Biomineralization

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.{{cite journal |title=Processes of carbonate precipitation in modern microbial mats |journal=Earth-Science Reviews |date=1 October 2009 |pages=141–162 |volume=96 |series=Microbial Mats in Earth's Fossil Record of Life: Geobiology |issue=3 |vauthors=Dupraz C, Reid RP, Braissant O, Decho AW, Norman RS, Visscher PT |bibcode=2009ESRv...96..141D |doi=10.1016/j.earscirev.2008.10.005}} 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), which provided a framework for differentiating these terms.

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.{{cite journal |vauthors=Sherman VR, Yang W, Meyers MA |title=The materials science of collagen |journal=Journal of the Mechanical Behavior of Biomedical Materials |volume=52 |pages=22–50 |date=December 2015 |pmid=26144973 |doi=10.1016/j.jmbbm.2015.05.023 |doi-access=free}}

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

• 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.{{cite journal |title=The oyster enigma variations: a hypothesis of microbial calcification |journal=Paleobiology |date=27 September 2013 |issn=0094-8373 |pages=1–13 |volume=40 |issue=1 |vauthors=Vermeij GJ |doi=10.1666/13002 |s2cid=67846463 |url=https://escholarship.org/uc/item/8kn6h5dg}}
• 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.{{cite journal |last1=Bindschedler |first1=Saskia |last2=Cailleau |first2=Guillaume |last3=Verrecchia |first3=Eric |title=Role of Fungi in the Biomineralization of Calcite |journal=Minerals |publisher=MDPI AG |volume=6 |issue=2 |date=2016-05-05 |issn=2075-163X |page=41 |bibcode=2016Mine....6...41B |doi=10.3390/min6020041 |doi-access=free}}{{cite journal |last1=Görgen |first1=Sigrid |last2=Benzerara |first2=Karim |last3=Skouri-Panet |first3=Fériel |last4=Gugger |first4=Muriel |last5=Chauvat |first5=Franck |last6=Cassier-Chauvat |first6=Corinne |title=The diversity of molecular mechanisms of carbonate biomineralization by bacteria |journal=Discover Materials |publisher=Springer Science and Business Media LLC |volume=1 |issue=1 |date=2020-11-30 |issn=2730-7727 |doi=10.1007/s43939-020-00001-9 |doi-access=free |s2cid=230631843}}

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

Among animals, biominerals composed of calcium carbonate, calcium phosphate, or silica perform a variety of roles such as support, defense, and feeding.{{cite journal |vauthors=Livingston BT, Killian CE, Wilt F, Cameron A, Landrum MJ, Ermolaeva O, Sapojnikov V, Maglott DR, Buchanan AM, Ettensohn CA |display-authors=6 |author-link8=Donna R. Maglott |title=A genome-wide analysis of biomineralization-related proteins in the sea urchin Strongylocentrotus purpuratus |journal=Developmental Biology |volume=300 |issue=1 |pages=335–348 |date=December 2006 |pmid=16987510 |doi=10.1016/j.ydbio.2006.07.047 |doi-access=free}}

File:Braarudosphaera bigelowii.jpg|Many protists, like this coccolithophore, have protective mineralised shells.

File:Foraminifères de Ngapali.jpg|Forams from a beach

File:Lobster NSRW rotated.jpg|Many invertebrate animals have external exoskeletons or shells, which achieve rigidity by a variety of mineralisations.

File:Elephant skeleton.jpg|Vertebrate animals have internal endoskeletons which achieve rigidity by binding calcium phosphate into hydroxylapatite.

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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. The matrix may be collagen, as in deuterostomes, or based on chitin or other polysaccharides, as in molluscs.{{cite journal |vauthors=Checa AG, Ramírez-Rico J, González-Segura A, Sánchez-Navas A |title=Nacre and false nacre (foliated aragonite) in extant monoplacophorans (=Tryblidiida: Mollusca) |journal=Die Naturwissenschaften |volume=96 |issue=1 |pages=111–122 |date=January 2009 |bibcode=2009NW.....96..111C |pmid=18843476 |doi=10.1007/s00114-008-0461-1 |s2cid=10214928}}

{{Further|Mollusc shell}}

File:Shells-IZE-006-Black.jpg

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.{{cite journal |vauthors=Currey JD |title=The design of mineralised hard tissues for their mechanical functions |journal=The Journal of Experimental Biology |volume=202 |issue=Pt 23 |pages=3285–3294 |date=December 1999 |pmid=10562511 |doi=10.1242/jeb.202.23.3285|bibcode=1999JExpB.202.3285C }} 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|date=May 2023}}

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. 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.{{cite book |title=Les étapes de la découverte des rapports entre la terre et la vie: une introduction à la paléontologie |vauthors=Cuif JP, Dauphin Y |publisher=Éditions scientifiques GB |year=2003 |isbn=978-2847030082 |location=Paris |oclc=77036366}}

File:Global involvement of fungi in some biogeochemical cycles.png Modified material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].}}(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.{{cite journal |vauthors=Gadd GM |title=Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation |journal=Mycological Research |volume=111 |issue=Pt 1 |pages=3–49 |date=January 2007 |pmid=17307120 |doi=10.1016/j.mycres.2006.12.001}}

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.{{cite journal |vauthors=Li Q, Gadd GM |author2-link=Geoffrey Michael Gadd |title=Biosynthesis of copper carbonate nanoparticles by ureolytic fungi |journal=Applied Microbiology and Biotechnology |volume=101 |issue=19 |pages=7397–7407 |date=October 2017 |pmid=28799032 |doi=10.1007/s00253-017-8451-x |pmc=5594056}} Fungal growth may produce a copper-containing mineral precipitate, such as copper carbonate produced from a mixture of (NH₄)₂CO₃ and CuCl₂. The production of the copper carbonate is produced in the presence of proteins made and secreted by the fungi. These fungal proteins that are found extracellularly aid in the size and morphology of the carbonate minerals precipitated by the fungi.

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.{{cite journal |vauthors=Liang X, Hillier S, Pendlowski H, Gray N, Ceci A, Gadd GM |title=Uranium phosphate biomineralization by fungi |journal=Environmental Microbiology |volume=17 |issue=6 |pages=2064–2075 |date=June 2015 |bibcode=2015EnvMi..17.2064L |pmid=25580878 |doi=10.1111/1462-2920.12771 |s2cid=9699895}} The fungi produce a hyphal matrix, also known as mycelium, that localizes and accumulates the uranium minerals that have been precipitated. Although uranium is often deemed toxic to living organisms, certain fungi such as Aspergillus niger and Paecilomyces javanicus can tolerate it.

Though minerals can be produced by fungi, they can also be degraded, mainly by oxalic acid–producing strains of fungi. Oxalic acid production is increased in the presence of glucose for three organic acid producing fungi: Aspergillus niger, Serpula himantioides, and Trametes versicolor. These fungi have been found to corrode apatite and galena minerals.{{cite journal |vauthors=Adeyemi AO, Gadd GM |title=Fungal degradation of calcium-, lead- and silicon-bearing minerals |journal=Biometals |volume=18 |issue=3 |pages=269–281 |date=June 2005 |pmid=15984571 |doi=10.1007/s10534-005-1539-2 |s2cid=35004304}} Degradation of minerals by fungi is carried out through a process known as neogenesis.{{cite journal |vauthors=Adamo P, Violante P |title=Weathering of rocks and neogenesis of minerals associated with lichen activity |date=1 May 2000 |journal=Applied Clay Science |volume=16 |issue=5 |pages=229–256 |bibcode=2000ApCS...16..229A |doi=10.1016/S0169-1317(99)00056-3}} The order of most to least oxalic acid secreted by the fungi studied are Aspergillus niger, followed by Serpula himantioides, and finally Trametes versicolor.

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.{{cite journal |vauthors=Fortin D |title=Geochemistry. What biogenic minerals tell us |journal=Science |volume=303 |issue=5664 |pages=1618–1619 |date=March 2004 |pmid=15016984 |doi=10.1126/science.1095177 |s2cid=41179538}}

File:White cliffs of dover 09 2004.jpg of the White Cliffs of Dover is almost entirely formed from fossil skeleton remains (coccoliths), biomineralized by planktonic microorganisms (coccolithophores).}}]]

Biomineralization plays significant global roles terraforming the planet, as well as in biogeochemical cycles and as a carbon sink.{{cite book |vauthors=Gwenzi W |title=Sustainable Agriculture Reviews 37 |chapter=Carbon Sequestration via Biomineralization: Processes, Applications and Future Directions |publisher=Springer International Publishing |publication-place=Cham |year=2019 |volume=37 |pages=93–106 |isbn=978-3-030-29297-3 |issn=2210-4410 |doi=10.1007/978-3-030-29298-0_5 |s2cid=214154330}}

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Most biominerals can be grouped by chemical composition into one of three distinct mineral classes: silicates, carbonates, or phosphates.

File:Collection Penard MHNG Specimen 533-2-1 Pamphagus granulatus.tif which has covered itself with protective diatom frustules]]

File:Odontodactylus scyllarus 2.png smash their prey by swinging club-like raptorial claws made of hydroxyapatite.{{cite journal |vauthors=Patek SN, Caldwell RL |title=Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp Odontodactylus scyllarus |date=October 2005 |journal=The Journal of Experimental Biology |volume=208 |issue=Pt 19 |pages=3655–3664 |pmid=16169943 |doi=10.1242/jeb.01831 |doi-access=free |bibcode=2005JExpB.208.3655P |s2cid=312009}}]]

Silicates (glass) are common in marine biominerals, where diatoms form frustules and radiolaria form capsules from hydrated amorphous silica (opal).{{cite book |chapter=Marine Silica Cycle |title=Encyclopedia of Ocean Sciences |year=2001 |vauthors=Demaster DJ |pages=1659–1667 |isbn=9780122274305 |doi=10.1006/rwos.2001.0278}}

The major carbonate in biominerals is CaCO₃. The most common polymorphs in biomineralization are calcite (e.g. foraminifera, coccolithophores) and aragonite (e.g. corals), although metastable vaterite and amorphous calcium carbonate can also be important, either structurally{{cite journal |vauthors=Pokroy B, Kabalah-Amitai L, Polishchuk I, DeVol RT, Blonsky AZ, Sun CY, Marcus MA, Scholl A, Gilbert PU |title=Narrowly Distributed Crystal Orientation in Biomineral Vaterite |date=13 October 2015 |journal=Chemistry of Materials |language=en |volume=27 |issue=19 |pages=6516–6523 |issn=0897-4756 |arxiv=1609.05449 |doi=10.1021/acs.chemmater.5b01542 |s2cid=118355403}}{{cite journal |vauthors=Neues F, Hild S, Epple M, Marti O, Ziegler A |title=Amorphous and crystalline calcium carbonate distribution in the tergite cuticle of moulting Porcellio scaber (Isopoda, Crustacea) |journal=Journal of Structural Biology |volume=175 |issue=1 |pages=10–20 |date=July 2011 |pmid=21458575 |doi=10.1016/j.jsb.2011.03.019 |url=https://bib-pubdb1.desy.de/record/140148/files/Amorphous.pdf}} or as intermediate phases in biomineralization.{{cite journal |vauthors=Jacob DE, Wirth R, Agbaje OB, Branson O, Eggins SM |title=Planktic foraminifera form their shells via metastable carbonate phases |journal=Nature Communications |volume=8 |issue=1 |pages=1265 |date=November 2017 |bibcode=2017NatCo...8.1265J |pmid=29097678 |doi=10.1038/s41467-017-00955-0 |pmc=5668319}}{{cite journal |vauthors=Mass T, Giuffre AJ, Sun CY, Stifler CA, Frazier MJ, Neder M, Tamura N, Stan CV, Marcus MA, Gilbert PU |display-authors=6 |title=Amorphous calcium carbonate particles form coral skeletons |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=114 |issue=37 |pages=E7670–E7678 |date=September 2017 |bibcode=2017PNAS..114E7670M |pmid=28847944 |doi=10.1073/pnas.1707890114 |doi-access=free |pmc=5604026}} Some biominerals include a mixture of these phases in distinct, organised structural components (e.g. bivalve shells). Carbonates are particularly prevalent in marine environments, but also present in freshwater and terrestrial organisms.{{cite journal |vauthors=Raven JA, Giordano M |title=Biomineralization by photosynthetic organisms: evidence of coevolution of the organisms and their environment? |journal=Geobiology |volume=7 |issue=2 |pages=140–154 |date=March 2009 |bibcode=2009Gbio....7..140R |pmid=19207569 |doi=10.1111/j.1472-4669.2008.00181.x |s2cid=42962176}}

The most common biogenic phosphate is hydroxyapatite (HA), a calcium phosphate (Ca₁₀(PO₄)₆(OH)₂) and a naturally occurring form of apatite. It is a primary constituent of bone, teeth, and fish scales.{{cite journal |vauthors=Onozato H, Watabe N |title=Studies on fish scale formation and resorption. III. Fine structure and calcification of the fibrillary plates of the scales in Carassius auratus (Cypriniformes: Cyprinidae) |journal=Cell and Tissue Research |volume=201 |issue=3 |pages=409–422 |date=October 1979 |pmid=574424 |doi=10.1007/BF00236999 |s2cid=2222515}} Bone is made primarily of HA crystals interspersed in a collagen matrix—65 to 70% of the mass of bone is HA. Similarly, HA is 70 to 80% of the mass of dentin and enamel in teeth. In enamel, the matrix for HA is formed by amelogenins and enamelins instead of collagen.{{cite book |vauthors=Habibah TU, Amlani DB, Brizuela M |chapter=Biomaterials, Hydroxyapatite |title=Stat Pearls |date=January 2018 |pmid=30020686 |chapter-url=https://www.ncbi.nlm.nih.gov/books/NBK513314/ |access-date=12 August 2018 |url-status=live |archive-url=https://web.archive.org/web/20200328145125/https://www.ncbi.nlm.nih.gov/books/NBK513314/ |archive-date=28 March 2020}} Remineralisation of tooth enamel involves the reintroduction of mineral ions into demineralised enamel.{{cite journal |vauthors=Abou Neel EA, Aljabo A, Strange A, Ibrahim S, Coathup M, Young AM, Bozec L, Mudera V |display-authors=6 |title=Demineralization-remineralization dynamics in teeth and bone |journal=International Journal of Nanomedicine |volume=11 |pages=4743–4763 |year=2016 |pmid=27695330 |doi=10.2147/IJN.S107624 |doi-access=free |pmc=5034904}} Hydroxyapatite is the main mineral component of enamel in teeth.{{cite journal |vauthors=Pepla E, Besharat LK, Palaia G, Tenore G, Migliau G |title=Nano-hydroxyapatite and its applications in preventive, restorative and regenerative dentistry: a review of literature |journal=Annali di Stomatologia |volume=5 |issue=3 |pages=108–114 |date=July 2014 |pmid=25506416 |pmc=4252862}} During demineralisation, calcium and phosphorus ions are drawn out from the hydroxyapatite. The mineral ions introduced during remineralisation restore the structure of the hydroxyapatite crystals.

The clubbing appendages of the peacock mantis shrimp are made of an extremely dense form of the mineral which has a higher specific strength; this has led to its investigation for potential synthesis and engineering use.{{cite journal |vauthors=Weaver JC, Milliron GW, Miserez A, Evans-Lutterodt K, Herrera S, Gallana I, Mershon WJ, Swanson B, Zavattieri P, DiMasi E, Kisailus D |display-authors=6 |title=The stomatopod dactyl club: a formidable damage-tolerant biological hammer |journal=Science |volume=336 |issue=6086 |pages=1275–1280 |date=June 2012 |bibcode=2012Sci...336.1275W |pmid=22679090 |doi=10.1126/science.1218764 |s2cid=8509385 |url=https://www.researchgate.net/publication/225280684 |access-date=2 December 2017 |url-status=live |archive-url=https://web.archive.org/web/20200913051518/https://www.researchgate.net/publication/225280684_The_Stomatopod_Dactyl_Club_A_Formidable_Damage-Tolerant_Biological_Hammer |archive-date=13 September 2020}} Their dactyl appendages have excellent impact resistance due to the impact region being composed of mainly crystalline hydroxyapatite, which offers significant hardness. A periodic layer underneath the impact layer composed of hydroxyapatite with lower calcium and phosphorus content (thus resulting in a much lower modulus) inhibits crack growth by forcing new cracks to change directions. This periodic layer also reduces the energy transferred across both layers due to the large difference in modulus, even reflecting some of the incident energy.{{cite journal |vauthors=Tanner KE |title=Materials science. Small but extremely tough |journal=Science |volume=336 |issue=6086 |pages=1237–1238 |date=June 2012 |bibcode=2012Sci...336.1237T |pmid=22679085 |doi=10.1126/science.1222642 |s2cid=206541609}}

File:Glomerula piloseta tube microstructure.jpg), longitudinal section of the tube showing aragonitic spherulitic prismatic structure]]

Beyond these main three categories, there are a number of less-common types of biominerals, usually resulting from a need for specific physical properties or the organism inhabiting an unusual environment. For example, teeth that are primarily used for scraping hard substrates may be reinforced with particularly tough minerals, such as the iron minerals magnetite in chitons{{cite book |vauthors=Joester D, Brooker LR |chapter=The Chiton Radula: A Model System for Versatile Use of Iron Oxides* |title=Iron Oxides |date=5 July 2016 |pages=177–206 |veditors=Faivre D |edition=1st |publisher=Wiley |language=en |isbn=978-3-527-33882-5 |doi=10.1002/9783527691395.ch8}} or goethite in limpets.{{cite journal |vauthors=Barber AH, Lu D, Pugno NM |title=Extreme strength observed in limpet teeth |journal=Journal of the Royal Society, Interface |volume=12 |issue=105 |pages=20141326 |date=April 2015 |pmid=25694539 |doi=10.1098/rsif.2014.1326 |pmc=4387522}} Gastropod molluscs living close to hydrothermal vents reinforce their carbonate shells with the iron-sulfur minerals pyrite and greigite.{{cite journal |vauthors=Chen C, Linse K, Copley JT, Rogers AD |title=The 'scaly-foot gastropod': a new genus and species of hydrothermal vent-endemic gastropod (Neomphalina: Peltospiridae) from the Indian Ocean |date=August 2015 |journal=Journal of Molluscan Studies |language=en |volume=81 |issue=3 |pages=322–334 |issn=0260-1230 |doi=10.1093/mollus/eyv013 |doi-access=free}} Magnetotactic bacteria also employ magnetic iron minerals magnetite and greigite to produce magnetosomes to aid orientation and distribution in the sediments.

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File:Chitonidae - Chiton squamosus.JPG|Chitons have aragonite shells and aragonite-based eyes,{{cite web |title=Weird Sea Mollusk Sports Hundreds of Eyes Made of Armor |date=19 November 2015 |website=Live Science |url=https://www.livescience.com/52857-mollusk-has-eyes-made-of-armor.html |access-date=28 July 2016 |url-status=live |archive-url=https://web.archive.org/web/20160817142003/http://www.livescience.com/52857-mollusk-has-eyes-made-of-armor.html |archive-date=17 August 2016}} as well as teeth coated with magnetite.

File:Common limpets1.jpg|Limpets have carbonate shells and teeth reinforced with goethite.

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File:Acantharia confocal micrograph 2.png|Acantharian radiolarians have celestine crystal shells.

File:Celestine - Sakoany deposit, Katsepy, Mitsinjo, Boeny, Madagascar.jpg |Celestine crystals, the heaviest mineral in the oceans

Celestine, the heaviest mineral in the ocean, consists of strontium sulfate, SrSO₄. The mineral is named for the delicate blue colour of its crystals. Planktic acantharean radiolarians form celestine crystal shells. The denseness of the celestite ensures their shells function as mineral ballast, resulting in fast sedimentation to bathypelagic depths. High settling fluxes of acantharian cysts have been observed at times in the Iceland Basin and the Southern Ocean, as much as half of the total gravitational organic carbon flux.{{cite journal |vauthors=Martin P, Allen JT, Cooper MJ, Johns DG, Lampitt RS, Sanders R, Teagle DA |title=Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies |year=2010 |journal=Limnology and Oceanography |volume=55 |issue=2 |pages=604–614 |doi=10.4319/lo.2009.55.2.0604 |doi-access=free}}{{cite journal |title=Acantharian cysts: High flux occurrence in the bathypelagic zone of the Scotia Sea, Southern Ocean |year=2018 |vauthors=Belcher A, Manno C, Thorpe S, Tarling G |journal=Marine Biology |volume=165 |issue=7 |page=117 |bibcode=2018MarBi.165..117B |doi=10.1007/s00227-018-3376-1 |s2cid=90349921 |url=https://nora.nerc.ac.uk/id/eprint/520421/2/Supplementary%20material.pdf}}{{cite journal |title=Pathways of Organic Carbon Downward Transport by the Oceanic Biological Carbon Pump |year=2019 |vauthors=Le Moigne FA |journal=Frontiers in Marine Science |volume=6 |page=634 |doi=10.3389/fmars.2019.00634 |doi-access=free|bibcode=2019FrMaS...6..634L }} 50px Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].

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| caption1 = The siliceous diatom frustule has the highest strength of any known biological material.

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| caption2 = Sponge spicules, like this from a siliceous glass sponge, form structures many times more flexible than equivalent structures made of pure silica.

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| caption3 = Transparent glass test or shell of a radiolarian

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In nature, there is a wide array of biominerals, ranging from iron oxide to strontium sulfate, with calcareous biominerals being particularly notable.{{cite journal |title=Biomineralization and Evolutionary History |year=2003 |vauthors=Knoll AH |journal=Reviews in Mineralogy and Geochemistry |volume=54 |issue=1 |pages=329–356 |bibcode=2003RvMG...54..329K |doi=10.2113/0540329}}{{cite book |chapter=Protistan Skeletons: A Geologic History of Evolution and Constraint |title=Evolution of Lightweight Structures |series=Biologically-Inspired Systems |year=2015 |vauthors=Knoll AH, Kotrc B |volume=6 |pages=1–16 |publisher=Springer |location=Dordrecht |isbn=978-94-017-9397-1 |doi=10.1007/978-94-017-9398-8_1 |s2cid=83376982 |url=https://dash.harvard.edu/handle/1/30403697}} However, the most taxonomically widespread biomineral is silica (SiO₂·nH₂O), being present in all eukaryotic supergroups.{{cite journal |vauthors=Marron AO, Ratcliffe S, Wheeler GL, Goldstein RE, King N, Not F, de Vargas C, Richter DJ |display-authors=6 |title=The Evolution of Silicon Transport in Eukaryotes |journal=Molecular Biology and Evolution |volume=33 |issue=12 |pages=3226–3248 |date=December 2016 |pmid=27729397 |doi=10.1093/molbev/msw209 |pmc=5100055}} Notwithstanding, the degree of silicification can vary even between closely related taxa, from being found in composite structures with other biominerals (e.g., limpet teeth;{{cite journal |vauthors=Sone ED, Weiner S, Addadi L |title=Biomineralization of limpet teeth: a cryo-TEM study of the organic matrix and the onset of mineral deposition |journal=Journal of Structural Biology |volume=158 |issue=3 |pages=428–444 |date=June 2007 |pmid=17306563 |doi=10.1016/j.jsb.2007.01.001}} to forming minor structures (e.g., ciliate granules;{{cite journal |vauthors=Foissner W, Weissenbacher B, Krautgartner WD, Lütz-Meindl U |title=A cover of glass: first report of biomineralized silicon in a ciliate, Maryna umbrellata (Ciliophora: Colpodea) |journal=The Journal of Eukaryotic Microbiology |volume=56 |issue=6 |pages=519–530 |year=2009 |pmid=19883440 |doi=10.1111/j.1550-7408.2009.00431.x |pmc=2917745}} or being a major structural constituent of the organism.{{cite journal |title=Siliceous structures and silicification in flagellated protists |year=1994 |vauthors=Preisig HR |journal=Protoplasma |volume=181 |issue=1–4 |pages=29–42 |doi=10.1007/BF01666387 |bibcode=1994Prpls.181...29P |s2cid=27698051}} The most extreme degree of silicification is evident in the diatoms, where almost all species have an obligate requirement for silicon to complete cell wall formation and cell division.{{cite journal |vauthors=Darley WM, Volcani BE |title=Role of silicon in diatom metabolism. A silicon requirement for deoxyribonucleic acid synthesis in the diatom Cylindrotheca fusiformis Reimann and Lewin |journal=Experimental Cell Research |volume=58 |issue=2 |pages=334–342 |date=December 1969 |pmid=5404077 |doi=10.1016/0014-4827(69)90514-X}}{{cite journal |vauthors=Martin-Jézéquel V, Hildebrand M, Brzezinski MA |title=Silicon Metabolism in Diatoms: Implications for Growth |year=2000 |journal=Journal of Phycology |volume=36 |issue=5 |pages=821–840 |bibcode=2000JPcgy..36..821M |doi=10.1046/j.1529-8817.2000.00019.x |s2cid=84525482}} Biogeochemically and ecologically, diatoms are the most important silicifiers in modern marine ecosystems, with radiolarians (polycystine and phaeodarian rhizarians), silicoflagellates (dictyochophyte and chrysophyte stramenopiles), and sponges with prominent roles as well. In contrast, the major silicifiers in terrestrial ecosystems are the land plants (embryophytes), with other silicifying groups (e.g., testate amoebae) having a minor role.

Broadly, biomineralized structures evolve and diversify when the energetic cost of biomineral production is less than the expense of producing an equivalent organic structure.{{cite book |title=Biomineralization: Principles and Concepts in Bioinorganic Materials Chemistry |isbn=9780198508823 |vauthors=Mann S |year=2001 |publisher=Oxford University Press |url=https://books.google.com/books?id=YLNl5Jm7XxwC&q=%22Biomineralization.+Principles+and+Concepts+in+Bioinorganic+Materials+Chemistry%22}}{{cite journal |title=The evolution of silicification in diatoms: Inescapable sinking and sinking as escape? |year=2004 |vauthors=Raven JA, Waite AM |journal=New Phytologist |volume=162 |issue=1 |pages=45–61 |doi=10.1111/j.1469-8137.2004.01022.x |doi-access=free|bibcode=2004NewPh.162...45R }}{{cite journal |title=Silica Use Through Time: Macroevolutionary Change in the Morphology of the Diatom Fustule |year=2010 |vauthors=Finkel ZV, Kotrc B |journal=Geomicrobiology Journal |volume=27 |issue=6–7 |pages=596–608 |bibcode=2010GmbJ...27..596F |doi=10.1080/01490451003702941 |s2cid=85218013}} The energetic costs of forming a silica structure from silicic acid are much less than forming the same volume from an organic structure (≈20-fold less than lignin or 10-fold less than polysaccharides like cellulose).{{cite journal |title=The Transport and Function of Silicon in Plants |year=1983 |vauthors=Raven JA |journal=Biological Reviews |volume=58 |issue=2 |pages=179–207 |doi=10.1111/j.1469-185X.1983.tb00385.x |s2cid=86067386}} Based on a structural model of biogenic silica,{{cite journal |title=The amino acid and sugar composition of diatom cell-walls |year=1973 |vauthors=Hecky RE, Mopper K, Kilham P, Degens ET |journal=Marine Biology |volume=19 |issue=4 |pages=323–331 |bibcode=1973MarBi..19..323H |doi=10.1007/BF00348902 |s2cid=84200496}} Lobel et al. (1996) identified by biochemical modeling a low-energy reaction pathway for nucleation and growth of silica.{{cite journal |title=Computational model for protein-mediated biomineralization of the diatom frustule |year=1996 |vauthors=Lobel KD, West JK, Hench LL |journal=Marine Biology |volume=126 |issue=3 |pages=353–360 |bibcode=1996MarBi.126..353L |doi=10.1007/BF00354617 |s2cid=84969529}} The combination of organic and inorganic components within biomineralized structures often results in enhanced properties compared to exclusively organic or inorganic materials. With respect to biogenic silica, this can result in the production of much stronger structures, such as siliceous diatom frustules having the highest strength per unit density of any known biological material,{{cite journal |vauthors=Hamm CE, Merkel R, Springer O, Jurkojc P, Maier C, Prechtel K, Smetacek V |title=Architecture and material properties of diatom shells provide effective mechanical protection |journal=Nature |volume=421 |issue=6925 |pages=841–843 |date=February 2003 |bibcode=2003Natur.421..841H |pmid=12594512 |doi=10.1038/nature01416 |s2cid=4336989 |url=https://epic.awi.de/id/eprint/5688/1/Ham2002b.pdf}}{{cite journal |vauthors=Aitken ZH, Luo S, Reynolds SN, Thaulow C, Greer JR |title=Microstructure provides insights into evolutionary design and resilience of Coscinodiscus sp. frustule |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=113 |issue=8 |pages=2017–2022 |date=February 2016 |bibcode=2016PNAS..113.2017A |pmid=26858446 |doi=10.1073/pnas.1519790113 |doi-access=free |pmc=4776537}} or sponge spicules being many times more flexible than an equivalent structure made of pure silica.{{cite journal |title=Nanostructural Organization of Naturally Occurring Composites—Part I: Silica-Collagen-Based Biocomposites |year=2008 |vauthors=Ehrlich H, Janussen D, Simon P, Bazhenov VV, Shapkin NP, Erler C, Mertig M, Born R, Heinemann S, Hanke T, Worch H |display-authors=6 |journal=Journal of Nanomaterials |volume=2008 |pages=1–8 |doi=10.1155/2008/623838 |doi-access=free}}{{cite journal |vauthors=Shimizu K, Amano T, Bari MR, Weaver JC, Arima J, Mori N |title=Glassin, a histidine-rich protein from the siliceous skeletal system of the marine sponge Euplectella, directs silica polycondensation |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=112 |issue=37 |pages=11449–11454 |date=September 2015 |bibcode=2015PNAS..11211449S |pmid=26261346 |doi=10.1073/pnas.1506968112 |doi-access=free |pmc=4577155}} As a result, biogenic silica structures are used for support,{{cite journal |vauthors=Weaver JC, Aizenberg J, Fantner GE, Kisailus D, Woesz A, Allen P, Fields K, Porter MJ, Zok FW, Hansma PK, Fratzl P, Morse DE |display-authors=6 |title=Hierarchical assembly of the siliceous skeletal lattice of the hexactinellid sponge Euplectella aspergillum |journal=Journal of Structural Biology |volume=158 |issue=1 |pages=93–106 |date=April 2007 |pmid=17175169 |doi=10.1016/j.jsb.2006.10.027}} feeding,{{cite journal |vauthors=Nesbit KT, Roer RD |title=Silicification of the medial tooth in the blue crab Callinectes sapidus |journal=Journal of Morphology |volume=277 |issue=12 |pages=1648–1660 |date=December 2016 |pmid=27650814 |doi=10.1002/jmor.20614 |s2cid=46840652}} predation defense{{cite journal |vauthors=Pondaven P, Gallinari M, Chollet S, Bucciarelli E, Sarthou G, Schultes S, Jean F |title=Grazing-induced changes in cell wall silicification in a marine diatom |journal=Protist |volume=158 |issue=1 |pages=21–28 |date=January 2007 |pmid=17081802 |doi=10.1016/j.protis.2006.09.002}}{{cite journal |title=Size and biomechanic properties of diatom frustules influence food uptake by copepods |year=2013 |vauthors=Friedrichs L, Hörnig M, Schulze L, Bertram A, Jansen S, Hamm C |journal=Marine Ecology Progress Series |volume=481 |pages=41–51 |bibcode=2013MEPS..481...41F |doi=10.3354/meps10227 |doi-access=free |url=https://www.int-res.com/abstracts/meps/v481/p41-51/}}{{cite journal |title=The ecology of herbivore-induced silicon defences in grasses |year=2016 |vauthors=Hartley S, Degabriel JL |journal=Functional Ecology |volume=30 |issue=8 |pages=1311–1322 |bibcode=2016FuEco..30.1311H |doi=10.1111/1365-2435.12706 |doi-access=free}} and environmental protection as a component of cyst walls. Biogenic silica also has useful optical properties for light transmission and modulation in organisms as diverse as plants,{{cite journal |vauthors=Schaller J, Brackhage C, Bäucker E, Dudel EG |title=UV-screening of grasses by plant silica layer? |journal=Journal of Biosciences |volume=38 |issue=2 |pages=413–416 |date=June 2013 |pmid=23660676 |doi=10.1007/s12038-013-9303-1 |s2cid=16034220}} diatoms,{{cite journal |title=Diatoms as living photonic crystals |year=2004 |vauthors=Fuhrmann T, Landwehr S, Rharbi-Kucki E, Sumper M |journal=Applied Physics B |volume=78 |issue=3–4 |pages=257–260 |bibcode=2004ApPhB..78..257F |doi=10.1007/s00340-004-1419-4 |s2cid=121002890}}{{cite journal |title=Optical properties of diatom silica frustule with special reference to blue light |year=2008 |vauthors=Yamanaka S, Yano R, Usami H, Hayashida N, Ohguchi M, Takeda H, Yoshino K |journal=Journal of Applied Physics |volume=103 |issue=7 |pages=074701–074701–5 |bibcode=2008JAP...103g4701Y |doi=10.1063/1.2903342}}{{cite journal |vauthors=Romann J, Valmalette JC, Chauton MS, Tranell G, Einarsrud MA, Vadstein O |title=Wavelength and orientation dependent capture of light by diatom frustule nanostructures |journal=Scientific Reports |volume=5 |pages=17403 |date=December 2015 |issue=1 |bibcode=2015NatSR...517403R |pmid=26627680 |doi=10.1038/srep17403 |pmc=4667171}} sponges,{{cite journal |vauthors=Sundar VC, Yablon AD, Grazul JL, Ilan M, Aizenberg J |title=Fibre-optical features of a glass sponge |journal=Nature |volume=424 |issue=6951 |pages=899–900 |date=August 2003 |bibcode=2003Natur.424..899S |pmid=12931176 |doi=10.1038/424899a |s2cid=4426508}} and molluscs.{{cite journal |vauthors=Dougherty LF, Johnsen S, Caldwell RL, Marshall NJ |title=A dynamic broadband reflector built from microscopic silica spheres in the 'disco' clam Ctenoides ales |journal=Journal of the Royal Society, Interface |volume=11 |issue=98 |pages=20140407 |date=September 2014 |pmid=24966236 |doi=10.1098/rsif.2014.0407 |pmc=4233689}} There is also evidence that silicification is used as a detoxification response in snails{{cite journal |vauthors=Desouky M, Jugdaohsingh R, McCrohan CR, White KN, Powell JJ |title=Aluminum-dependent regulation of intracellular silicon in the aquatic invertebrate Lymnaea stagnalis |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=99 |issue=6 |pages=3394–3399 |date=March 2002 |bibcode=2002PNAS...99.3394D |pmid=11891333 |doi=10.1073/pnas.062478699 |doi-access=free |pmc=122534}} and plants,{{cite journal |vauthors=Neumann D, zur Nieden U |title=Silicon and heavy metal tolerance of higher plants |journal=Phytochemistry |volume=56 |issue=7 |pages=685–692 |date=April 2001 |bibcode=2001PChem..56..685N |pmid=11314953 |doi=10.1016/S0031-9422(00)00472-6}} biosilica has even been suggested to play a role as a pH buffer for the enzymatic activity of carbonic anhydrase, aiding the acquisition of inorganic carbon for photosynthesis.{{cite journal |vauthors=Milligan AJ, Morel FM |title=A proton buffering role for silica in diatoms |journal=Science |volume=297 |issue=5588 |pages=1848–1850 |date=September 2002 |bibcode=2002Sci...297.1848M |pmid=12228711 |doi=10.1126/science.1074958 |s2cid=206507070}}

File:Diversity of biomineralization across the eukaryotes.jpg |{{center|Diversity of biomineralization across the eukaryotes{{cite journal |title=Competition between Silicifiers and Non-silicifiers in the Past and Present Ocean and Its Evolutionary Impacts |year=2018 |vauthors=Hendry KR, Marron AO, Vincent F, Conley DJ, Gehlen M, Ibarbalz FM, Quéguiner B, Bowler C |journal=Frontiers in Marine Science |volume=5 |page=22 |doi=10.3389/fmars.2018.00022 |doi-access=free |bibcode=2018FrMaS...5...22H |s2cid=12447257}} 50px Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].}} The phylogeny shown in this diagram is based on Adl et al. (2012),{{cite journal |vauthors=Adl SM, Simpson AG, Lane CE, Lukeš J, Bass D, Bowser SS, Brown MW, Burki F, Dunthorn M, Hampl V, Heiss A, Hoppenrath M, Lara E, Le Gall L, Lynn DH, McManus H, Mitchell EA, Mozley-Stanridge SE, Parfrey LW, Pawlowski J, Rueckert S, Shadwick L, Shadwick L, Schoch CL, Smirnov A, Spiegel FW |display-authors=6 |title=The revised classification of eukaryotes |journal=The Journal of Eukaryotic Microbiology |volume=59 |issue=5 |pages=429–493 |date=September 2012 |pmid=23020233 |doi=10.1111/j.1550-7408.2012.00644.x |pmc=3483872}} with major eukaryotic supergroups named in boxes. Letters next to taxon names denote the presence of biomineralization, with circled letters indicating the prominent and widespread use of that biomineral. S, silica; C, calcium carbonate; P, calcium phosphate; I, iron (magnetite/goethite); X, calcium oxalate; SO₄, sulfates (calcium/barium/strontium), ? denotes uncertainty in the report.{{cite journal |vauthors=Ensikat HJ, Geisler T, Weigend M |title=A first report of hydroxylated apatite as structural biomineral in Loasaceae - plants' teeth against herbivores |journal=Scientific Reports |volume=6 |pages=26073 |date=May 2016 |issue=1 |bibcode=2016NatSR...626073E |pmid=27194462 |doi=10.1038/srep26073 |pmc=4872142}}{{cite journal |vauthors=Gal A, Hirsch A, Siegel S, Li C, Aichmayer B, Politi Y, Fratzl P, Weiner S, Addadi L |display-authors=6 |title=Plant cystoliths: a complex functional biocomposite of four distinct silica and amorphous calcium carbonate phases |journal=Chemistry: A European Journal |volume=18 |issue=33 |pages=10262–10270 |date=August 2012 |pmid=22696477 |doi=10.1002/chem.201201111}}{{cite journal |title=Non-Skeletal Biomineralization by Eukaryotes: Matters of Moment and Gravity |year=2010 |vauthors=Raven JA, Knoll AH |journal=Geomicrobiology Journal |volume=27 |issue=6–7 |pages=572–584 |bibcode=2010GmbJ...27..572R |doi=10.1080/01490451003702990 |s2cid=37809270 |url=https://dash.harvard.edu/handle/1/4795339}}{{cite journal |vauthors=Weich RG, Lundberg P, Vogel HJ, Jensén P |title=Phosphorus-31 NMR Studies of Cell Wall-Associated Calcium-Phosphates in Ulva lactuca |journal=Plant Physiology |volume=90 |issue=1 |pages=230–236 |date=May 1989 |pmid=16666741 |doi=10.1104/pp.90.1.230 |pmc=1061703}}

There are questions which have yet to be resolved, such as why some organisms biomineralize while others do not, and why is there such a diversity of biominerals besides silicon when silicon is so abundant, comprising 28% of the Earth's crust. The answer to these questions lies in the evolutionary interplay between biomineralization and geochemistry, and in the competitive interactions that have arisen from these dynamics. Fundamentally whether an organism produces silica or not involves evolutionary trade-offs and competition between silicifiers themselves, and non-silicifying organisms (both those which use other biominerals, and non-mineralizing groups). Mathematical models and controlled experiments of resource competition in phytoplankton have demonstrated the rise to dominance of different algal species based on nutrient backgrounds in defined media. These have been part of fundamental studies in ecology.{{cite journal |jstor=1935608 |title=Resource Competition between Plankton Algae: An Experimental and Theoretical Approach |vauthors=Tilman D |journal=Ecology |year=1977 |volume=58 |issue=2 |pages=338–348 |bibcode=1977Ecol...58..338T |doi=10.2307/1935608}}{{cite journal |title=The impact of light intensity and daylength on silicate and nitrate competition among marine phytoplankton |year=1994 |vauthors=Sommer U |journal=Limnology and Oceanography |volume=39 |issue=7 |pages=1680–1688 |bibcode=1994LimOc..39.1680S |doi=10.4319/lo.1994.39.7.1680 |url=https://oceanrep.geomar.de/id/eprint/14122/1/Sommer_1994.pdf}} However, the vast diversity of organisms that thrive in a complex array of biotic and abiotic interactions in oceanic ecosystems are a challenge to such minimal models and experimental designs, whose parameterization and possible combinations, respectively, limit the interpretations that can be built on them.

{{Clear}}

File:Haeckel Calcispongiae.jpg, Kunstformen der Natur)]]

The first evidence of biomineralization dates to some {{Ma|750}},{{cite journal |vauthors=Porter S |title=The rise of predators |journal=Geology |volume=39 |issue=6 |pages=607–608 |year=2011 |bibcode=2011Geo....39..607P |doi=10.1130/focus062011.1 |doi-access=free}}{{cite journal |vauthors=Cohen PA, Schopf JW, Butterfield NJ, Kudryavtsev AB, Macdonald FA |title=Phosphate biomineralization in mid-Neoproterozoic protists |journal=Geology |year=2011 |bibcode=2011Geo....39..539C |volume=39 |issue=6 |pages=539–542 |doi=10.1130/G31833.1 |s2cid=32229787}} and sponge-grade organisms may have formed calcite skeletons {{Ma|630}}.{{cite journal |vauthors=Maloof AC, Rose CV, Beach R, Samuels BM, Calmet CC, Erwin DH, Poirier GR, Yao N, Simons FJ |display-authors=6 |title=Possible animal-body fossils in pre-Marinoan limestones from South Australia |journal=Nature Geoscience |volume=3 |issue=9 |pages=653–659 |year=2010 |bibcode=2010NatGe...3..653M |doi=10.1038/ngeo934 |s2cid=13171894}} But in most lineages, biomineralization first occurred in the Cambrian or Ordovician periods.{{cite journal |vauthors=Wood RA, Grotzinger JP, Dickson JA |title=Proterozoic modular biomineralized metazoan from the Nama Group, Namibia |journal=Science |volume=296 |issue=5577 |pages=2383–2386 |date=June 2002 |bibcode=2002Sci...296.2383W |pmid=12089440 |doi=10.1126/science.1071599 |s2cid=9515357}} Organisms used whichever form of calcium carbonate was more stable in the water column at the point in time when they became biomineralized, and stuck with that form for the remainder of their biological history{{cite journal |vauthors=Porter SM |title=Seawater chemistry and early carbonate biomineralization |journal=Science |volume=316 |issue=5829 |pages=1302 |date=June 2007 |bibcode=2007Sci...316.1302P |pmid=17540895 |doi=10.1126/science.1137284 |s2cid=27418253}} (but see{{cite journal |vauthors=Maloof AC, Porter SM, Moore JL, Dudás FÖ, Bowring SA, Higgins JA, Fike DA, Eddy MP |title=The earliest Cambrian record of animals and ocean geochemical change |journal=Geological Society of America Bulletin |volume=122 |issue=11–12 |pages=1731–1774 |year=2010 |bibcode=2010GSAB..122.1731M |doi=10.1130/B30346.1 |s2cid=6694681}} for a more detailed analysis). The stability is dependent on the Ca/Mg ratio of seawater, which is thought to be controlled primarily by the rate of sea floor spreading, although atmospheric {{CO2}} levels may also play a role.{{cite journal |title=Eve of biomineralization: Controls on skeletal mineralogy |year=2008 |vauthors=Zhuravlev AY, Wood RA |journal=Geology |volume=36 |issue=12 |pages=923 |bibcode=2008Geo....36..923Z |doi=10.1130/G25094A.1 |url=http://wzar.unizar.es/murero/activos/pdfs/2008_Zhuravlev%26Wood_Geology.pdf |access-date=28 August 2015 |url-status=live |archive-url=https://web.archive.org/web/20160304032652/http://wzar.unizar.es/murero/activos/pdfs/2008_Zhuravlev%26Wood_Geology.pdf |archive-date=4 March 2016}}

Biomineralization evolved multiple times, independently,{{cite journal |vauthors=Murdock DJ, Donoghue PC |title=Evolutionary origins of animal skeletal biomineralization |journal=Cells Tissues Organs |volume=194 |issue=2–4 |pages=98–102 |year=2011 |pmid=21625061 |doi=10.1159/000324245 |s2cid=45466684}} and most animal lineages first expressed biomineralized components in the Cambrian period.{{cite journal |vauthors=Kouchinsky A, Bengtson S, Runnegar B, Skovsted C, Steiner M, Vendrasco M |title=Chronology of early Cambrian biomineralization |journal=Geological Magazine |pages=221–251 |year=2011 |volume=149 |issue=2 |bibcode=2012GeoM..149..221K |doi=10.1017/S0016756811000720 |doi-access=free}} Many of the same processes are used in unrelated lineages, which suggests that biomineralization machinery was assembled from pre-existing "off-the-shelf" components already used for other purposes in the organism.{{cite book |author=Knoll, A.H. |year=2004 |chapter=Biomineralization and evolutionary history |veditors=Dove PM, DeYoreo JJ, Weiner S |editor-link1=Patricia M. Dove |title=Reviews in Mineralogy and Geochemistry |chapter-url=http://www.geochem.geos.vt.edu/bgep/pubs/Chapter_11_Knoll.pdf |url-status=dead |archive-url=https://web.archive.org/web/20100620222809/http://www.geochem.geos.vt.edu/bgep/pubs/Chapter_11_Knoll.pdf |archive-date=20 June 2010}} Although the biomachinery facilitating biomineralization is complex – involving signalling transmitters, inhibitors, and transcription factors – many elements of this 'toolkit' are shared between phyla as diverse as corals, molluscs, and vertebrates.{{cite journal |vauthors=Westbroek P, Marin F |title=A marriage of bone and nacre |journal=Nature |volume=392 |issue=6679 |pages=861–862 |date=April 1998 |bibcode=1998Natur.392..861W |pmid=9582064 |doi=10.1038/31798 |doi-access=free |s2cid=4348775}} The shared components tend to perform quite fundamental tasks, such as designating that cells will be used to create the minerals, whereas genes controlling more finely tuned aspects that occur later in the biomineralization process, such as the precise alignment and structure of the crystals produced, tend to be uniquely evolved in different lineages.{{cite journal |vauthors=Jackson DJ, McDougall C, Woodcroft B, Moase P, Rose RA, Kube M, Reinhardt R, Rokhsar DS, Montagnani C, Joubert C, Piquemal D, Degnan BM |display-authors=6 |title=Parallel evolution of nacre building gene sets in molluscs |journal=Molecular Biology and Evolution |volume=27 |issue=3 |pages=591–608 |date=March 2010 |pmid=19915030 |doi=10.1093/molbev/msp278 |doi-access=free}} This suggests that Precambrian organisms were employing the same elements, albeit for a different purpose – perhaps to avoid the inadvertent precipitation of calcium carbonate from the supersaturated Proterozoic oceans. Forms of mucus that are involved in inducing mineralization in most animal lineages appear to have performed such an anticalcifatory function in the ancestral state.{{cite journal |vauthors=Marin F, Smith M, Isa Y, Muyzer G, Westbroek P |title=Skeletal matrices, muci, and the origin of invertebrate calcification |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=93 |issue=4 |pages=1554–1559 |date=February 1996 |bibcode=1996PNAS...93.1554M |pmid=11607630 |doi=10.1073/pnas.93.4.1554 |doi-access=free |pmc=39979}} Further, certain proteins that would originally have been involved in maintaining calcium concentrations within cells{{cite journal |vauthors=Lowenstam HA, Margulis L |title=Evolutionary prerequisites for early Phanerozoic calcareous skeletons |journal=Bio Systems |volume=12 |issue=1–2 |pages=27–41 |year=1980 |bibcode=1980BiSys..12...27L |author-link=Lynn Margulis |pmid=6991017 |doi=10.1016/0303-2647(80)90036-2}} are homologous in all animals, and appear to have been co-opted into biomineralization after the divergence of the animal lineages.{{cite journal |vauthors=Lowenstam HA, Margulis L |title=Evolutionary prerequisites for early Phanerozoic calcareous skeletons |journal=Bio Systems |volume=12 |issue=1–2 |pages=27–41 |year=1980 |bibcode=1980BiSys..12...27L |pmid=6991017 |doi=10.1016/0303-2647(80)90036-2}} The galaxins are one probable example of a gene being co-opted from a different ancestral purpose into controlling biomineralization, in this case, being 'switched' to this purpose in the Triassic scleractinian corals; the role performed appears to be functionally identical to that of the unrelated pearlin gene in molluscs.{{cite journal |vauthors=Reyes-Bermudez A, Lin Z, Hayward DC, Miller DJ, Ball EE |title=Differential expression of three galaxin-related genes during settlement and metamorphosis in the scleractinian coral Acropora millepora |journal=BMC Evolutionary Biology |volume=9 |pages=178 |date=July 2009 |issue=1 |bibcode=2009BMCEE...9..178R |pmid=19638240 |doi=10.1186/1471-2148-9-178 |doi-access=free |pmc=2726143}} Carbonic anhydrase serves a role in mineralization broadly in the animal kingdom, including in sponges, implying an ancestral role.{{cite journal |vauthors=Jackson DJ, Macis L, Reitner J, Degnan BM, Wörheide G |title=Sponge paleogenomics reveals an ancient role for carbonic anhydrase in skeletogenesis |journal=Science |volume=316 |issue=5833 |pages=1893–1895 |date=June 2007 |bibcode=2007Sci...316.1893J |pmid=17540861 |doi=10.1126/science.1141560 |doi-access=free |s2cid=7042860}} Far from being a rare trait that evolved a few times and remained stagnant, biomineralization pathways in fact evolved many times and are still evolving rapidly today; even within a single genus, it is possible to detect great variation within a single gene family.

File:Highborme Cay-Stromatolite-04 (cropped) 2.jpgs made by bacteria. Fossilized stromatolites record some of the earliest life.]]

The homology of biomineralization pathways is underlined by a remarkable experiment whereby the nacreous layer of a molluscan shell was implanted into a human tooth, and rather than experiencing an immune response, the molluscan nacre was incorporated into the host bone matrix. This points to the exaptation of an original biomineralization pathway. The biomineralisation capacity of brachiopods and molluscs has also been demonstrated to be homologous, building on a conserved set of genes.{{cite journal |vauthors=Wernström JV, Gąsiorowski L, Hejnol A |title=Brachiopod and mollusc biomineralisation is a conserved process that was lost in the phoronid-bryozoan stem lineage |journal=EvoDevo |volume=13 |issue=1 |pages=17 |date=September 2022 |pmid=36123753 |doi=10.1186/s13227-022-00202-8 |doi-access=free |pmc=9484238}} This indicates that biomineralisation is likely ancestral to all lophotrochozoans.

The most ancient example of biomineralization, dating back 2 billion years, is the deposition of magnetite, which is observed in some bacteria, as well as the teeth of chitons and the brains of vertebrates; it is possible that this pathway, which performed a magnetosensory role in the common ancestor of all bilaterians, was duplicated and modified in the Cambrian to form the basis for calcium-based biomineralization pathways.{{cite book |vauthors=Kirschvink JL, Hagadorn JW |chapter=10 A Grand Unified theory of Biomineralization. |veditors=Bäuerlein E |title=The Biomineralisation of Nano- and Micro-Structures |publisher=Wiley-VCH |location=Weinheim, Germany |pages=139–150 |year=2000}} Iron is stored in close proximity to magnetite-coated chiton teeth, so that the teeth can be renewed as they wear. Not only is there a marked similarity between the magnetite deposition process and enamel deposition in vertebrates, but some vertebrates even have comparable iron storage facilities near their teeth.{{cite journal |vauthors=Towe KM, Lowenstam HA |title=Ultrastructure and development of iron mineralization in the radular teeth of Cryptochiton stelleri (Mollusca) |journal=Journal of Ultrastructure Research |volume=17 |issue=1 |pages=1–13 |date=January 1967 |pmid=6017357 |doi=10.1016/S0022-5320(67)80015-7}}

Most traditional approaches to the synthesis of nanoscale materials are energy inefficient, requiring stringent conditions (e.g., high temperature, pressure, or pH), and often produce toxic byproducts. Furthermore, the quantities produced are small, and the resultant material is usually irreproducible because of the difficulties in controlling agglomeration.{{cite book |vauthors=Thomas GB, Komarneni S, Parker J |title=Nanophase and Nanocomposite Materials: Symposium Held December 1–3, 1992, Boston, Massachusetts, U.S.A. (Materials Research Society Symposium Proceedings) |publisher=Materials Research Society |location=Pittsburgh, Pa |year=1993 |isbn=978-1-55899-181-1}} In contrast, materials produced by organisms have properties that usually surpass those of analogous synthetically manufactured materials with similar phase composition. Biological materials are assembled in aqueous environments under mild conditions by using macromolecules. Organic macromolecules collect and transport raw materials and assemble these substrates and into short- and long-range ordered composites with consistency and uniformity.{{cite book |title=Biomineralization: From Nature to Application |isbn=9780470986318 |vauthors=Sigel A, Sigel H, Sigel RK |date=30 April 2008 |publisher=John Wiley & Sons |url=https://books.google.com/books?id=TiwK2VQhPMkC&q=biomineralization+applications}}{{cite book |title=Biomineralization and Biomaterials: Fundamentals and Applications |isbn=9781782423560 |vauthors=Aparicio C, Ginebra MP |date=28 September 2015 |publisher=Woodhead |url=https://books.google.com/books?id=UhGpBAAAQBAJ&q=biomineralization+applications}}

The aim of biomimetics is to mimic the natural way of producing minerals such as apatites. Many man-made crystals require elevated temperatures and strong chemical solutions, whereas the organisms have long been able to lay down elaborate mineral structures at ambient temperatures. Often, the mineral phases are not pure but are made as composites that entail an organic part, often protein, which takes part in and controls the biomineralization. These composites are often not only as hard as the pure mineral but also tougher, as the micro-environment controls biomineralization.

File:SEM image of Bacillus megaterium.jpg|Bacillus megaterium

File:Bacillus subtilis.jpg|Bacillus subtilis

One biological system that might be of key importance in the future development of architecture is bacterial biofilm. The term biofilm refers to complex heterogeneous structures comprising different populations of microorganisms that attach and form a community on inert (e.g. rocks, glass, plastic) or organic (e.g. skin, cuticle, mucosa) surfaces.{{cite journal |vauthors=Kolter R, Greenberg EP |title=Microbial sciences: the superficial life of microbes |journal=Nature |volume=441 |issue=7091 |pages=300–302 |date=May 2006 |bibcode=2006Natur.441..300K |pmid=16710410 |doi=10.1038/441300a |doi-access=free |s2cid=4430171}}

The properties of the surface, such as charge, hydrophobicity, and roughness, determine initial bacterial attachment.{{cite journal |vauthors=Palmer J, Flint S, Brooks J |title=Bacterial cell attachment, the beginning of a biofilm |journal=Journal of Industrial Microbiology & Biotechnology |volume=34 |issue=9 |pages=577–588 |date=September 2007 |pmid=17619090 |doi=10.1007/s10295-007-0234-4 |doi-access=free |s2cid=978396}} A common principle of all biofilms is the production of extracellular matrix (ECM) composed of different organic substances, such as extracellular proteins, exopolysaccharides, and nucleic acids.{{cite journal |vauthors=Branda SS, Vik S, Friedman L, Kolter R |title=Biofilms: the matrix revisited |journal=Trends in Microbiology |volume=13 |issue=1 |pages=20–26 |date=January 2005 |pmid=15639628 |doi=10.1016/j.tim.2004.11.006}} While the ability to generate ECM appears to be a common feature of multicellular bacterial communities, the means by which these matrices are constructed and function are diverse.{{cite journal |vauthors=Steinberg N, Kolodkin-Gal I |title=The Matrix Reloaded: Probing the Extracellular Matrix Synchronizes Bacterial Communities |journal=Journal of Bacteriology |volume=197 |issue=13 |pages=2092–2103 |date=July 2015 |pmid=25825428 |doi=10.1128/JB.02516-14 |pmc=4455261}}{{cite journal |vauthors=Dragoš A, Kovács ÁT |title=The Peculiar Functions of the Bacterial Extracellular Matrix |journal=Trends in Microbiology |volume=25 |issue=4 |pages=257–266 |date=April 2017 |pmid=28089324 |doi=10.1016/j.tim.2016.12.010}}

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Biomineralization‐mediated scaffolding of bacterial biofilms.jpg |{{center|Model for biomineralization-mediated scaffolding
of bacterial biofilms}} A directed growth of the calcium carbonate crystals allows mechanical support of the 3D structure. The bacterial extracellular matrix (brown) promotes the crystals' growth in specific directions.{{cite journal |vauthors=Oppenheimer-Shaanan Y, Sibony-Nevo O, Bloom-Ackermann Z, Suissa R, Steinberg N, Kartvelishvily E, Brumfeld V, Kolodkin-Gal I |display-authors=6 |title=Spatio-temporal assembly of functional mineral scaffolds within microbial biofilms |journal=npj Biofilms and Microbiomes |volume=2 |pages=15031 |year=2016 |pmid=28721240 |doi=10.1038/npjbiofilms.2015.31 |pmc=5515261}} 50px Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].{{cite journal |vauthors=Dade-Robertson M, Keren-Paz A, Zhang M, Kolodkin-Gal I |title=Architects of nature: growing buildings with bacterial biofilms |journal=Microbial Biotechnology |volume=10 |issue=5 |pages=1157–1163 |date=September 2017 |pmid=28815998 |doi=10.1111/1751-7915.12833 |pmc=5609236}} 50px Material was copied from this source, which is available under a [https://creativecommons.org/licenses/by/4.0/ Creative Commons Attribution 4.0 International License].

Bacterially induced calcium carbonate precipitation can be used to produce "self-healing" concrete. Bacillus megaterium spores and suitable dried nutrients are mixed and applied to steel-reinforced concrete. When the concrete cracks, water ingress dissolves the nutrients and the bacteria germinate triggering calcium carbonate precipitation, resealing the crack and protecting the steel reinforcement from corrosion.{{cite book |vauthors=Jonkers HM |veditors=van der Zwaag S |title=Self Healing Materials: An Alternative Approach to 20 Centuries of Materials Science |publisher=Springer |date=2007 |pages=195–204 |chapter=Self healing concrete: a biological approach |isbn=9781402062506 |chapter-url=https://books.google.com/books?id=v4zDNBtACIsC&q=Self+healing+concrete%3A+a+biological+approach.&pg=PA194}} This process can also be used to manufacture new hard materials, such as bio-cement.{{cite patent |inventor=Dosier GK |assign1=Biomason Inc |gdate=2014 |title=Methods for making construction material using enzyme producing bacteria |country=US |number=8728365 |postscript=.}}

However, the full potential of bacteria-driven biomineralization is yet to be realized, as it is currently used as a passive filling rather than as a smart designable material. A future challenge is to develop ways to control the timing and the location of mineral formation, as well as the physical properties of the mineral itself, by environmental input. Bacillus subtilis has already been shown to respond to its environment, by changing the production of its ECM. It uses the polymers produced by single cells during biofilm formation as a physical cue to coordinate ECM production by the bacterial community.{{cite journal |vauthors=Rubinstein SM, Kolodkin-Gal I, McLoon A, Chai L, Kolter R, Losick R, Weitz DA |title=Osmotic pressure can regulate matrix gene expression in Bacillus subtilis |journal=Molecular Microbiology |volume=86 |issue=2 |pages=426–436 |date=October 2012 |pmid=22882172 |doi=10.1111/j.1365-2958.2012.08201.x |pmc=3828655}}{{cite journal |vauthors=Chan JM, Guttenplan SB, Kearns DB |title=Defects in the flagellar motor increase synthesis of poly-γ-glutamate in Bacillus subtilis |journal=Journal of Bacteriology |volume=196 |issue=4 |pages=740–753 |date=February 2014 |pmid=24296669 |doi=10.1128/JB.01217-13 |pmc=3911173}}

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Autunite-69257.jpg |{{center|Autunite crystal}}

Biomineralization may be used to remediate groundwater contaminated with uranium.{{cite journal |vauthors=Newsome L, Morris K, Lloyd JR |title=The biogeochemistry and bioremediation of uranium and other priority radionuclides |journal=Chemical Geology |year=2014 |volume=363 |pages=164–184 |bibcode=2014ChGeo.363..164N |doi=10.1016/j.chemgeo.2013.10.034 |doi-access=free}} The biomineralization of uranium primarily involves the precipitation of uranium phosphate minerals associated with the release of phosphate by microorganisms. Negatively charged ligands at the surface of the cells attract the positively charged uranyl ion (UO₂²⁺). If the concentrations of phosphate and UO₂²⁺ are sufficiently high, minerals such as autunite (Ca(UO₂)₂(PO₄)₂•10-12H₂O) or polycrystalline HUO₂PO₄ may form thus reducing the mobility of UO₂²⁺. Compared to the direct addition of inorganic phosphate to contaminated groundwater, biomineralization has the advantage that the ligands produced by microbes will target uranium compounds more specifically rather than react actively with all aqueous metals. Stimulating bacterial phosphatase activity to liberate phosphate under controlled conditions limits the rate of bacterial hydrolysis of organophosphate and the release of phosphate to the system, thus avoiding clogging of the injection location with metal phosphate minerals. The high concentration of ligands near the cell surface also provides nucleation foci for precipitation, which leads to higher efficiency than chemical precipitation.{{cite book |vauthors=Lloyd JR, Macaskie LE |title=Environmental microbe-metal interactions: Bioremediation of radionuclide-containing wastewaters |publisher=ASM Press |location=Washington, DC |year=2000 |pages=277–327 |isbn=978-1-55581-195-2}}

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{{See also|Mineral#Biogenic minerals}}

The geological definition of mineral normally excludes compounds that occur only in living beings. However, some minerals are often biogenic (such as calcite) or are organic compounds in the sense of chemistry (such as mellite). Moreover, living beings often synthesize inorganic minerals (such as hydroxylapatite) that also occur in rocks.{{citation needed|date=May 2023}}

The International Mineralogical Association (IMA) is the generally recognized standard body for the definition and nomenclature of mineral species. {{As of|2020|12}}, the IMA recognizes 5,650 official mineral species{{cite web |title=The New IMA List of Minerals A Work in Progress |work=The New IMA List of Minerals |publisher=IMA – CNMNC (Commission on New Minerals Nomenclature and Classification) |date=November 2020 |vauthors=Pasero M |display-authors=etal |url=http://cnmnc.main.jp/IMA_Master_List_%282020-11%29.pdf |access-date=11 December 2020 |url-status=dead |archive-url=https://web.archive.org/web/20201210045439/http://cnmnc.main.jp/IMA_Master_List_(2020-11).pdf |archive-date=10 December 2020}} out of 5,862 proposed or traditional ones.{{cite web |title=IMA Database of Mineral Properties/ RRUFF Project |publisher=Department of Geosciences, University of Arizona |url=https://rruff.info/ima/ |access-date=11 December 2020}}

The IMA's decision to exclude biogenic crystalline substances is a topic of contention among geologists and mineralogists. For example, Lowenstam (1981) stated that "organisms are capable of forming a diverse array of minerals, some of which cannot be formed inorganically in the biosphere."{{cite journal |vauthors=Lowenstam HA |title=Minerals formed by organisms |journal=Science |volume=211 |issue=4487 |pages=1126–1131 |date=March 1981 |bibcode=1981Sci...211.1126L |jstor=1685216 |pmid=7008198 |doi=10.1126/science.7008198}}

Skinner (2005) views all solids as potential minerals and includes biominerals in the mineral kingdom, which are created by organisms' metabolic activities. Skinner expanded the previous definition of a mineral to classify "element or compound, amorphous or crystalline, formed through biogeochemical processes," as a mineral.

Recent advances in high-resolution genetics and X-ray absorption spectroscopy are providing revelations on the biogeochemical relations between microorganisms and minerals that may shed new light on this question.{{cite journal |vauthors=Nickel EH |title=The definition of a mineral |journal=The Canadian Mineralogist |volume=33 |issue=3 |pages=689–90 |year=1995 |url=https://pubs.geoscienceworld.org/canmin/article-abstract/33/3/689/12679/the-definition-of-a-mineral?redirectedFrom=fulltext}}{{cite journal |vauthors=Skinner HC |title=Biominerals |journal=Mineralogical Magazine |volume=69 |issue=5 |pages=621–41 |year=2005 |bibcode=2005MinM...69..621S |doi=10.1180/0026461056950275 |s2cid=232388764}} For example, the IMA-commissioned "Working Group on Environmental Mineralogy and Geochemistry " deals with minerals in the hydrosphere, atmosphere, and biosphere.{{cite web |title=Working Group on Environmental Mineralogy and Geochemistry |publisher=International Mineralogical Association |language=en |date=3 August 2011 |website=Commissions, working groups and committees |url=https://www.ima-mineralogy.org/WGEMG_objectives.htm |access-date=4 April 2018}} The group's scope includes mineral-forming microorganisms, which exist on nearly every rock, soil, and particle surface spanning the globe to depths of at least 1,600 metres below the sea floor and 70 kilometres into the stratosphere (possibly entering the mesosphere).{{cite book |vauthors=Takai K |chapter=Limits of life and the biosphere: Lessons from the detection of microorganisms in the deep sea and deep subsurface of the Earth. |title=Origins and Evolution of Life: An Astrobiological Perspective |veditors=Gargaud M, Lopez-Garcia P, Martin H |pages=469–86 |date=2010 |publisher=Cambridge University Press |location=Cambridge |isbn=978-1-139-49459-5}}{{cite journal |vauthors=Roussel EG, Bonavita MA, Querellou J, Cragg BA, Webster G, Prieur D, Parkes RJ |title=Extending the sub-sea-floor biosphere |journal=Science |volume=320 |issue=5879 |pages=1046 |date=May 2008 |bibcode=2008Sci...320.1046R |pmid=18497290 |doi=10.1126/science.1154545 |s2cid=23374807 |url=https://archimer.ifremer.fr/doc/00000/4209/|url-access=subscription }}{{cite journal |vauthors=Pearce DA, Bridge PD, Hughes KA, Sattler B, Psenner R, Russell NJ |title=Microorganisms in the atmosphere over Antarctica |journal=FEMS Microbiology Ecology |volume=69 |issue=2 |pages=143–157 |date=August 2009 |bibcode=2009FEMME..69..143P |pmid=19527292 |doi=10.1111/j.1574-6941.2009.00706.x |doi-access=free}}

Biogeochemical cycles have contributed to the formation of minerals for billions of years. Microorganisms can precipitate metals from solution, contributing to the formation of ore deposits. They can also catalyze the dissolution of minerals.{{cite journal |vauthors=Newman DK, Banfield JF |title=Geomicrobiology: how molecular-scale interactions underpin biogeochemical systems |journal=Science |volume=296 |issue=5570 |pages=1071–1077 |date=May 2002 |bibcode=2002Sci...296.1071N |pmid=12004119 |doi=10.1126/science.1010716 |s2cid=1235688}}{{cite journal |vauthors=Warren LA, Kauffman ME |title=Geoscience. Microbial geoengineers |journal=Science |volume=299 |issue=5609 |pages=1027–1029 |date=February 2003 |jstor=3833546 |pmid=12586932 |doi=10.1126/science.1072076 |s2cid=19993145}}{{cite journal |vauthors=González-Muñoz MT, Rodriguez-Navarro C, Martínez-Ruiz F, Arias JM, Merroun ML, Rodriguez-Gallego M |title=Bacterial biomineralization: new insights from Myxococcus-induced mineral precipitation |journal=Geological Society, London, Special Publications |volume=336 |issue=1 |pages=31–50 |bibcode=2010GSLSP.336...31G |year=2010 |doi=10.1144/SP336.3 |s2cid=130343033}}

Before the International Mineralogical Association's listing, over 60 biominerals had been discovered, named, and published.{{cite journal |title=Biomineralization. Cell Biology and Mineral Deposition. by Kenneth Simkiss; Karl M. Wilbur On Biomineralization. by Heinz A. Lowenstam; Stephen Weiner |journal=Science |volume=247 |issue=4946 |pages=1129–30 |year=1990 |bibcode=1990Sci...247.1129S |jstor=2874281 |vauthors=Veis A |pmid=17800080 |doi=10.1126/science.247.4946.1129}} These minerals (a sub-set tabulated in Lowenstam (1981)) are considered minerals proper according to Skinner's (2005) definition. These biominerals are not listed in the International Mineral Association official list of mineral names,{{cite web |title=Official IMA list of mineral names |date=March 2009 |work=uws.edu.au |url=http://pubsites.uws.edu.au/ima-cnmnc/IMA%20mineral%20list%20update%20BB%20Upload%208%20April%202011.pdf |url-status=dead |archive-url=https://web.archive.org/web/20110706121228/http://pubsites.uws.edu.au/ima-cnmnc/IMA%20mineral%20list%20update%20BB%20Upload%208%20April%202011.pdf |archive-date=6 July 2011}} however, many of these biomineral representatives are distributed among the 78 mineral classes listed in the Dana classification scheme.

Skinner's (2005) definition of a mineral considers this matter by stating that a mineral can be crystalline or amorphous. Although biominerals are not the most common form of minerals,{{cite book |vauthors=Hefferan K, O'Brien J |title=Earth Materials |date=2010 |isbn=978-1-4443-3460-9 |publisher=Wiley-Blackwell}} they help to define the limits of what constitutes a mineral properly. Nickel's (1995) formal definition explicitly mentioned crystallinity as a key to defining a substance as a mineral. A 2011 article defined icosahedrite, an aluminium-iron-copper alloy as mineral; named for its unique natural icosahedral symmetry, it is a quasicrystal. Unlike a true crystal, quasicrystals are ordered but not periodic.{{cite journal |vauthors=Bindi L, Steinhardt PJ, Yao N, Lu PJ |author-link=Luca Bindi |title=Icosahedrite, Al₆₃Cu₂₄Fe₁₃, the first natural quasicrystal |journal=American Mineralogist |volume=96 |issue=5–6 |pages=928–31 |year=2011 |bibcode=2011AmMin..96..928B |doi=10.2138/am.2011.3758 |s2cid=101152220}}{{cite web |work=Commission on New Minerals and Mineral Names |title=Approved as new mineral |url=http://pubsites.uws.edu.au/ima-cnmnc/newminerals2010.pdf |url-status=dead |archive-url=https://web.archive.org/web/20120320182918/http://pubsites.uws.edu.au/ima-cnmnc/newminerals2010.pdf |archive-date=20 March 2012}}

Examples of biogenic minerals include:{{cite journal |vauthors=Corliss WR |title=Biogenic Minerals |journal=Science Frontiers |date=Nov–Dec 1989 |volume=66 |url=https://www.science-frontiers.com/sf066/sf066g13.htm}}

• Apatite in bones and teeth
• Aragonite, calcite, fluorite in vestibular systems (part of the inner ear) of vertebrates
• Aragonite and calcite in travertine and biogenic silica (siliceous sinter, opal) deposited through algal action
• Goethite found as filaments in limpet teeth
• Hydroxyapatite formed by mitochondria
• Magnetite and greigite formed by magnetotactic bacteria
• Oxalate and calcium carbonate raphides, silica bodies, strontium and barium sulfate in some plants{{cite journal |last1=He |first1=Honghua |last2=Veneklaas |first2=Erik J. |last3=Kuo |first3=John |last4=Lambers |first4=Hans |title=Physiological and ecological significance of biomineralization in plants |date=2014-03-01 |journal=Trends in Plant Science |volume=19 |issue=3 |pages=166–174 |issn=1360-1385 |doi=10.1016/j.tplants.2013.11.002 |pmid=24291440 |bibcode=2014TPS....19..166H |url=https://www.sciencedirect.com/science/article/abs/pii/S1360138513002537|url-access=subscription }}
• Pyrite and marcasite in sedimentary rocks deposited by sulfate-reducing bacteria
• Quartz formed from bacterial action on fossil fuels (gas, oil, coal)

Biominerals could be important indicators of extraterrestrial life and thus could play an essential role in the search for past or present life on Mars. Furthermore, organic components (biosignatures) that are often associated with biominerals are believed to play crucial roles in both pre-biotic and biotic reactions.{{cite book |veditors=Steele A, Beaty D |contribution=Final report of the MEPAG Astrobiology Field Laboratory Science Steering Group (AFL-SSG) |title=The Astrobiology Field Laboratory |publisher=Mars Exploration Program Analysis Group (MEPAG) - NASA |place=U.S.A. |pages=72 |date=26 September 2006 |collaboration=MEPAG Astrobiology Field Laboratory Science Steering Group |contribution-url=http://mepag.jpl.nasa.gov/reports/AFL_SSG_WHITE_PAPER_v3.doc |chapter-format=.doc |access-date=22 July 2009}}

On 24 January 2014, NASA reported that current studies by the Curiosity and Opportunity rovers on the planet Mars will now be searching for evidence of ancient life, including a biosphere based on autotrophic, chemotrophic and chemolithoautotrophic microorganisms, as well as ancient water, including fluvio-lacustrine environments (plains related to ancient rivers or lakes) that may have been habitable.{{cite journal |vauthors=Grotzinger JP |title=Exploring martian habitability. Habitability, taphonomy, and the search for organic carbon on Mars. Introduction |journal=Science |volume=343 |issue=6169 |pages=386–387 |date=January 2014 |bibcode=2014Sci...343..386G |pmid=24458635 |doi=10.1126/science.1249944 |doi-access=free}}{{cite journal |author=Various |title=Special Issue - Table of Contents - Exploring Martian Habitability |date=24 January 2014 |journal=Science |volume=343 |pages=345–452 |url=https://www.science.org/toc/science/343/6169 |access-date=24 January 2014}}{{cite journal |author=Various |title=Special Collection - Curiosity - Exploring Martian Habitability |date=24 January 2014 |journal=Science |url=https://www.science.org/action/doSearch?AllField=Curiosity+Mars |access-date=24 January 2014}}{{cite journal |vauthors=Grotzinger JP, Sumner DY, Kah LC, Stack K, Gupta S, Edgar L, Rubin D, Lewis K, Schieber J, Mangold N, Milliken R, Conrad PG, DesMarais D, Farmer J, Siebach K, Calef F, Hurowitz J, McLennan SM, Ming D, Vaniman D, Crisp J, Vasavada A, Edgett KS, Malin M, Blake D, Gellert R, Mahaffy P, Wiens RC, Maurice S, Grant JA, Wilson S, Anderson RC, Beegle L, Arvidson R, Hallet B, Sletten RS, Rice M, Bell J, Griffes J, Ehlmann B, Anderson RB, Bristow TF, Dietrich WE, Dromart G, Eigenbrode J, Fraeman A, Hardgrove C, Herkenhoff K, Jandura L, Kocurek G, Lee S, Leshin LA, Leveille R, Limonadi D, Maki J, McCloskey S, Meyer M, Minitti M, Newsom H, Oehler D, Okon A, Palucis M, Parker T, Rowland S, Schmidt M, Squyres S, Steele A, Stolper E, Summons R, Treiman A, Williams R, Yingst A |display-authors=6 |title=A habitable fluvio-lacustrine environment at Yellowknife Bay, Gale crater, Mars |journal=Science |volume=343 |issue=6169 |pages=1242777 |date=January 2014 |citeseerx=10.1.1.455.3973 |bibcode=2014Sci...343A.386G |pmid=24324272 |doi=10.1126/science.1242777 |s2cid=52836398}} The search for evidence of habitability, taphonomy (related to fossils), and organic carbon on the planet Mars is now a primary NASA objective.

{{Div col|colwidth=30em}}

• Biocrystallization
• Biofilm
• Biointerface
• Biomineralising polychaetes
• Bone mineral
• Microbiologically induced calcite precipitation
• Micropaleontology
• Mineralized tissues
• Phytolith
• Raphide and Druse (botany)
• Susannah M. Porter history of biomineralization

{{Div col end}}

{{Notelist}}

{{Reflist|35em}}

{{Refbegin|30em}}

• {{cite journal |vauthors=Addadi L, Weiner S |author-link1=Lia Addadi |title=Control And Design Principles In Biological Mineralization |journal=Angewandte Chemie International Edition in English |year=1992 |volume=31 |issue=2 |pages=153–169 |format=abstract |doi=10.1002/anie.199201531 |url=https://onlinelibrary.wiley.com/doi/10.1002/anie.199201531 |url-status=live |archive-url=https://archive.today/20121217192224/http://www3.interscience.wiley.com/cgi-bin/abstract/106587877/ABSTRACT?CRETRY=1&SRETRY=0 |archive-date=17 December 2012|url-access=subscription }}
• {{cite journal |vauthors=Boskey AL |title=Biomineralization: an overview |journal=Connective Tissue Research |volume=44 |issue=Supplement 1 |pages=5–9 |year=2003 |pmid=12952166 |doi=10.1080/713713622}}
• {{cite journal |vauthors=Cuif JP, Sorauf JE |title=Biomineralization and diagenesis in the Scleractinia: part I, biomineralization |journal=Bull. Tohoku Univ. Museum |year=2001 |volume=1 |pages=144–151}}
• {{cite journal |vauthors=Dauphin Y |title=Structures, organo mineral compositions and diagenetic changes in biominerals |journal=Current Opinion in Colloid & Interface Science |year=2002 |volume=7 |issue=1–2 |pages=133–138 |doi=10.1016/S1359-0294(02)00013-4}}
• {{cite journal |vauthors=Dauphin Y |title=Biomineralization |journal=Encyclopedia of Inorganic Chemistry |veditors=King RB |publisher=Wiley & Sons |year=2005 |volume=1 |pages=391–404 |isbn=978-0-521-87473-1}}
• {{cite journal |vauthors=Kupriyanova EK, Vinn O, Taylor PD, Schopf JW, Kudryavtsev AB, Bailey-Brock J |title=Serpulids living deep: calcareous tubeworms beyond the abyss |journal=Deep-Sea Research Part I |year=2014 |volume=90 |pages=91–104 |bibcode=2014DSRI...90...91K |doi=10.1016/j.dsr.2014.04.006 |url=https://www.academia.edu/8904126 |access-date=9 January 2014}}
• {{cite journal |vauthors=Lowenstam HA |title=Minerals formed by organisms |journal=Science |volume=211 |issue=4487 |pages=1126–1131 |date=March 1981 |bibcode=1981Sci...211.1126L |jstor=1685216 |pmid=7008198 |doi=10.1126/science.7008198 |s2cid=31036238}}
• {{cite journal |vauthors=McPhee, Joseph |title=The Little Workers of the Mining Industry |journal=Science Creative Quarterly |year=2006 |issue=2 |url=https://www.scq.ubc.ca/the-little-workers-of-the-mining-industry/ |access-date=3 November 2006}}
• {{cite journal |vauthors=Schmittner KE, Giresse P |title=Micro-environmental controls on biomineralization: superficial processes of apatite and calcite precipitation in Quaternary soils, Roussillon, France |journal=Sedimentology |year=1999 |volume=46 |issue=3 |pages=463–476 |bibcode=1999Sedim..46..463S |doi=10.1046/j.1365-3091.1999.00224.x |s2cid=140680495}}
• {{cite book |vauthors=Uebe R, Schüler D |chapter=The Formation of Iron Biominerals |pages=159–184 |veditors=Kroneck PM, Sosa Torres ME |title=Metals, Microbes, and Minerals - The Biogeochemical Side of Life |date=2021 |publisher=De Gruyter |location=Berlin |isbn=978-3-11-058977-1 |doi=10.1515/9783110589771-006}}
• {{cite journal |vauthors=Vinn O |title=Occurrence, formation and function of organic sheets in the mineral tube structures of Serpulidae (polychaeta, Annelida) |journal=PLOS ONE |volume=8 |issue=10 |pages=e75330 |year=2013 |bibcode=2013PLoSO...875330V |pmid=24116035 |doi=10.1371/journal.pone.0075330 |doi-access=free |pmc=3792063}}
• {{cite journal |vauthors=Vinn O, ten Hove HA, Mutvei H |title=Ultrastructure and mineral composition of serpulid tubes (Polychaeta, Annelida) |journal=Zoological Journal of the Linnean Society |year=2008 |volume=154 |issue=4 |pages=633–650 |doi=10.1111/j.1096-3642.2008.00421.x |doi-access=free |url=https://www.researchgate.net/publication/222089798 |access-date=9 January 2014}}
• {{cite journal |vauthors=Weiner S, Addadi L |author-link2=Lia Addadi |title=Design strategies in mineralized biological materials |journal=Journal of Materials Chemistry |year=1997 |volume=7 |issue=5 |pages=689–702 |doi=10.1039/a604512j}}

{{Refend}}

• [https://biomineralisation.blogspot.com/ 'Data and literature on modern and fossil Biominerals']
• [https://www.scq.ubc.ca/the-little-workers-of-the-mining-industry/ The Little Workers of the Mining Industry] (an overview of the bacteria involved in biomineralization), Science Creative Quarterly
• [https://bio-mineral.org/ Biomineralization web-book: bio-mineral.org]
• [https://archive.today/20140627092658/https://webcast.stsci.edu/webcast/detail.xhtml?talkid=4006 Minerals and the Origins of Life] (Robert Hazen, NASA) (video, 60m, April 2014).
• [https://web.archive.org/web/20161220213016/http://www.spp-biomineralisation.de/ Special German Research Project About the Principles of Biomineralization]

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