Biomineralization - Knowledge

2372: 1865: 1359: 2081: 1960: 2494:. 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. 1123: 2080: 1812: 2225: 1850: 1013: 1542: 1029: 1087:

remaining 1–5%. The resulting composite has a fracture toughness ≈3000 times greater than that of the crystals themselves. 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.

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irreproducible because of the difficulties in controlling agglomeration. 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.

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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

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composition, and growth. 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.

9778: 2003:; to forming minor structures (e.g., ciliate granules; or being a major structural constituent of the organism. 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. Biogeochemically and ecologically, diatoms are the most important silicifiers in modern marine ecosystems, with radiolarians ( 2061:, Lobel et al. (1996) identified by biochemical modeling a low-energy reaction pathway for nucleation and growth of silica. 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 2430: 9754: 8119: 8060: 5457: 4934: 3676: 1106:). 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 ( 2371: 2193:. 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. This suggests that 932:, 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. 1781: 867:

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,

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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

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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, they help to define the limits of what constitutes a mineral properly. Nickel's (1995) formal definition explicitly

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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

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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

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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

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that are involved in inducing mineralization in most animal lineages appear to have performed such an anticalcifatory function in the ancestral state. Further, certain proteins that would originally have been involved in maintaining calcium concentrations within cells are homologous in all animals,

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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. However, the vast diversity of organisms that thrive in a complex array of biotic and abiotic interactions in oceanic ecosystems

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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

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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

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A topic of contention among geologists and mineralogists has been the IMA's decision to exclude biogenic crystalline substances. 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."

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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.

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transmission and modulation in organisms as diverse as plants, diatoms, sponges, and molluscs. There is also evidence that silicification is used as a detoxification response in snails and plants, biosilica has even been suggested to play a role as a pH buffer for the enzymatic activity of

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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

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being many times more flexible than an equivalent structure made of pure silica. As a result, biogenic silica structures are used for support, feeding, predation defense and environmental protection as a component of cyst walls. Biogenic silica also has useful optical properties for light

8170: 2221:, implying an ancestral role. 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. 1524:

of tooth enamel involves the reintroduction of mineral ions into demineralised enamel. Hydroxyapatite is the main mineral component of enamel in teeth. During demineralisation, calcium and phosphorus ions are drawn out from the hydroxyapatite. The mineral ions introduced during remineralisation

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Before the International Mineralogical Association's listing, over 60 biominerals had been discovered, named, and published. 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

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The phylogeny shown in this diagram is based on Adl et al. (2012), 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

2181:"off-the-shelf" components already used for other purposes in the organism. 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 911:

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

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and appear to have been co-opted into biomineralization after the divergence of the animal lineages. 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

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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

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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

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Biomineralization evolved multiple times, independently, and most animal lineages first expressed biomineralized components in the Cambrian period. Many of the same processes are used in unrelated lineages, which suggests that biomineralization machinery was assembled from pre-existing

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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. Fungal growth may produce a copper-containing mineral precipitate, such as

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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

2267:, was duplicated and modified in the Cambrian to form the basis for calcium-based biomineralization pathways. 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 1047:

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

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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. This indicates that biomineralisation is likely ancestral to all lophotrochozoans.

2291:. 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 2044:

Broadly, biomineralized structures evolve and diversify when the energetic cost of biomineral production is less than the expense of producing an equivalent organic structure. The energetic costs of forming a silica structure from

1900:, 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 2345:

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.

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at the point in time when they became biomineralized, and stuck with that form for the remainder of their biological history (but see for a more detailed analysis). The stability is dependent on the Ca/Mg ratio of

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Onozato H, Watabe N (October 1979). "Studies on fish scale formation and resorption. III. Fine structure and calcification of the fibrillary plates of the scales in Carassius auratus (Cypriniformes: Cyprinidae)".

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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.

1145:. 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 1532:

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. Their dactyl appendages have excellent

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International Mineral Association official list of mineral names, however, many of these biomineral representatives are distributed amongst the 78 mineral classes listed in the Dana classification scheme.

1204:. 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 5907:

Weaver JC, Aizenberg J, Fantner GE, Kisailus D, Woesz A, Allen P, et al. (April 2007). "Hierarchical assembly of the siliceous skeletal lattice of the hexactinellid sponge Euplectella aspergillum".

4626: 736:. It is an extremely widespread phenomenon: all six taxonomic kingdoms contain members that are able to form minerals, and over 60 different minerals have been identified in organisms. Examples include 6640:

Gal A, Hirsch A, Siegel S, Li C, Aichmayer B, Politi Y, et al. (August 2012). "Plant cystoliths: a complex functional biocomposite of four distinct silica and amorphous calcium carbonate phases".

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González-Muñoz MT, Rodriguez-Navarro C, Martínez-Ruiz F, Arias JM, Merroun ML, Rodriguez-Gallego M (2010). "Bacterial biomineralization: new insights from Myxococcus-induced mineral precipitation".

880:, 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 1028: 8572:

Takai K (2010). "Limits of life and the biosphere: Lessons from the detection of microorganisms in the deep sea and deep subsurface of the Earth.". In Gargaud M, Lopez-Garcia P, Martin H (eds.).

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Darley WM, Volcani BE (December 1969). "Role of silicon in diatom metabolism. A silicon requirement for deoxyribonucleic acid synthesis in the diatom Cylindrotheca fusiformis Reimann and Lewin".

2299:, 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. 1538:

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.

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can also be important, either structurally or as intermediate phases in biomineralization. Some biominerals include a mixture of these phases in distinct, organised structural components (e.g.

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Pokroy B, Kabalah-Amitai L, Polishchuk I, DeVol RT, Blonsky AZ, Sun CY, Marcus MA, Scholl A, Gilbert PU (13 October 2015). "Narrowly Distributed Crystal Orientation in Biomineral Vaterite".

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Schmittner KE, Giresse P (1999). "Micro-environmental controls on biomineralization: superficial processes of apatite and calcite precipitation in Quaternary soils, Roussillon, France".

2580:. 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 1811: 2365:. 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. 2108:

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.

8897: 2446:. The biomineralization of uranium primarily involves the precipitation of uranium phosphate minerals associated with the release of phosphate by microorganisms. Negatively charged 1864: 2568:

and minerals that may shed new light on this question. For example, the IMA-commissioned "Working Group on Environmental Mineralogy and Geochemistry " deals with minerals in the

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Checa AG, Ramírez-Rico J, González-Segura A, Sánchez-Navas A (January 2009). "Nacre and false nacre (foliated aragonite) in extant monoplacophorans (=Tryblidiida: Mollusca)".

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Grotzinger JP, Sumner DY, Kah LC, Stack K, Gupta S, Edgar L, et al. (January 2014). "A habitable fluvio-lacustrine environment at Yellowknife Bay, Gale crater, Mars".

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Pondaven P, Gallinari M, Chollet S, Bucciarelli E, Sarthou G, Schultes S, Jean F (January 2007). "Grazing-induced changes in cell wall silicification in a marine diatom".

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minerals. Degradation of minerals by fungi is carried out through a process known as neogenesis. The order of most to least oxalic acid secreted by the fungi studied are

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Veis A (1990). "Biomineralization. Cell Biology and Mineral Deposition. by Kenneth Simkiss; Karl M. Wilbur On Biomineralization. by Heinz A. Lowenstam; Stephen Weiner".

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organisms were employing the same elements, albeit for a different purpose – perhaps to avoid the inadvertent precipitation of calcium carbonate from the supersaturated

4953:"Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies" 1227:, that localizes and accumulates the uranium minerals that have been precipitated. Although uranium is often deemed as toxic to living organisms, certain fungi such as 6189:

Yamanaka S, Yano R, Usami H, Hayashida N, Ohguchi M, Takeda H, Yoshino K (2008). "Optical properties of diatom silica frustule with special reference to blue light".

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Maloof AC, Porter SM, Moore JL, Dudás FÖ, Bowring SA, Higgins JA, Fike DA, Eddy MP (2010). "The earliest Cambrian record of animals and ocean geochemical change".

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Maloof AC, Rose CV, Beach R, Samuels BM, Calmet CC, Erwin DH, et al. (2010). "Possible animal-body fossils in pre-Marinoan limestones from South Australia".

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Gadd GM (January 2007). "Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation".

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Nanophase and Nanocomposite Materials: Symposium Held December 1–3, 1992, Boston, Massachusetts, U.S.A. (Materials Research Society Symposium Proceedings)

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Sone ED, Weiner S, Addadi L (June 2007). "Biomineralization of limpet teeth: a cryo-TEM study of the organic matrix and the onset of mineral deposition".

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Towe KM, Lowenstam HA (January 1967). "Ultrastructure and development of iron mineralization in the radular teeth of Cryptochiton stelleri (Mollusca)".

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Raven JA, Giordano M (March 2009). "Biomineralization by photosynthetic organisms: evidence of coevolution of the organisms and their environment?".

2924: 2535:(IMA) is the generally recognized standard body for the definition and nomenclature of mineral species. As of December 2020, the IMA recognizes 676: 8137: 3233:

Dupraz C, Reid RP, Braissant O, Decho AW, Norman RS, Visscher PT (1 October 2009). "Processes of carbonate precipitation in modern microbial mats".

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Biominerals could be important indicators of extraterrestrial life and thus could play an essential role in the search for past or present life on

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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

2892: 393: 8168:, Dosier GK, "Methods for making construction material using enzyme producing bacteria", issued 2014, assigned to Biomason Inc. 4783:"The 'scaly-foot gastropod': a new genus and species of hydrothermal vent-endemic gastropod (Neomphalina: Peltospiridae) from the Indian Ocean" 1170:

domain. Studies of their significant roles in geological processes, "geomycology", have shown that fungi are involved with biomineralization,

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deposition process and enamel deposition in vertebrates, but some vertebrates even have comparable iron storage facilities near their teeth.

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of biomineralization pathways is underlined by a remarkable experiment whereby the nacreous layer of a molluscan shell was implanted into a

1831: 4027: 8908: 7479:"Differential expression of three galaxin-related genes during settlement and metamorphosis in the scleractinian coral Acropora millepora" 1350:

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

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defines biomineralization as "mineralization caused by cell-mediated phenomena" and notes that it "is a process generally concomitant to

888:. Additionally, the mineralization of collagen provides crucial compressive strength for the bones, cartilage, and teeth of vertebrates. 6850:

Cohen PA, Schopf JW, Butterfield NJ, Kudryavtsev AB, Macdonald FA (2011). "Phosphate biomineralization in mid-Neoproterozoic protists".

5850:"Glassin, a histidine-rich protein from the siliceous skeletal system of the marine sponge Euplectella, directs silica polycondensation" 614: 9008: 3405:

Görgen, Sigrid; Benzerara, Karim; Skouri-Panet, Fériel; Gugger, Muriel; Chauvat, Franck; Cassier-Chauvat, Corinne (30 November 2020).

2243:, and rather than experiencing an immune response, the molluscan nacre was incorporated into the host bone matrix. This points to the 1136:, leading to the release of bioavailable metals or nutrients, which eventually may be uptaken by living organisms or precipitated as 2508:

The geological definition of mineral normally excludes compounds that occur only in living beings. However, some minerals are often

4196:"Amorphous and crystalline calcium carbonate distribution in the tergite cuticle of moulting Porcellio scaber (Isopoda, Crustacea)" 1114:), though some amorphous calcium carbonate occurs as well; and although they react as crystals, they never show angles and facets. 9723: 4061:"Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp Odontodactylus scyllarus" 9071: 8363: 2532: 655: 8367: 9298: 4699:

Joester D, Brooker LR (5 July 2016). "The Chiton Radula: A Model System for Versatile Use of Iron Oxides*". In Faivre D (ed.).

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One biological system that might be of key importance in the future development of architecture is bacterial biofilm. The term

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to give structural support to bones and shells. The structures of these biocomposite materials are highly controlled from the

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Wood RA, Grotzinger JP, Dickson JA (June 2002). "Proterozoic modular biomineralized metazoan from the Nama Group, Namibia".

4464: 1456:). Carbonates are particularly prevalent in marine environments, but also present in freshwater and terrestrial organisms. 9591:"Occurrence, formation and function of organic sheets in the mineral tube structures of Serpulidae (polychaeta, Annelida)" 3075:"Occurrence, formation and function of organic sheets in the mineral tube structures of Serpulidae (polychaeta, Annelida)" 5655:

Lobel KD, West JK, Hench LL (1996). "Computational model for protein-mediated biomineralization of the diatom frustule".

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Newman DK, Banfield JF (May 2002). "Geomicrobiology: how molecular-scale interactions underpin biogeochemical systems".

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Finkel ZV, Kotrc B (2010). "Silica Use Through Time: Macroevolutionary Change in the Morphology of the Diatom Fustule".

4542:"Nano-hydroxyapatite and its applications in preventive, restorative and regenerative dentistry: a review of literature" 3793:

Liang X, Hillier S, Pendlowski H, Gray N, Ceci A, Gadd GM (June 2015). "Uranium phosphate biomineralization by fungi".

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Oppenheimer-Shaanan Y, Sibony-Nevo O, Bloom-Ackermann Z, Suissa R, Steinberg N, Kartvelishvily E, et al. (2016).

7757: 7727: 686:: Complete conversion of organic substances to inorganic derivatives by living organisms, especially micro-organisms. 8149: 7767: 7737: 7595:"Brachiopod and mollusc biomineralisation is a conserved process that was lost in the phoronid-bryozoan stem lineage" 5485: 4125: 17: 9338:

Cuif JP, Sorauf JE (2001). "Biomineralization and diagenesis in the Scleractinia : part I, biomineralization".

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Adamo P, Violante P (1 May 2000). "Weathering of rocks and neogenesis of minerals associated with lichen activity".

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can vary even between closely related taxa, from being found in composite structures with other biominerals (e.g.,

9737: 9098:"Exploring martian habitability. Habitability, taphonomy, and the search for organic carbon on Mars. Introduction" 9837: 6585:"A first report of hydroxylated apatite as structural biomineral in Loasaceae - plants' teeth against herbivores" 5371:

Martin-Jézéquel V, Hildebrand M, Brzezinski MA (2000). "Silicon Metabolism in Diatoms: Implications for Growth".

5251:"A cover of glass: first report of biomineralized silicon in a ciliate, Maryna umbrellata (Ciliophora: Colpodea)" 2384:

A directed growth of the calcium carbonate crystals allows mechanical support of the 3D structure. The bacterial

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5425:"Competition between Silicifiers and Non-silicifiers in the Past and Present Ocean and Its Evolutionary Impacts" 2033:

with prominent roles as well. In contrast, the major silicifiers in terrestrial ecosystems are the land plants (

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Nesbit KT, Roer RD (December 2016). "Silicification of the medial tooth in the blue crab Callinectes sapidus".

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and roughness, determine initial bacterial attachment. A common principle of all biofilms is the production of

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Gwenzi W (2019). "Carbon Sequestration via Biomineralization: Processes, Applications and Future Directions".

3626:"Programming pluripotent precursor cells derived from Xenopus embryos to generate specific tissues and organs" 3463:"A genome-wide analysis of biomineralization-related proteins in the sea urchin Strongylocentrotus purpuratus" 9842: 5612:

Hecky RE, Mopper K, Kilham P, Degens ET (1973). "The amino acid and sugar composition of diatom cell-walls".

2561: 2213:; the role performed appears to be functionally identical to that of the unrelated pearlin gene in molluscs. 525: 520: 353: 154: 7644:

Kirschvink JL, Hagadorn JW (2000). "10 A Grand Unified theory of Biomineralization.". In Bäuerlein E (ed.).

6769:"The impact of light intensity and daylength on silicate and nitrate competition among marine phytoplankton" 6103:

Schaller J, Brackhage C, Bäucker E, Dudel EG (June 2013). "UV-screening of grasses by plant silica layer?".

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9782: 4985: 2882: 1449: 348: 6334:"A dynamic broadband reflector built from microscopic silica spheres in the 'disco' clam Ctenoides ales" 5758:"Microstructure provides insights into evolutionary design and resilience of Coscinodiscus sp. frustule" 9822: 9744: 4873: 3851:

Adeyemi AO, Gadd GM (June 2005). "Fungal degradation of calcium-, lead- and silicon-bearing minerals".

2776:) that are often associated with biominerals are believed to play crucial roles in both pre-biotic and 635: 582: 8165: 6383:"Aluminum-dependent regulation of intracellular silicon in the aquatic invertebrate Lymnaea stagnalis" 2251:

The most ancient example of biomineralization, dating back 2 billion years, is the deposition of

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Uebe R, Schüler D (2021). "The Formation of Iron Biominerals". In Kroneck PM, Sosa Torres ME (eds.).

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Weaver JC, Milliron GW, Miserez A, Evans-Lutterodt K, Herrera S, Gallana I, et al. (June 2012).

2599: 1521: 576: 235: 9403: 9231: 4986:"Acantharian cysts: High flux occurrence in the bathypelagic zone of the Scotia Sea, Southern Ocean" 3553:

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Livingston BT, Killian CE, Wilt F, Cameron A, Landrum MJ, Ermolaeva O, et al. (December 2006).

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In nature, there is a wide array of biominerals, ranging from iron oxide to strontium sulfate, with

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Dauphin Y (2002). "Structures, organo mineral compositions and diagenetic changes in biominerals".

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Dragoš A, Kovács ÁT (April 2017). "The Peculiar Functions of the Bacterial Extracellular Matrix".

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Hendry KR, Marron AO, Vincent F, Conley DJ, Gehlen M, Ibarbalz FM, Quéguiner B, Bowler C (2018).

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Knoll AH, Kotrc B (2015). "Protistan Skeletons: A Geologic History of Evolution and Constraint".

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biominerals being particularly notable. However, the most taxonomically widespread biomineral is

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Fuhrmann T, Landwehr S, Rharbi-Kucki E, Sumper M (2004). "Diatoms as living photonic crystals".

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Environmental microbe-metal interactions: Bioremediation of radionuclide-containing wastewaters

5700:"Architecture and material properties of diatom shells provide effective mechanical protection" 2834: 2830: 2722: 1896:

form celestine crystal shells. The denseness of the celestite ensures their shells function as

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and the brains of vertebrates; it is possible that this pathway, which performed a

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