hinge line. In many bivalve shells, the two valves are symmetrical along the hinge line—when truly symmetrical, such an animal is said to be equivalved; if the valves vary from each other in size or shape, inequivalved. If symmetrical front-to-back, the valves are said to be equilateral, and are otherwise considered inequilateral.

Ammolite

Ammolite is an opal-like organic gemstone found primarily along the eastern slopes of the Rocky Mountains of North America. It is made of the fossilized shells of ammonites, which in turn are composed primarily of aragonite, the same mineral contained in nacre, with a microstructure inherited from the shell. It is one of few biogenic gemstones; others include amber and pearl.1 In 1981, ammolite was given official gemstone status by the World Jewellery Confederation (CIBJO), the same year commercial mining of ammolite began. It was designated the official gemstone of the City of Lethbridge, Alberta in 2007.

Mollusc shell Exoskeleton of an animal in the phylum Mollusca

The molluscshell 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 small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) are mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end of the Early Cambrian Period. They are very diverse, and there is no formal definition of "small shelly fauna" or "small shelly fossils". Almost all are from earlier rocks than more familiar fossils such as trilobites. Since most SSFs were preserved by being covered quickly with phosphate and this method of preservation is mainly limited to the Late Ediacaran and Early Cambrian periods, the animals that made them may actually have arisen earlier and persisted after this time span.

Mollusca Large phylum of invertebrate animals

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda. The members are known as molluscs or mollusks. Around 85,000 extant species of molluscs are recognized. The number of fossil species is estimated between 60,000 and 100,000 additional species. The proportion of undescribed species is very high. Many taxa remain poorly studied.

Mineralized tissues Biological tissues incorporating minerals

Mineralized tissues are biological tissues that incorporate minerals into soft matrices. Typically these tissues form a protective shield or structural support. Bone, mollusc shells, deep sea sponge Euplectella species, radiolarians, diatoms, antler bone, tendon, cartilage, tooth enamel and dentin are some examples of mineralized tissues.

Lustrin A is an insoluble protein used in the production of a nacreous layer in bivalve molluscs. It contributes to the properties of the nacreous layer, imparting resistance to cracking and elasticity. This is accomplished by its structure; it consists of many spring-like units which can expand when the shell is under extensional pressure. Its structure is similar to that of proteins involved in silica deposition in diatoms. It consists of 1428 amino acid residues. Its molecular weight is estimated to be 142 kDa. Its terminus consists of a protease inhibitor, which contributes to its longevity in the molluscan shell matrix.

Imitation pearl Manmade objects resembling pearls

Imitation pearls are man-made objects that are designed to resemble real pearls. Several methods are used to create imitation pearls from starting materials that include glass, plastic, and actual mollusc shell. Some beads are coated with a pearlescent substance to imitate the natural iridescence of nacre or mother of pearl. Varieties of imitation pearls include:

Molluscs in culture

Culture consists of the social behaviour and norms in human societies transmitted through social learning. Molluscs play a variety of roles in culture, including but not limited to art and literature, with both practical interactions—whether useful or harmful—and symbolic uses.

Lia Addadi Italian biochemist

Lia Addadi is a professor of structural biology at the Weizmann Institute of Science. She works on crystallisation in biology, including biomineralization, interactions with cells and crystallisation in cell membranes. She was elected a member of the National Academy of Sciences (NAS) in 2017 for “distinguished and continuing achievements in original research”, and the American Philosophical Society (2020).

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Further reading