Biomineralization, also written biomineralisation, is the process by which living organisms produce

minerals,[a] often resulting in hardened or stiffened 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.[2][3][4] 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. Calcium
carbonates and calcium 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).

In terms of taxonomic distribution, the most common biominerals are the phosphate and carbonate
salts of calcium that are used in conjunction with organic polymers such as collagen and chitin to give
structural support to bones and shells.[5] The structures of these biocomposite materials are highly
controlled from the nanometer to the macroscopic level, resulting in complex architectures that
provide multifunctional properties. Because this range of control over mineral growth is desirable for
materials engineering applications, there is interest in understanding and elucidating the
mechanisms of biologically-controlled biomineralization.[6][7]

Types

Mineralization can be subdivided into different categories depending on the following: the organisms
or processes that create chemical conditions necessary for mineral formation, the origin of the
substrate at the site of mineral precipitation, and the degree of control that the substrate has on
crystal morphology, composition, and growth.[8] These subcategories include biomineralization,
organomineralization, and inorganic mineralization, which can be subdivided further. However, the
usage of these terms varies widely in the scientific literature because there are no standardized
definitions. The following definitions are based largely on a paper written by Dupraz et al. (2009),[8]
which provided a framework for differentiating these terms.

Biomineralization

Biomineralization, biologically controlled mineralization, occurs when crystal morphology, growth,

composition, and location are completely controlled by the cellular processes of a specific organism.
Examples include the shells of invertebrates, such as molluscs and brachiopods. Additionally, the
mineralization of collagen provides crucial compressive strength for the bones, cartilage, and teeth of
vertebrates.[9]

Organomineralization
This type of mineralization includes both biologically induced mineralization and biologically
influenced mineralization.

Living Stromatolites in Exumas, Bahamas.

Fossil skeletal parts from extinct belemnite cephalopods of the Jurassic – these contain mineralized
calcite and aragonite.

Biologically induced mineralization occurs when the metabolic activity of microbes (e.g. bacteria)
produces chemical conditions favorable for mineral formation. The substrate for mineral growth is
the organic matrix, secreted by the microbial community, and affects crystal morphology and
composition. Examples of this type of mineralization include calcareous or siliceous stromatolites and
other microbial mats. A more specific type of biologically induced mineralization, remote calcification
or remote mineralization, takes place when calcifying microbes occupy a shell-secreting organism
and alter the chemical environment surrounding the area of shell formation. The result is mineral
formation not strongly controlled by the cellular processes of the animal host (i.e., remote
mineralization); this may lead to unusual crystal morphologies.[10]

Biologically influenced mineralization takes place when chemical conditions surrounding the site of
mineral formation are influenced by abiotic processes (e.g., evaporation or degassing). However, the
organic matrix (secreted by microorganisms) is responsible for crystal morphology and composition.
Examples include micro- to nanometer-scale crystals of various morphologies.[11][12]

Biological mineralization can also take place as a result of fossilization. See also calcification.

Biological roles

Among animals, biominerals composed of calcium carbonate, calcium phosphate, or silica perform a
variety of roles such as support, defense, and feeding.[13]

Many protists, like this coccolithophore, have protective mineralised shells

Many protists, like this coccolithophore, have protective mineralised shells

Forams from a beach

Forams from a beach

Many invertebrate animals have external exoskeletons or shells, which achieve rigidity by a variety of
mineralisations

Many invertebrate animals have external exoskeletons or shells, which achieve rigidity by a variety of
mineralisations
Vertebrate animals have internal endoskeletons which achieve rigidity by binding calcium phosphate
into hydroxylapatite

Vertebrate animals have internal endoskeletons which achieve rigidity by binding calcium phosphate
into hydroxylapatite

If present on a supracellular scale, biominerals are usually deposited by a dedicated organ, which is
often defined very early in embryological development. This organ will contain an organic matrix that
facilitates and directs the deposition of crystals.[13] The matrix may be collagen, as in
deuterostomes,[13] or based on chitin or other polysaccharides, as in molluscs.[14]

In molluscs

Further information: Mollusc shell

A variety of mollusc shells

The mollusc shell is a biogenic composite material that has been the subject of much interest in
materials science because of its unusual properties and its model character for biomineralization.
Molluscan shells consist of 95–99% calcium carbonate by weight, while an organic component makes
up the remaining 1–5%. The resulting composite has a fracture toughness ≈3000 times greater than
that of the crystals themselves.[15] In the biomineralization of the mollusc shell, specialized proteins
are responsible for directing crystal nucleation, phase, morphology, and growths dynamics and
ultimately give the shell its remarkable mechanical strength. The application of biomimetic principles
elucidated from mollusc shell assembly and structure may help in fabricating new composite
materials with enhanced optical, electronic, or structural properties.[citation needed]

The most described arrangement in mollusc shells is the nacre, known in large shells such as Pinna or
the pearl oyster (Pinctada). Not only does the structure of the layers differ, but so do their
mineralogy and chemical composition. Both contain organic components (proteins, sugars, and
lipids), and the organic components are characteristic of the layer and of the species.[4] The
structures and arrangements of mollusc shells are diverse, but they share some features: the main
part of the shell is crystalline calcium carbonate (aragonite, calcite), though some amorphous
calcium carbonate occurs as well; and although they react as crystals, they never show angles and
facets.[16]

In fungi

Global involvement of fungi in some biogeochemical cycles [17]

(a) Fungi contribute substantially to mineral weathering, leading to the release of bioavailable metals
or nutrients, which eventually may be uptaken by living organisms or precipitated as secondary
minerals
(b) Fungi as heterotrophs, recycle organic matter. While doing so, they produce metabolites such as
organic acids that can also precipitate as secondary minerals (salts). Recycling organic matter
eventually releases constitutive elements such as C, N, P, and S

(c) CO2 produced by heterotrophic fungal respiration can dissolve into H2O and depending on the
physicochemical conditions precipitate as CaCO3 leading to the formation of a secondary mineral.

Fungi are a diverse group of organisms that belong to the eukaryotic domain. Studies of their
significant roles in geological processes, "geomycology", have shown that fungi are involved with
biomineralization, biodegradation, and metal-fungal interactions.[18]

In studying fungi's roles in biomineralization, it has been found that fungi deposit minerals with the
help of an organic matrix, such as a protein, that provides a nucleation site for the growth of
biominerals.[19] Fungal growth may produce a copper-containing mineral precipitate, such as copper
carbonate produced from a mixture of (NH4)2CO3 and CuCl2.[19] The production of the copper
carbonate is produced in the presence of proteins made and secreted by the fungi.[19] These fungal
proteins that are found extracellularly aid in the size and morphology of the carbonate minerals
precipitated by the fungi.[19]

In addition to precipitating carbonate minerals, fungi can also precipitate uranium-containing

phosphate biominerals in the presence of organic phosphorus that acts as a substrate for the
process.[20] The fungi produce a hyphal matrix, also known as mycelium, that localizes and
accumulates the uranium minerals that have been precipitated.[20] Although uranium is often
deemed as toxic to living organisms, certain fungi such as Aspergillus niger and Paecilomyces
javanicus can tolerate it.[20]

Though minerals can be produced by fungi, they can also be degraded, mainly by oxalic acid–
producing strains of fungi.[21] Oxalic acid production is increased in the presence of glucose for
three organic acid producing fungi: Aspergillus niger, Serpula himantioides, and Trametes versicolor.
[21] These fungi have been found to corrode apatite and galena minerals.[21] Degradation of
minerals by fungi is carried out through a process known as neogenesis.[22] The order of most to
least oxalic acid secreted by the fungi studied are Aspergillus niger, followed by Serpula himantioides,
and finally Trametes versicolor.[21]

In bacteria

It is less clear what purpose biominerals serve in bacteria. One hypothesis is that cells create them to
avoid entombment by their own metabolic byproducts. Iron oxide particles may also enhance their
metabolism.[23]