Biomineralization

Last updated

IUPAC definition

Biomineralization: Complete conversion of organic substances to inorganic derivatives by living organisms, especially micro-organisms. [1]

Contents

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

Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, [a] often resulting in hardened or stiffened mineralized tissues . It is an extremely widespread phenomenon: all six taxonomic kingdoms contain members that are able to form minerals, and over 60 different minerals have been identified in organisms. [2] [3] [4] Examples include silicates in algae and diatoms, carbonates in invertebrates, and calcium phosphates and carbonates in vertebrates. These minerals often form structural features such as sea shells and the bone in mammals and birds.

Organisms have been producing mineralized skeletons for the past 550 million years. Calcium carbonates and calcium phosphates are usually crystalline, but silica organisms (such as sponges and diatoms) are always non-crystalline minerals. Other examples include copper, iron, and gold deposits involving bacteria. Biologically formed minerals often have special uses such as magnetic sensors in magnetotactic bacteria (Fe3O4), gravity-sensing devices (CaCO3, CaSO4, BaSO4) and iron storage and mobilization (Fe2O3•H2O in the protein ferritin).

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

Types

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

Biomineralization

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

Organomineralization

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

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

Biological roles

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

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

In molluscs

A variety of mollusc shells Shells-IZE-006-Black.jpg
A variety of mollusc shells

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

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

In fungi

Global involvement of fungi in some biogeochemical cycles (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. Global involvement of fungi in some biogeochemical cycles.png
Global involvement of fungi in some biogeochemical cycles 
(a) Fungi contribute substantially to mineral weathering, leading to the release of bioavailable metals or nutrients, which eventually may be uptaken by living organisms or precipitated as secondary minerals
(b) Fungi as heterotrophs, recycle organic matter. While doing so, they produce metabolites such as organic acids that can also precipitate as secondary minerals (salts). Recycling organic matter eventually releases constitutive elements such as C, N, P, and S
(c) CO2 produced by heterotrophic fungal respiration can dissolve into H2O and depending on the physicochemical conditions precipitate as CaCO3 leading to the formation of a secondary mineral.

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

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

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

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

In bacteria

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

Other roles

The chalk of the White Cliffs of Dover is almost entirely formed from fossil skeleton remains (coccoliths), biomineralized by planktonic microorganisms (coccolithophores). White cliffs of dover 09 2004.jpg
The chalk 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   [17] and as a carbon sink. [24]

Composition

Most biominerals can be grouped by chemical composition into one of three distinct mineral classes: silicates, carbonates, or phosphates. [25]

Silicates

A testate amoeba which has covered itself with protective diatom frustules Collection Penard MHNG Specimen 533-2-1 Pamphagus granulatus.tif
A testate amoeba which has covered itself with protective diatom frustules
Peacock mantis shrimp smash their prey by swinging club-like raptorial claws made of hydroxyapatite . Odontodactylus scyllarus 2.png
Peacock mantis shrimp smash their prey by swinging club-like raptorial claws made of hydroxyapatite  .

Silicates (glass) are common in marine biominerals, where diatoms form frustules and radiolaria form capsules from hydrated amorphous silica (opal). [27]

Carbonates

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 [28] [29] or as intermediate phases in biomineralization. [30] [31] 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. [32]

Phosphates

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. [33] 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. [34] Remineralisation of tooth enamel involves the reintroduction of mineral ions into demineralised enamel. [35] Hydroxyapatite is the main mineral component of enamel in teeth. [36] 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. [36]

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. [37] 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. [38]

Glomerula piloseta (Sabellidae), longitudinal section of the tube showing aragonitic spherulitic prismatic structure Glomerula piloseta tube microstructure.jpg
Glomerula piloseta (Sabellidae), longitudinal section of the tube showing aragonitic spherulitic prismatic structure
CompositionExample organisms
Calcium carbonate
(calcite or aragonite)
Silica
(silicate/glass/opal)
Apatite
(phosphate minerals)

Other minerals

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 [39] or goethite in limpets. [40] Gastropod molluscs living close to hydrothermal vents reinforce their carbonate shells with the iron-sulphur minerals pyrite and greigite. [41] Magnetotactic bacteria also employ magnetic iron minerals magnetite and greigite to produce magnetosomes to aid orientation and distribution in the sediments.

Small magnetosome.jpg
Magnetotactic bacterium containing a chain of magnetosomes
Magnetosome chain.png
Magnetosome chain with octahedral habits modelled lower right [42]

Celestine, the heaviest mineral in the ocean, consists of strontium sulfate, SrSO4. The mineral is named for the delicate blue colour of its crystals. [44] 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. [45] [46] [44]

Diversity

The most widespread biomineral is silica
Diatom - Triceratium favus.jpg
The siliceous diatom frustule has the highest strength of any known biological material.
Sponge-spicule hg.jpg
Sponge spicules, like this from a siliceous glass sponge, form structures many times more flexible than equivalent structures made of pure silica.
Radiolarian - Podocyrtis (Lampterium) mitra Ehrenberg - 160x.jpg
Transparent glass test or shell of a radiolarian

In nature, there is a wide array of biominerals, ranging from iron oxide to strontium sulfate, [47] with calcareous biominerals being particularly notable. [48] [49] However, the most taxonomically widespread biomineral is silica (SiO2·nH2O), being present in all eukaryotic supergroups. [50] 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; [51] to forming minor structures (e.g., ciliate granules; [52] or being a major structural constituent of the organism. [53] 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. [54] [55] 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. [56]

Broadly, biomineralized structures evolve and diversify when the energetic cost of biomineral production is less than the expense of producing an equivalent organic structure. [57] [58] [59] 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). [60] Based on a structural model of biogenic silica, [61] Lobel et al. (1996) identified by biochemical modeling a low-energy reaction pathway for nucleation and growth of silica. [62] 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, [63] [64] or sponge spicules being many times more flexible than an equivalent structure made of pure silica. [65] [66] As a result, biogenic silica structures are used for support, [67] feeding, [68] predation defense  [69] [70] [71] and environmental protection as a component of cyst walls. [53] Biogenic silica also has useful optical properties for light transmission and modulation in organisms as diverse as plants, [72] diatoms, [73] [74] [75] sponges, [76] and molluscs. [77] There is also evidence that silicification is used as a detoxification response in snails  [78] and plants, [79] 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. [80] [56]

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. [56] 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. [85] [86] 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. [56]

Evolution

Some calcareous sponges (Ernst Haeckel, Kunstformen der Natur) Haeckel Calcispongiae.jpg
Some calcareous sponges (Ernst Haeckel, Kunstformen der Natur )

The first evidence of biomineralization dates to some 750  million years ago, [87] [88] and sponge-grade organisms may have formed calcite skeletons 630  million years ago. [89] But in most lineages, biomineralization first occurred in the Cambrian or Ordovician periods. [90] Organisms used whichever form of calcium carbonate was more stable in the water column at the point in time when they became biomineralized, [91] and stuck with that form for the remainder of their biological history [92] (but see [93] 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. [91]

Biomineralization evolved multiple times, independently, [94] and most animal lineages first expressed biomineralized components in the Cambrian period. [95] 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. [25] 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. [96] 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. [13] [97] 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. [96] 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. [98] Further, certain proteins that would originally have been involved in maintaining calcium concentrations within cells [99] are homologous in all animals, and appear to have been co-opted into biomineralization after the divergence of the animal lineages. [100] 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. [101] Carbonic anhydrase serves a role in mineralization broadly in the animal kingdom, including in sponges, implying an ancestral role. [102] 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. [97]

Stromatolites made by bacteria. Fossilized stromatolites record some of the earliest life. Highborme Cay-Stromatolite-04 (cropped) 2.jpg
Stromatolites 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. [103] 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. [104] 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. [105]

Potential applications

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. [106] 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. [107] [108]

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. [107] [108]

Architecture

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. [109]

The properties of the surface, such as charge, hydrophobicity and roughness, determine initial bacterial attachment. [110] 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. [111] 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. [111] [112] [113] [114]

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. [116] This process can also be used to manufacture new hard materials, such as bio-cement. [117] [114]

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. [118] [119] [114]

Uranium contaminants

Biomineralization may be used to remediate groundwater contaminated with uranium. [120] 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. [120] The high concentration of ligands near the cell surface also provides nucleation foci for precipitation, which leads to higher efficiency than chemical precipitation. [121]

Biogenic mineral controversy

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 ]

The International Mineralogical Association (IMA) is the generally recognized standard body for the definition and nomenclature of mineral species. As of December 2020, the IMA recognizes 5,650 official mineral species [122] out of 5,862 proposed or traditional ones. [123]

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." [124]

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. [125]

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. [126] [125] For example, the IMA-commissioned "Working Group on Environmental Mineralogy and Geochemistry " deals with minerals in the hydrosphere, atmosphere, and biosphere. [127] 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). [128] [129] [130]

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. [131] [132] [133]

Before the International Mineralogical Association's listing, over 60 biominerals had been discovered, named, and published. [134] These minerals (a sub-set tabulated in Lowenstam (1981) [124] ) are considered minerals proper according to Skinner's (2005) definition. [125] These biominerals are not listed in the International Mineral Association official list of mineral names, [135] however, many of these biomineral representatives are distributed amongst the 78 mineral classes listed in the Dana classification scheme. [125]

Skinner's (2005) definition of a mineral considers this matter by stating that a mineral can be crystalline or amorphous. [125] Although biominerals are not the most common form of minerals, [136] 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. [126] 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. [137] [138]

List of minerals

Examples of biogenic minerals include: [139]

Astrobiology

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. [140]

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. [141] [142] [143] [144] The search for evidence of habitability, taphonomy (related to fossils), and organic carbon on the planet Mars is now a primary NASA objective. [141] [142]

See also

Notes

  1. ↑ The International Union of Pure and Applied Chemistry defines biomineralization as "mineralization caused by cell-mediated phenomena" and notes that it "is a process generally concomitant to biodegradation". [1]

Related Research Articles

<span class="mw-page-title-main">Calcite</span> Calcium carbonate mineral

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratch hardness comparison. Large calcite crystals are used in optical equipment, and limestone composed mostly of calcite has numerous uses.

<span class="mw-page-title-main">Exoskeleton</span> External skeleton of an organism

An exoskeleton is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton which is enclosed underneath other soft tissues. Some large, hard and non-flexible protective exoskeletons are known as shell or armour.

<span class="mw-page-title-main">Aragonite</span> Calcium carbonate mineral

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

<i>Nanobacterium</i> Genus of bacteria

Nanobacterium is the unit or member name of a former proposed class of living organisms, specifically cell-walled microorganisms, now discredited, with a size much smaller than the generally accepted lower limit for life. Originally based on observed nano-scale structures in geological formations, the status of nanobacteria was controversial, with some researchers suggesting they are a new class of living organism capable of incorporating radiolabeled uridine, and others attributing to them a simpler, abiotic nature. One skeptic dubbed them "the cold fusion of microbiology", in reference to a notorious episode of supposed erroneous science. The term "calcifying nanoparticles" (CNPs) has also been used as a conservative name regarding their possible status as a life form.

<span class="mw-page-title-main">Microfossil</span> Fossil that requires the use of a microscope to see it

A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, such as a hand lens, is referred to as a macrofossil.

<span class="mw-page-title-main">Marine sediment</span> Accumulated material on seafloor

Marine sediment, or ocean sediment, or seafloor sediment, are deposits of insoluble particles that have accumulated on the seafloor. These particles either have their origins in soil and rocks and have been transported from the land to the sea, mainly by rivers but also by dust carried by wind and by the flow of glaciers into the sea, or they are biogenic deposits from marine organisms or from chemical precipitation in seawater, as well as from underwater volcanoes and meteorite debris.

<span class="mw-page-title-main">Calcite sea</span> Sea chemistry favouring low-magnesium calcite as the inorganic calcium carbonate precipitate

A calcite sea is a sea in which low-magnesium calcite is the primary inorganic marine calcium carbonate precipitate. An aragonite sea is the alternate seawater chemistry in which aragonite and high-magnesium calcite are the primary inorganic carbonate precipitates. The Early Paleozoic and the Middle to Late Mesozoic oceans were predominantly calcite seas, whereas the Middle Paleozoic through the Early Mesozoic and the Cenozoic are characterized by aragonite seas.

<span class="mw-page-title-main">Siliceous ooze</span> Biogenic pelagic sediment located on the deep ocean floor

Siliceous ooze is a type of biogenic pelagic sediment located on the deep ocean floor. Siliceous oozes are the least common of the deep sea sediments, and make up approximately 15% of the ocean floor. Oozes are defined as sediments which contain at least 30% skeletal remains of pelagic microorganisms. Siliceous oozes are largely composed of the silica based skeletons of microscopic marine organisms such as diatoms and radiolarians. Other components of siliceous oozes near continental margins may include terrestrially derived silica particles and sponge spicules. Siliceous oozes are composed of skeletons made from opal silica SiO2·nH2O, as opposed to calcareous oozes, which are made from skeletons of calcium carbonate (CaCO3·nH2O) organisms (i.e. coccolithophores). Silica (Si) is a bioessential element and is efficiently recycled in the marine environment through the silica cycle. Distance from land masses, water depth and ocean fertility are all factors that affect the opal silica content in seawater and the presence of siliceous oozes.

<span class="mw-page-title-main">Mollusc shell</span> Exoskeleton of an animal in the phylum Mollusca

The molluscshell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes. Not all shelled molluscs live in the sea; many live on the land and in freshwater.

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

<span class="mw-page-title-main">Mineralized tissues</span> Biological tissues incorporating minerals

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

<span class="mw-page-title-main">Amorphous calcium carbonate</span>

Amorphous calcium carbonate (ACC) is the amorphous and least stable polymorph of calcium carbonate. ACC is extremely unstable under normal conditions and is found naturally in taxa as wide-ranging as sea urchins, corals, mollusks, and foraminifera. It is usually found as a monohydrate, holding the chemical formula CaCO3·H2O; however, it can also exist in a dehydrated state, CaCO3. ACC has been known to science for over 100 years when a non-diffraction pattern of calcium carbonate was discovered by Sturcke Herman, exhibiting its poorly-ordered nature.

<span class="mw-page-title-main">Pupa Gilbert</span> Italian-American biophysicist and geobiologist

Pupa Gilbert is an American biophysicist and geobiologist. She has been pioneering synchrotron spectromicroscopy methods since 1989, and she continues to use and develop them today. Since 2004 she has focused on biomineralization in sea urchins, mollusk shells, and tunicates. She and her group are frequent users of the Berkeley-Advanced Light Source.

<span class="mw-page-title-main">Marine biogenic calcification</span> Shell formation mechanism

Marine biogenic calcification is the production of calcium carbonate by organisms in the global ocean.

<span class="mw-page-title-main">Silica cycle</span> Biogeochemical cycle

The silica cycle is the biogeochemical cycle in which biogenic silica is transported between the Earth's systems. Silicon is considered a bioessential element and is one of the most abundant elements on Earth. The silica cycle has significant overlap with the carbon cycle and plays an important role in the sequestration of carbon through continental weathering, biogenic export and burial as oozes on geologic timescales.

<span class="mw-page-title-main">Lia Addadi</span> Israeli biochemist

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

<span class="mw-page-title-main">Patricia Dove</span> American geochemist and crystal growth researcher

Patricia Martin Dove is an American geochemist. She is a university distinguished professor and the C.P. Miles Professor of Science at Virginia Tech with appointments in the department of Geosciences, department of Chemistry, and department of Materials Science and Engineering. Her research focuses on the kinetics and thermodynamics of mineral reactions with aqueous solutions in biogeochemical systems. Much of her work is on crystal nucleation and growth during biomineralization and biomaterial interactions with mineralogical systems. She was elected a member of the National Academy of Sciences (NAS) in 2012 and currently serves as chair of Class I, Physical and Mathematical Sciences.

<span class="mw-page-title-main">Protist shell</span> Protective shell of a type of eukaryotic organism

Many protists have protective shells or tests, usually made from silica (glass) or calcium carbonate (chalk). Protists are a diverse group of eukaryote organisms that are not plants, animals, or fungi. They are typically microscopic unicellular organisms that live in water or moist environments.

<span class="mw-page-title-main">Particulate inorganic carbon</span>

Particulate inorganic carbon (PIC) can be contrasted with dissolved inorganic carbon (DIC), the other form of inorganic carbon found in the ocean. These distinctions are important in chemical oceanography. Particulate inorganic carbon is sometimes called suspended inorganic carbon. In operational terms, it is defined as the inorganic carbon in particulate form that is too large to pass through the filter used to separate dissolved inorganic carbon.

<span class="mw-page-title-main">Silicification</span> Geological petrification process

In geology, silicification is a petrification process in which silica-rich fluids seep into the voids of Earth materials, e.g., rocks, wood, bones, shells, and replace the original materials with silica (SiO2). Silica is a naturally existing and abundant compound found in organic and inorganic materials, including Earth's crust and mantle. There are a variety of silicification mechanisms. In silicification of wood, silica permeates into and occupies cracks and voids in wood such as vessels and cell walls. The original organic matter is retained throughout the process and will gradually decay through time. In the silicification of carbonates, silica replaces carbonates by the same volume. Replacement is accomplished through the dissolution of original rock minerals and the precipitation of silica. This leads to a removal of original materials out of the system. Depending on the structures and composition of the original rock, silica might replace only specific mineral components of the rock. Silicic acid (H4SiO4) in the silica-enriched fluids forms lenticular, nodular, fibrous, or aggregated quartz, opal, or chalcedony that grows within the rock. Silicification happens when rocks or organic materials are in contact with silica-rich surface water, buried under sediments and susceptible to groundwater flow, or buried under volcanic ashes. Silicification is often associated with hydrothermal processes. Temperature for silicification ranges in various conditions: in burial or surface water conditions, temperature for silicification can be around 25°−50°; whereas temperatures for siliceous fluid inclusions can be up to 150°−190°. Silicification could occur during a syn-depositional or a post-depositional stage, commonly along layers marking changes in sedimentation such as unconformities or bedding planes.

References

  1. 1 2 Vert M, Doi Y, Hellwich KH, Hess M, Hodge P, Kubisa P, Rinaudo M, Schué F (11 January 2012). "Terminology for biorelated polymers and applications (IUPAC Recommendations 2012)". Pure and Applied Chemistry. 84 (2): 377–410. doi: 10.1351/PAC-REC-10-12-04 . S2CID   98107080.
  2. ↑ Sigel A, Sigel H, Sigel RK, eds. (2008). Biomineralization: From Nature to Application. Metal Ions in Life Sciences. Vol. 4. Wiley. ISBN   978-0-470-03525-2.
  3. ↑ Weiner S, Lowenstam HA (1989). On biomineralization. Oxford [Oxfordshire]: Oxford University Press. ISBN   978-0-19-504977-0.
  4. 1 2 Cuif JP, Dauphin Y, Sorauf JE (2011). Biominerals and fossils through time. Cambridge. ISBN   978-0-521-87473-1.
  5. ↑ Vinn O (2013). "Occurrence, formation and function of organic sheets in the mineral tube structures of Serpulidae (polychaeta, Annelida)". PLOS ONE. 8 (10): e75330. Bibcode:2013PLoSO...875330V. doi: 10.1371/journal.pone.0075330 . PMC   3792063 . PMID   24116035.
  6. ↑ Boskey AL (1998). "Biomineralization: conflicts, challenges, and opportunities". Journal of Cellular Biochemistry. 30–31 (S30-31): 83–91. doi:10.1002/(SICI)1097-4644(1998)72:30/31+<83::AID-JCB12>3.0.CO;2-F. PMID   9893259. S2CID   46004807.
  7. ↑ Sarikaya M (December 1999). "Biomimetics: materials fabrication through biology". Proceedings of the National Academy of Sciences of the United States of America. 96 (25): 14183–14185. Bibcode:1999PNAS...9614183S. doi: 10.1073/pnas.96.25.14183 . PMC   33939 . PMID   10588672.
  8. 1 2 Dupraz C, Reid RP, Braissant O, Decho AW, Norman RS, Visscher PT (1 October 2009). "Processes of carbonate precipitation in modern microbial mats". Earth-Science Reviews. Microbial Mats in Earth's Fossil Record of Life: Geobiology. 96 (3): 141–162. Bibcode:2009ESRv...96..141D. doi:10.1016/j.earscirev.2008.10.005.
  9. ↑ Sherman VR, Yang W, Meyers MA (December 2015). "The materials science of collagen". Journal of the Mechanical Behavior of Biomedical Materials. 52: 22–50. doi: 10.1016/j.jmbbm.2015.05.023 . PMID   26144973.
  10. ↑ Vermeij GJ (27 September 2013). "The oyster enigma variations: a hypothesis of microbial calcification". Paleobiology. 40 (1): 1–13. doi:10.1666/13002. ISSN   0094-8373. S2CID   67846463.
  11. ↑ Bindschedler, Saskia; Cailleau, Guillaume; Verrecchia, Eric (5 May 2016). "Role of Fungi in the Biomineralization of Calcite". Minerals. 6 (2). MDPI AG: 41. Bibcode:2016Mine....6...41B. doi: 10.3390/min6020041 . ISSN   2075-163X.
  12. ↑ Görgen, Sigrid; Benzerara, Karim; Skouri-Panet, Fériel; Gugger, Muriel; Chauvat, Franck; Cassier-Chauvat, Corinne (30 November 2020). "The diversity of molecular mechanisms of carbonate biomineralization by bacteria". Discover Materials. 1 (1). Springer Science and Business Media LLC. doi: 10.1007/s43939-020-00001-9 . ISSN   2730-7727. S2CID   230631843.
  13. 1 2 3 4 Livingston BT, Killian CE, Wilt F, Cameron A, Landrum MJ, Ermolaeva O, et al. (December 2006). "A genome-wide analysis of biomineralization-related proteins in the sea urchin Strongylocentrotus purpuratus". Developmental Biology. 300 (1): 335–348. doi: 10.1016/j.ydbio.2006.07.047 . PMID   16987510.
  14. ↑ Checa AG, Ramírez-Rico J, González-Segura A, Sánchez-Navas A (January 2009). "Nacre and false nacre (foliated aragonite) in extant monoplacophorans (=Tryblidiida: Mollusca)". Die Naturwissenschaften. 96 (1): 111–122. Bibcode:2009NW.....96..111C. doi:10.1007/s00114-008-0461-1. PMID   18843476. S2CID   10214928.
  15. ↑ Currey JD (December 1999). "The design of mineralised hard tissues for their mechanical functions". The Journal of Experimental Biology. 202 (Pt 23): 3285–3294. doi:10.1242/jeb.202.23.3285. PMID   10562511.
  16. ↑ Cuif JP, Dauphin Y (2003). Les étapes de la découverte des rapports entre la terre et la vie : une introduction à la paléontologie. Paris: Éditions scientifiques GB. ISBN   978-2847030082. OCLC   77036366.
  17. 1 2 Borchers A, Pieler T (November 2010). "Programming pluripotent precursor cells derived from Xenopus embryos to generate specific tissues and organs". Genes. 1 (3). MDPI AG: 413–426. Bibcode:2016Mine....6...41B. doi: 10.3390/min6020041 . PMC   3966229 . PMID   24710095. CC-BY icon.svg Modified material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  18. ↑ Gadd GM (January 2007). "Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation". Mycological Research. 111 (Pt 1): 3–49. doi:10.1016/j.mycres.2006.12.001. PMID   17307120.
  19. 1 2 3 4 Li Q, Gadd GM (October 2017). "Biosynthesis of copper carbonate nanoparticles by ureolytic fungi". Applied Microbiology and Biotechnology. 101 (19): 7397–7407. doi:10.1007/s00253-017-8451-x. PMC   5594056 . PMID   28799032.
  20. 1 2 3 Liang X, Hillier S, Pendlowski H, Gray N, Ceci A, Gadd GM (June 2015). "Uranium phosphate biomineralization by fungi". Environmental Microbiology. 17 (6): 2064–2075. Bibcode:2015EnvMi..17.2064L. doi:10.1111/1462-2920.12771. PMID   25580878. S2CID   9699895.
  21. 1 2 3 4 Adeyemi AO, Gadd GM (June 2005). "Fungal degradation of calcium-, lead- and silicon-bearing minerals". Biometals. 18 (3): 269–281. doi:10.1007/s10534-005-1539-2. PMID   15984571. S2CID   35004304.
  22. ↑ Adamo P, Violante P (1 May 2000). "Weathering of rocks and neogenesis of minerals associated with lichen activity". Applied Clay Science. 16 (5): 229–256. Bibcode:2000ApCS...16..229A. doi:10.1016/S0169-1317(99)00056-3.
  23. ↑ Fortin D (March 2004). "Geochemistry. What biogenic minerals tell us". Science. 303 (5664): 1618–1619. doi:10.1126/science.1095177. PMID   15016984. S2CID   41179538.
  24. ↑ Gwenzi W (2019). "Carbon Sequestration via Biomineralization: Processes, Applications and Future Directions". Sustainable Agriculture Reviews 37. Vol. 37. Cham: Springer International Publishing. pp. 93–106. doi:10.1007/978-3-030-29298-0_5. ISBN   978-3-030-29297-3. ISSN   2210-4410. S2CID   214154330.
  25. 1 2 3 Knoll, A.H. (2004). "Biomineralization and evolutionary history" (PDF). In Dove PM, DeYoreo JJ, Weiner S (eds.). Reviews in Mineralogy and Geochemistry. Archived from the original (PDF) on 20 June 2010.
  26. ↑ Patek SN, Caldwell RL (October 2005). "Extreme impact and cavitation forces of a biological hammer: strike forces of the peacock mantis shrimp Odontodactylus scyllarus". The Journal of Experimental Biology. 208 (Pt 19): 3655–3664. doi: 10.1242/jeb.01831 . PMID   16169943. S2CID   312009.
  27. ↑ Demaster DJ (2001). "Marine Silica Cycle". Encyclopedia of Ocean Sciences. pp. 1659–1667. doi:10.1006/rwos.2001.0278. ISBN   9780122274305.
  28. ↑ 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". Chemistry of Materials. 27 (19): 6516–6523. arXiv: 1609.05449 . doi:10.1021/acs.chemmater.5b01542. ISSN   0897-4756. S2CID   118355403.
  29. ↑ Neues F, Hild S, Epple M, Marti O, Ziegler A (July 2011). "Amorphous and crystalline calcium carbonate distribution in the tergite cuticle of moulting Porcellio scaber (Isopoda, Crustacea)" (PDF). Journal of Structural Biology. 175 (1): 10–20. doi:10.1016/j.jsb.2011.03.019. PMID   21458575.
  30. ↑ Jacob DE, Wirth R, Agbaje OB, Branson O, Eggins SM (November 2017). "Planktic foraminifera form their shells via metastable carbonate phases". Nature Communications. 8 (1): 1265. Bibcode:2017NatCo...8.1265J. doi:10.1038/s41467-017-00955-0. PMC   5668319 . PMID   29097678.
  31. ↑ Mass T, Giuffre AJ, Sun CY, Stifler CA, Frazier MJ, Neder M, et al. (September 2017). "Amorphous calcium carbonate particles form coral skeletons". Proceedings of the National Academy of Sciences of the United States of America. 114 (37): E7670–E7678. Bibcode:2017PNAS..114E7670M. doi: 10.1073/pnas.1707890114 . PMC   5604026 . PMID   28847944.
  32. ↑ Raven JA, Giordano M (March 2009). "Biomineralization by photosynthetic organisms: evidence of coevolution of the organisms and their environment?". Geobiology. 7 (2): 140–154. Bibcode:2009Gbio....7..140R. doi:10.1111/j.1472-4669.2008.00181.x. PMID   19207569. S2CID   42962176.
  33. ↑ 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)". Cell and Tissue Research. 201 (3): 409–422. doi:10.1007/BF00236999. PMID   574424. S2CID   2222515.
  34. ↑ Habibah TU, Amlani DB, Brizuela M (January 2018). "Biomaterials, Hydroxyapatite". Stat Pearls. PMID   30020686. Archived from the original on 28 March 2020. Retrieved 12 August 2018.
  35. ↑ Abou Neel EA, Aljabo A, Strange A, Ibrahim S, Coathup M, Young AM, et al. (2016). "Demineralization-remineralization dynamics in teeth and bone". International Journal of Nanomedicine. 11: 4743–4763. doi: 10.2147/IJN.S107624 . PMC   5034904 . PMID   27695330.
  36. 1 2 Pepla E, Besharat LK, Palaia G, Tenore G, Migliau G (July 2014). "Nano-hydroxyapatite and its applications in preventive, restorative and regenerative dentistry: a review of literature". Annali di Stomatologia. 5 (3): 108–114. PMC   4252862 . PMID   25506416.
  37. ↑ Weaver JC, Milliron GW, Miserez A, Evans-Lutterodt K, Herrera S, Gallana I, et al. (June 2012). "The stomatopod dactyl club: a formidable damage-tolerant biological hammer". Science. 336 (6086): 1275–1280. Bibcode:2012Sci...336.1275W. doi:10.1126/science.1218764. PMID   22679090. S2CID   8509385. Archived from the original on 13 September 2020. Retrieved 2 December 2017.
  38. ↑ Tanner KE (June 2012). "Materials science. Small but extremely tough". Science. 336 (6086): 1237–1238. Bibcode:2012Sci...336.1237T. doi:10.1126/science.1222642. PMID   22679085. S2CID   206541609.
  39. ↑ Joester D, Brooker LR (5 July 2016). "The Chiton Radula: A Model System for Versatile Use of Iron Oxides*". In Faivre D (ed.). Iron Oxides (1st ed.). Wiley. pp. 177–206. doi:10.1002/9783527691395.ch8. ISBN   978-3-527-33882-5.
  40. ↑ Barber AH, Lu D, Pugno NM (April 2015). "Extreme strength observed in limpet teeth". Journal of the Royal Society, Interface. 12 (105): 20141326. doi:10.1098/rsif.2014.1326. PMC   4387522 . PMID   25694539.
  41. ↑ Chen C, Linse K, Copley JT, Rogers AD (August 2015). "The 'scaly-foot gastropod': a new genus and species of hydrothermal vent-endemic gastropod (Neomphalina: Peltospiridae) from the Indian Ocean". Journal of Molluscan Studies. 81 (3): 322–334. doi: 10.1093/mollus/eyv013 . ISSN   0260-1230.
  42. ↑ Pósfai M, Lefèvre CT, Trubitsyn D, Bazylinski DA, Frankel RB (November 2013). "Phylogenetic significance of composition and crystal morphology of magnetosome minerals". Frontiers in Microbiology. 4: 344. doi: 10.3389/fmicb.2013.00344 . PMC   3840360 . PMID   24324461.
  43. ↑ "Weird Sea Mollusk Sports Hundreds of Eyes Made of Armor". Live Science . 19 November 2015. Archived from the original on 17 August 2016. Retrieved 28 July 2016.
  44. 1 2 Le Moigne FA (2019). "Pathways of Organic Carbon Downward Transport by the Oceanic Biological Carbon Pump". Frontiers in Marine Science. 6. doi: 10.3389/fmars.2019.00634 . CC-BY icon.svg Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  45. ↑ Martin P, Allen JT, Cooper MJ, Johns DG, Lampitt RS, Sanders R, Teagle DA (2010). "Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies". Limnology and Oceanography. 55 (2): 604–614. doi: 10.4319/lo.2009.55.2.0604 .
  46. ↑ Belcher A, Manno C, Thorpe S, Tarling G (2018). "Acantharian cysts: High flux occurrence in the bathypelagic zone of the Scotia Sea, Southern Ocean" (PDF). Marine Biology. 165 (7): 117. Bibcode:2018MarBi.165..117B. doi:10.1007/s00227-018-3376-1. S2CID   90349921.
  47. 1 2 Raven JA, Knoll AH (2010). "Non-Skeletal Biomineralization by Eukaryotes: Matters of Moment and Gravity". Geomicrobiology Journal. 27 (6–7): 572–584. Bibcode:2010GmbJ...27..572R. doi:10.1080/01490451003702990. S2CID   37809270.
  48. ↑ Knoll AH (2003). "Biomineralization and Evolutionary History". Reviews in Mineralogy and Geochemistry. 54 (1): 329–356. Bibcode:2003RvMG...54..329K. doi:10.2113/0540329.
  49. ↑ Knoll AH, Kotrc B (2015). "Protistan Skeletons: A Geologic History of Evolution and Constraint". Evolution of Lightweight Structures. Biologically-Inspired Systems. Vol. 6. Dordrecht: Springer. pp. 1–16. doi:10.1007/978-94-017-9398-8_1. ISBN   978-94-017-9397-1. S2CID   83376982.
  50. 1 2 Marron AO, Ratcliffe S, Wheeler GL, Goldstein RE, King N, Not F, et al. (December 2016). "The Evolution of Silicon Transport in Eukaryotes". Molecular Biology and Evolution. 33 (12): 3226–3248. doi:10.1093/molbev/msw209. PMC   5100055 . PMID   27729397.
  51. ↑ 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". Journal of Structural Biology. 158 (3): 428–444. doi:10.1016/j.jsb.2007.01.001. PMID   17306563.
  52. ↑ Foissner W, Weissenbacher B, Krautgartner WD, Lütz-Meindl U (2009). "A cover of glass: first report of biomineralized silicon in a ciliate, Maryna umbrellata (Ciliophora: Colpodea)". The Journal of Eukaryotic Microbiology. 56 (6): 519–530. doi:10.1111/j.1550-7408.2009.00431.x. PMC   2917745 . PMID   19883440.
  53. 1 2 Preisig HR (1994). "Siliceous structures and silicification in flagellated protists". Protoplasma. 181 (1–4): 29–42. doi:10.1007/BF01666387. S2CID   27698051.
  54. ↑ 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". Experimental Cell Research. 58 (2): 334–342. doi:10.1016/0014-4827(69)90514-X. PMID   5404077.
  55. ↑ Martin-Jézéquel V, Hildebrand M, Brzezinski MA (2000). "Silicon Metabolism in Diatoms: Implications for Growth". Journal of Phycology. 36 (5): 821–840. Bibcode:2000JPcgy..36..821M. doi:10.1046/j.1529-8817.2000.00019.x. S2CID   84525482.
  56. 1 2 3 4 5 Hendry KR, Marron AO, Vincent F, Conley DJ, Gehlen M, Ibarbalz FM, Quéguiner B, Bowler C (2018). "Competition between Silicifiers and Non-silicifiers in the Past and Present Ocean and Its Evolutionary Impacts". Frontiers in Marine Science. 5. doi: 10.3389/fmars.2018.00022 . S2CID   12447257. CC-BY icon.svg Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  57. ↑ Mann S (2001). Biomineralization: Principles and Concepts in Bioinorganic Materials Chemistry. Oxford University Press. ISBN   9780198508823.
  58. ↑ Raven JA, Waite AM (2004). "The evolution of silicification in diatoms: Inescapable sinking and sinking as escape?". New Phytologist. 162 (1): 45–61. doi: 10.1111/j.1469-8137.2004.01022.x .
  59. ↑ Finkel ZV, Kotrc B (2010). "Silica Use Through Time: Macroevolutionary Change in the Morphology of the Diatom Fustule". Geomicrobiology Journal. 27 (6–7): 596–608. Bibcode:2010GmbJ...27..596F. doi:10.1080/01490451003702941. S2CID   85218013.
  60. ↑ Raven JA (1983). "The Transport and Function of Silicon in Plants". Biological Reviews. 58 (2): 179–207. doi:10.1111/j.1469-185X.1983.tb00385.x. S2CID   86067386.
  61. ↑ Hecky RE, Mopper K, Kilham P, Degens ET (1973). "The amino acid and sugar composition of diatom cell-walls". Marine Biology. 19 (4): 323–331. Bibcode:1973MarBi..19..323H. doi:10.1007/BF00348902. S2CID   84200496.
  62. ↑ Lobel KD, West JK, Hench LL (1996). "Computational model for protein-mediated biomineralization of the diatom frustule". Marine Biology. 126 (3): 353–360. Bibcode:1996MarBi.126..353L. doi:10.1007/BF00354617. S2CID   84969529.
  63. ↑ Hamm CE, Merkel R, Springer O, Jurkojc P, Maier C, Prechtel K, Smetacek V (February 2003). "Architecture and material properties of diatom shells provide effective mechanical protection" (PDF). Nature. 421 (6925): 841–843. Bibcode:2003Natur.421..841H. doi:10.1038/nature01416. PMID   12594512. S2CID   4336989.
  64. ↑ Aitken ZH, Luo S, Reynolds SN, Thaulow C, Greer JR (February 2016). "Microstructure provides insights into evolutionary design and resilience of Coscinodiscus sp. frustule". Proceedings of the National Academy of Sciences of the United States of America. 113 (8): 2017–2022. Bibcode:2016PNAS..113.2017A. doi: 10.1073/pnas.1519790113 . PMC   4776537 . PMID   26858446.
  65. ↑ Ehrlich H, Janussen D, Simon P, Bazhenov VV, Shapkin NP, Erler C, et al. (2008). "Nanostructural Organization of Naturally Occurring Composites—Part I: Silica-Collagen-Based Biocomposites". Journal of Nanomaterials. 2008: 1–8. doi: 10.1155/2008/623838 .
  66. ↑ Shimizu K, Amano T, Bari MR, Weaver JC, Arima J, Mori N (September 2015). "Glassin, a histidine-rich protein from the siliceous skeletal system of the marine sponge Euplectella, directs silica polycondensation". Proceedings of the National Academy of Sciences of the United States of America. 112 (37): 11449–11454. Bibcode:2015PNAS..11211449S. doi: 10.1073/pnas.1506968112 . PMC   4577155 . PMID   26261346.
  67. ↑ 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". Journal of Structural Biology. 158 (1): 93–106. doi:10.1016/j.jsb.2006.10.027. PMID   17175169.
  68. ↑ Nesbit KT, Roer RD (December 2016). "Silicification of the medial tooth in the blue crab Callinectes sapidus". Journal of Morphology. 277 (12): 1648–1660. doi:10.1002/jmor.20614. PMID   27650814. S2CID   46840652.
  69. ↑ 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". Protist. 158 (1): 21–28. doi:10.1016/j.protis.2006.09.002. PMID   17081802.
  70. ↑ Friedrichs L, Hörnig M, Schulze L, Bertram A, Jansen S, Hamm C (2013). "Size and biomechanic properties of diatom frustules influence food uptake by copepods". Marine Ecology Progress Series. 481: 41–51. Bibcode:2013MEPS..481...41F. doi: 10.3354/meps10227 .
  71. ↑ Hartley S, Degabriel JL (2016). "The ecology of herbivore-induced silicon defences in grasses". Functional Ecology. 30 (8): 1311–1322. Bibcode:2016FuEco..30.1311H. doi: 10.1111/1365-2435.12706 .
  72. ↑ Schaller J, Brackhage C, Bäucker E, Dudel EG (June 2013). "UV-screening of grasses by plant silica layer?". Journal of Biosciences. 38 (2): 413–416. doi:10.1007/s12038-013-9303-1. PMID   23660676. S2CID   16034220.
  73. ↑ Fuhrmann T, Landwehr S, Rharbi-Kucki E, Sumper M (2004). "Diatoms as living photonic crystals". Applied Physics B. 78 (3–4): 257–260. Bibcode:2004ApPhB..78..257F. doi:10.1007/s00340-004-1419-4. S2CID   121002890.
  74. ↑ 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". Journal of Applied Physics. 103 (7): 074701–074701–5. Bibcode:2008JAP...103g4701Y. doi:10.1063/1.2903342.
  75. ↑ Romann J, Valmalette JC, Chauton MS, Tranell G, Einarsrud MA, Vadstein O (December 2015). "Wavelength and orientation dependent capture of light by diatom frustule nanostructures". Scientific Reports. 5 (1): 17403. Bibcode:2015NatSR...517403R. doi:10.1038/srep17403. PMC   4667171 . PMID   26627680.
  76. ↑ Sundar VC, Yablon AD, Grazul JL, Ilan M, Aizenberg J (August 2003). "Fibre-optical features of a glass sponge". Nature. 424 (6951): 899–900. Bibcode:2003Natur.424..899S. doi:10.1038/424899a. PMID   12931176. S2CID   4426508.
  77. ↑ Dougherty LF, Johnsen S, Caldwell RL, Marshall NJ (September 2014). "A dynamic broadband reflector built from microscopic silica spheres in the 'disco' clam Ctenoides ales". Journal of the Royal Society, Interface. 11 (98): 20140407. doi:10.1098/rsif.2014.0407. PMC   4233689 . PMID   24966236.
  78. ↑ Desouky M, Jugdaohsingh R, McCrohan CR, White KN, Powell JJ (March 2002). "Aluminum-dependent regulation of intracellular silicon in the aquatic invertebrate Lymnaea stagnalis". Proceedings of the National Academy of Sciences of the United States of America. 99 (6): 3394–3399. Bibcode:2002PNAS...99.3394D. doi: 10.1073/pnas.062478699 . PMC   122534 . PMID   11891333.
  79. ↑ Neumann D, zur Nieden U (April 2001). "Silicon and heavy metal tolerance of higher plants". Phytochemistry. 56 (7): 685–692. Bibcode:2001PChem..56..685N. doi:10.1016/S0031-9422(00)00472-6. PMID   11314953.
  80. ↑ Milligan AJ, Morel FM (September 2002). "A proton buffering role for silica in diatoms". Science. 297 (5588): 1848–1850. Bibcode:2002Sci...297.1848M. doi:10.1126/science.1074958. PMID   12228711. S2CID   206507070.
  81. ↑ Adl SM, Simpson AG, Lane CE, Lukeš J, Bass D, Bowser SS, et al. (September 2012). "The revised classification of eukaryotes". The Journal of Eukaryotic Microbiology. 59 (5): 429–493. doi:10.1111/j.1550-7408.2012.00644.x. PMC   3483872 . PMID   23020233.
  82. ↑ Ensikat HJ, Geisler T, Weigend M (May 2016). "A first report of hydroxylated apatite as structural biomineral in Loasaceae - plants' teeth against herbivores". Scientific Reports. 6 (1): 26073. Bibcode:2016NatSR...626073E. doi:10.1038/srep26073. PMC   4872142 . PMID   27194462.
  83. ↑ 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". Chemistry: A European Journal. 18 (33): 10262–10270. doi:10.1002/chem.201201111. PMID   22696477.
  84. ↑ Weich RG, Lundberg P, Vogel HJ, Jensén P (May 1989). "Phosphorus-31 NMR Studies of Cell Wall-Associated Calcium-Phosphates in Ulva lactuca". Plant Physiology. 90 (1): 230–236. doi:10.1104/pp.90.1.230. PMC   1061703 . PMID   16666741.
  85. ↑ Tilman D (1977). "Resource Competition between Plankton Algae: An Experimental and Theoretical Approach". Ecology. 58 (2): 338–348. Bibcode:1977Ecol...58..338T. doi:10.2307/1935608. JSTOR   1935608.
  86. ↑ Sommer U (1994). "The impact of light intensity and daylength on silicate and nitrate competition among marine phytoplankton" (PDF). Limnology and Oceanography. 39 (7): 1680–1688. Bibcode:1994LimOc..39.1680S. doi:10.4319/lo.1994.39.7.1680.
  87. ↑ Porter S (2011). "The rise of predators". Geology. 39 (6): 607–608. Bibcode:2011Geo....39..607P. doi: 10.1130/focus062011.1 .
  88. ↑ Cohen PA, Schopf JW, Butterfield NJ, Kudryavtsev AB, Macdonald FA (2011). "Phosphate biomineralization in mid-Neoproterozoic protists". Geology. 39 (6): 539–542. Bibcode:2011Geo....39..539C. doi:10.1130/G31833.1. S2CID   32229787.
  89. ↑ 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". Nature Geoscience. 3 (9): 653–659. Bibcode:2010NatGe...3..653M. doi:10.1038/ngeo934. S2CID   13171894.
  90. ↑ Wood RA, Grotzinger JP, Dickson JA (June 2002). "Proterozoic modular biomineralized metazoan from the Nama Group, Namibia". Science. 296 (5577): 2383–2386. Bibcode:2002Sci...296.2383W. doi:10.1126/science.1071599. PMID   12089440. S2CID   9515357.
  91. 1 2 Zhuravlev AY, Wood RA (2008). "Eve of biomineralization: Controls on skeletal mineralogy" (PDF). Geology. 36 (12): 923. Bibcode:2008Geo....36..923Z. doi:10.1130/G25094A.1. Archived from the original (PDF) on 4 March 2016. Retrieved 28 August 2015.
  92. ↑ Porter SM (June 2007). "Seawater chemistry and early carbonate biomineralization". Science. 316 (5829): 1302. Bibcode:2007Sci...316.1302P. doi:10.1126/science.1137284. PMID   17540895. S2CID   27418253.
  93. ↑ 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". Geological Society of America Bulletin. 122 (11–12): 1731–1774. Bibcode:2010GSAB..122.1731M. doi:10.1130/B30346.1. S2CID   6694681.
  94. ↑ Murdock DJ, Donoghue PC (2011). "Evolutionary origins of animal skeletal biomineralization". Cells Tissues Organs. 194 (2–4): 98–102. doi:10.1159/000324245. PMID   21625061. S2CID   45466684.
  95. ↑ Kouchinsky A, Bengtson S, Runnegar B, Skovsted C, Steiner M, Vendrasco M (2011). "Chronology of early Cambrian biomineralization". Geological Magazine. 149 (2): 221–251. Bibcode:2012GeoM..149..221K. doi: 10.1017/S0016756811000720 .
  96. 1 2 Westbroek P, Marin F (April 1998). "A marriage of bone and nacre". Nature. 392 (6679): 861–862. Bibcode:1998Natur.392..861W. doi: 10.1038/31798 . PMID   9582064. S2CID   4348775.
  97. 1 2 Jackson DJ, McDougall C, Woodcroft B, Moase P, Rose RA, Kube M, et al. (March 2010). "Parallel evolution of nacre building gene sets in molluscs". Molecular Biology and Evolution. 27 (3): 591–608. doi: 10.1093/molbev/msp278 . PMID   19915030.
  98. ↑ Marin F, Smith M, Isa Y, Muyzer G, Westbroek P (February 1996). "Skeletal matrices, muci, and the origin of invertebrate calcification". Proceedings of the National Academy of Sciences of the United States of America. 93 (4): 1554–1559. Bibcode:1996PNAS...93.1554M. doi: 10.1073/pnas.93.4.1554 . PMC   39979 . PMID   11607630.
  99. ↑ Lowenstam HA, Margulis L (1980). "Evolutionary prerequisites for early Phanerozoic calcareous skeletons". Bio Systems. 12 (1–2): 27–41. Bibcode:1980BiSys..12...27L. doi:10.1016/0303-2647(80)90036-2. PMID   6991017.
  100. ↑ Lowenstam HA, Margulis L (1980). "Evolutionary prerequisites for early Phanerozoic calcareous skeletons". Bio Systems. 12 (1–2): 27–41. Bibcode:1980BiSys..12...27L. doi:10.1016/0303-2647(80)90036-2. PMID   6991017.
  101. ↑ Reyes-Bermudez A, Lin Z, Hayward DC, Miller DJ, Ball EE (July 2009). "Differential expression of three galaxin-related genes during settlement and metamorphosis in the scleractinian coral Acropora millepora". BMC Evolutionary Biology. 9 (1): 178. Bibcode:2009BMCEE...9..178R. doi: 10.1186/1471-2148-9-178 . PMC   2726143 . PMID   19638240.
  102. ↑ Jackson DJ, Macis L, Reitner J, Degnan BM, Wörheide G (June 2007). "Sponge paleogenomics reveals an ancient role for carbonic anhydrase in skeletogenesis". Science. 316 (5833): 1893–1895. Bibcode:2007Sci...316.1893J. doi: 10.1126/science.1141560 . PMID   17540861. S2CID   7042860.
  103. ↑ Wernström JV, Gąsiorowski L, Hejnol A (September 2022). "Brachiopod and mollusc biomineralisation is a conserved process that was lost in the phoronid-bryozoan stem lineage". EvoDevo. 13 (1): 17. doi: 10.1186/s13227-022-00202-8 . PMC   9484238 . PMID   36123753.
  104. ↑ Kirschvink JL, Hagadorn JW (2000). "10 A Grand Unified theory of Biomineralization.". In Bäuerlein E (ed.). The Biomineralisation of Nano- and Micro-Structures. Weinheim, Germany: Wiley-VCH. pp. 139–150.
  105. ↑ Towe KM, Lowenstam HA (January 1967). "Ultrastructure and development of iron mineralization in the radular teeth of Cryptochiton stelleri (Mollusca)". Journal of Ultrastructure Research. 17 (1): 1–13. doi:10.1016/S0022-5320(67)80015-7. PMID   6017357.
  106. ↑ Thomas GB, Komarneni S, Parker J (1993). Nanophase and Nanocomposite Materials: Symposium Held December 1–3, 1992, Boston, Massachusetts, U.S.A. (Materials Research Society Symposium Proceedings). Pittsburgh, Pa: Materials Research Society. ISBN   978-1-55899-181-1.
  107. 1 2 Sigel A, Sigel H, Sigel RK (30 April 2008). Biomineralization: From Nature to Application. John Wiley & Sons. ISBN   9780470986318.
  108. 1 2 Aparicio C, Ginebra MP (28 September 2015). Biomineralization and Biomaterials: Fundamentals and Applications. Woodhead. ISBN   9781782423560.
  109. ↑ Kolter R, Greenberg EP (May 2006). "Microbial sciences: the superficial life of microbes". Nature. 441 (7091): 300–302. Bibcode:2006Natur.441..300K. doi: 10.1038/441300a . PMID   16710410. S2CID   4430171.
  110. ↑ Palmer J, Flint S, Brooks J (September 2007). "Bacterial cell attachment, the beginning of a biofilm". Journal of Industrial Microbiology & Biotechnology. 34 (9): 577–588. doi: 10.1007/s10295-007-0234-4 . PMID   17619090. S2CID   978396.
  111. 1 2 Branda SS, Vik S, Friedman L, Kolter R (January 2005). "Biofilms: the matrix revisited". Trends in Microbiology. 13 (1): 20–26. doi:10.1016/j.tim.2004.11.006. PMID   15639628.
  112. ↑ Steinberg N, Kolodkin-Gal I (July 2015). "The Matrix Reloaded: Probing the Extracellular Matrix Synchronizes Bacterial Communities". Journal of Bacteriology. 197 (13): 2092–2103. doi:10.1128/JB.02516-14. PMC   4455261 . PMID   25825428.
  113. ↑ Dragoš A, Kovács ÁT (April 2017). "The Peculiar Functions of the Bacterial Extracellular Matrix". Trends in Microbiology. 25 (4): 257–266. doi:10.1016/j.tim.2016.12.010. PMID   28089324.
  114. 1 2 3 4 Dade-Robertson M, Keren-Paz A, Zhang M, Kolodkin-Gal I (September 2017). "Architects of nature: growing buildings with bacterial biofilms". Microbial Biotechnology. 10 (5): 1157–1163. doi:10.1111/1751-7915.12833. PMC   5609236 . PMID   28815998. CC-BY icon.svg Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  115. ↑ Oppenheimer-Shaanan Y, Sibony-Nevo O, Bloom-Ackermann Z, Suissa R, Steinberg N, Kartvelishvily E, et al. (2016). "Spatio-temporal assembly of functional mineral scaffolds within microbial biofilms". npj Biofilms and Microbiomes. 2: 15031. doi:10.1038/npjbiofilms.2015.31. PMC   5515261 . PMID   28721240. CC-BY icon.svg Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  116. ↑ Jonkers HM (2007). "Self healing concrete: a biological approach". In van der Zwaag S (ed.). Self Healing Materials: An Alternative Approach to 20 Centuries of Materials Science. Springer. pp. 195–204. ISBN   9781402062506.
  117. ↑ US 8728365,Dosier GK,"Methods for making construction material using enzyme producing bacteria",issued 2014, assigned to Biomason Inc.
  118. ↑ Rubinstein SM, Kolodkin-Gal I, McLoon A, Chai L, Kolter R, Losick R, Weitz DA (October 2012). "Osmotic pressure can regulate matrix gene expression in Bacillus subtilis". Molecular Microbiology. 86 (2): 426–436. doi:10.1111/j.1365-2958.2012.08201.x. PMC   3828655 . PMID   22882172.
  119. ↑ Chan JM, Guttenplan SB, Kearns DB (February 2014). "Defects in the flagellar motor increase synthesis of poly-γ-glutamate in Bacillus subtilis". Journal of Bacteriology. 196 (4): 740–753. doi:10.1128/JB.01217-13. PMC   3911173 . PMID   24296669.
  120. 1 2 Newsome L, Morris K, Lloyd JR (2014). "The biogeochemistry and bioremediation of uranium and other priority radionuclides". Chemical Geology. 363: 164–184. Bibcode:2014ChGeo.363..164N. doi: 10.1016/j.chemgeo.2013.10.034 .
  121. ↑ Lloyd JR, Macaskie LE (2000). Environmental microbe-metal interactions: Bioremediation of radionuclide-containing wastewaters. Washington, DC: ASM Press. pp. 277–327. ISBN   978-1-55581-195-2.
  122. ↑ Pasero M, et al. (November 2020). "The New IMA List of Minerals A Work in Progress" (PDF). The New IMA List of Minerals. IMA – CNMNC (Commission on New Minerals Nomenclature and Classification). Archived (PDF) from the original on 10 December 2020. Retrieved 11 December 2020.
  123. ↑ "IMA Database of Mineral Properties/ RRUFF Project". Department of Geosciences, University of Arizona. Retrieved 11 December 2020.
  124. 1 2 Lowenstam HA (March 1981). "Minerals formed by organisms". Science. 211 (4487): 1126–1131. Bibcode:1981Sci...211.1126L. doi:10.1126/science.7008198. JSTOR   1685216. PMID   7008198.
  125. 1 2 3 4 5 Skinner HC (2005). "Biominerals". Mineralogical Magazine. 69 (5): 621–41. Bibcode:2005MinM...69..621S. doi:10.1180/0026461056950275. S2CID   232388764.
  126. 1 2 Nickel EH (1995). "The definition of a mineral". The Canadian Mineralogist. 33 (3): 689–90.
  127. ↑ "Working Group on Environmental Mineralogy and Geochemistry". Commissions, working groups and committees. International Mineralogical Association. 3 August 2011. Retrieved 4 April 2018.
  128. ↑ 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.). Origins and Evolution of Life: An Astrobiological Perspective. Cambridge: Cambridge University Press. pp. 469–86. ISBN   978-1-139-49459-5.
  129. ↑ Roussel EG, Bonavita MA, Querellou J, Cragg BA, Webster G, Prieur D, Parkes RJ (May 2008). "Extending the sub-sea-floor biosphere". Science. 320 (5879): 1046. Bibcode:2008Sci...320.1046R. doi:10.1126/science.1154545. PMID   18497290. S2CID   23374807.
  130. ↑ Pearce DA, Bridge PD, Hughes KA, Sattler B, Psenner R, Russell NJ (August 2009). "Microorganisms in the atmosphere over Antarctica". FEMS Microbiology Ecology. 69 (2): 143–157. Bibcode:2009FEMME..69..143P. doi: 10.1111/j.1574-6941.2009.00706.x . PMID   19527292.
  131. ↑ Newman DK, Banfield JF (May 2002). "Geomicrobiology: how molecular-scale interactions underpin biogeochemical systems". Science. 296 (5570): 1071–1077. Bibcode:2002Sci...296.1071N. doi:10.1126/science.1010716. PMID   12004119. S2CID   1235688.
  132. ↑ Warren LA, Kauffman ME (February 2003). "Geoscience. Microbial geoengineers". Science. 299 (5609): 1027–1029. doi:10.1126/science.1072076. JSTOR   3833546. PMID   12586932. S2CID   19993145.
  133. ↑ 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". Geological Society, London, Special Publications. 336 (1): 31–50. Bibcode:2010GSLSP.336...31G. doi:10.1144/SP336.3. S2CID   130343033.
  134. ↑ Veis A (1990). "Biomineralization. Cell Biology and Mineral Deposition. by Kenneth Simkiss; Karl M. Wilbur On Biomineralization. by Heinz A. Lowenstam; Stephen Weiner". Science. 247 (4946): 1129–30. Bibcode:1990Sci...247.1129S. doi:10.1126/science.247.4946.1129. JSTOR   2874281. PMID   17800080.
  135. ↑ "Official IMA list of mineral names" (PDF). uws.edu.au. March 2009. Archived from the original (PDF) on 6 July 2011.
  136. ↑ Hefferan K, O'Brien J (2010). Earth Materials. Wiley-Blackwell. ISBN   978-1-4443-3460-9.
  137. ↑ Bindi L, Steinhardt PJ, Yao N, Lu PJ (2011). "Icosahedrite, Al63Cu24Fe13, the first natural quasicrystal". American Mineralogist. 96 (5–6): 928–31. Bibcode:2011AmMin..96..928B. doi:10.2138/am.2011.3758. S2CID   101152220.
  138. ↑ "Approved as new mineral" (PDF). Commission on New Minerals and Mineral Names. Archived from the original (PDF) on 20 March 2012.
  139. ↑ Corliss WR (November–December 1989). "Biogenic Minerals". Science Frontiers. 66.
  140. ↑ Steele A, Beaty D, eds. (26 September 2006). "Final report of the MEPAG Astrobiology Field Laboratory Science Steering Group (AFL-SSG)" (.doc). The Astrobiology Field Laboratory. U.S.A.: Mars Exploration Program Analysis Group (MEPAG) - NASA. p. 72. Retrieved 22 July 2009.
  141. 1 2 Grotzinger JP (January 2014). "Exploring martian habitability. Habitability, taphonomy, and the search for organic carbon on Mars. Introduction". Science. 343 (6169): 386–387. Bibcode:2014Sci...343..386G. doi: 10.1126/science.1249944 . PMID   24458635.
  142. 1 2 Various (24 January 2014). "Special Issue - Table of Contents - Exploring Martian Habitability". Science . 343: 345–452. Retrieved 24 January 2014.
  143. ↑ Various (24 January 2014). "Special Collection - Curiosity - Exploring Martian Habitability". Science . Retrieved 24 January 2014.
  144. ↑ 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". Science. 343 (6169): 1242777. Bibcode:2014Sci...343A.386G. CiteSeerX   10.1.1.455.3973 . doi:10.1126/science.1242777. PMID   24324272. S2CID   52836398.

Further reading

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.