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1

Becker, Wilhelm, Julia Marxen, Matthias Epple, and Oliver Reelsen. "Influence of microgravity on crystal formation in biomineralization." Journal of Applied Physiology 89, no. 4 (October 1, 2000): 1601–7. http://dx.doi.org/10.1152/jappl.2000.89.4.1601.

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Biomineralized tissues are widespread in animals. They are essential elements in skeletons and in statocysts. The function of both can only be understood with respect to gravitational force, which has always been present. Therefore, it is not astonishing to identify microgravity as a factor influencing biomineralization, normally resulting in the reduction of biomineralized materials. All known biominerals are composite materials, in which the organic matrix and the inorganic materials, organized in crystals, interact. If, during remodeling and turnover processes under microgravity, a defective organization of these crystals occurs, a reduction in biomineralized materials could be the result. To understand the influence of microgravity on the formation of biocrystals, we studied the shell-building process of the snail Biomphalaria glabrataas a model system. We show that, under microgravity (space shuttle flights STS-89 and STS-90), shell material is built in a regular way in both adult snails and snail embryos during the beginning of shell development. Microgravity does not influence crystal formation. Because gravity has constantly influenced evolution, the organization of biominerals with densities near 3 must have gained independence from gravitational forces, possibly early in evolution.
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Schmahl, Wolfgang, Erika Griesshaber, Lurdes Fernandez-Diaz, Andreas Ziegler, Klemens Kelm, Bernd Maier, Fitriana Nindiyasari, and Guntram Jordan. "Hierarchical structure of CaCO3biominerals – mesocrystals and functionalization." Acta Crystallographica Section A Foundations and Advances 70, a1 (August 5, 2014): C239. http://dx.doi.org/10.1107/s2053273314097605.

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Skeletal parts and teeth of marine organisms, avian eggshells, trilobite and isopod eyes, and many more biomineralized tissues consist of bio-calcite or bio-aragonite crystals. We explore the nano- to micro-scale architectures of these materials by electron backscatter diffraction (EBSD) and complementary techniques. In contrast to their inorganic cousins the biogenic "crystals" are hybrid composites with small amounts of organic matrix controlling morphogenesis and critically improving mechanical performance or other functions. For the biominerals meso-crystal-like structures are ubiquitous, consisting of co-oriented nano-blocks with a mosaic-spread of a few degrees, depending on the organism and on the size of the mesocrystal entity[1, 2, 3]. The nano-mosaic can be attributed to growth by nano-particle accretion from an amorphous or gel-like precursor, where relics of organic matrix cause misorientations between the crystallized nano-blocks. Recently we were able to reproduce this feature in gel-grown calcite [Nindiyasari et al., Crystal Growth and Design, in press]. The mesocrystal-co-orientation spreads on to the micro- and even millimeter-scale, frequently with a fractal nature of co-oriented hierarchical units [Maier et al., Acta Biomaterialia, accepted for publication]. The hierarchically structured morphology of the composite crystal or polycrystal is always directed by organic matrix membranes. Sea urchin teeth show a multiplex composite crystal architecture, where different subunits of engineered shapes, Mg-contents, and small misalignments are essential prerequisites for self-sharpening [1]. The figure shows an EBSD map of dendritic interdigitating calcite crystals in an avian egg shell (color coding for crystal orientation) with an misorientation profile along the grey line.
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3

Du, Tianming, Yumiao Niu, Youjun Liu, Haisheng Yang, Aike Qiao, and Xufeng Niu. "Physical and Chemical Characterization of Biomineralized Collagen with Different Microstructures." Journal of Functional Biomaterials 13, no. 2 (May 13, 2022): 57. http://dx.doi.org/10.3390/jfb13020057.

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Mineralized collagen is the basic unit in hierarchically organized natural bone with different structures. Polyacrylic acid (PAA) and periodic fluid shear stress (FSS) are the most common chemical and physical means to induce intrafibrillar mineralization. In the present study, non-mineralized collagen, extrafibrillar mineralized (EM) collagen, intrafibrillar mineralized (IM) collagen, and hierarchical intrafibrillar mineralized (HIM) collagen induced by PAA and FSS were prepared, respectively. The physical and chemical properties of these mineralized collagens with different microstructures were systematically investigated afterwards. Transmission electron microscopy (TEM) and scanning electron microscopy (SEM) showed that mineralized collagen with different microstructures was prepared successfully. The pore density of the mineralized collagen scaffold is higher under the action of periodic FSS. Fourier transform infrared spectroscopy (FTIR) analysis showed the formation of the hydroxyapatite (HA) crystal. A significant improvement in the pore density, hydrophilicity, enzymatic stability, and thermal stability of the mineralized collagen indicated that the IM collagen under the action of periodic FSS was beneficial for maintaining collagen activity. HIM collagen fibers, which are prepared under the co-action of periodic FSS and sodium tripolyphosphate (TPP), may pave the way for new bone substitute material applications.
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Schoeppler, Vanessa, Robert Lemanis, Elke Reich, Tamás Pusztai, László Gránásy, and Igor Zlotnikov. "Crystal growth kinetics as an architectural constraint on the evolution of molluscan shells." Proceedings of the National Academy of Sciences 116, no. 41 (September 24, 2019): 20388–97. http://dx.doi.org/10.1073/pnas.1907229116.

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Molluscan shells are a classic model system to study formation–structure–function relationships in biological materials and the process of biomineralized tissue morphogenesis. Typically, each shell consists of a number of highly mineralized ultrastructures, each characterized by a specific 3D mineral–organic architecture. Surprisingly, in some cases, despite the lack of a mutual biochemical toolkit for biomineralization or evidence of homology, shells from different independently evolved species contain similar ultrastructural motifs. In the present study, using a recently developed physical framework, which is based on an analogy to the process of directional solidification and simulated by phase-field modeling, we compare the process of ultrastructural morphogenesis of shells from 3 major molluscan classes: A bivalve Unio pictorum, a cephalopod Nautilus pompilius, and a gastropod Haliotis asinina. We demonstrate that the fabrication of these tissues is guided by the organisms by regulating the chemical and physical boundary conditions that control the growth kinetics of the mineral phase. This biomineralization concept is postulated to act as an architectural constraint on the evolution of molluscan shells by defining a morphospace of possible shell ultrastructures that is bounded by the thermodynamics and kinetics of crystal growth.
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5

Sugiura, Yuki, Kunio Ishikawa, Kazuo Onuma, and Yoji Makita. "PO4 adsorption on the calcite surface modulates calcite formation and crystal size." American Mineralogist 104, no. 10 (October 1, 2019): 1381–88. http://dx.doi.org/10.2138/am-2019-7015.

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Abstract Calcium carbonate (CaCO3) and particularly its stable phase, calcite, is of great geological significance in the deep carbon cycle since CaCO3 from biomineralized shells and corals form sedimentary rocks. Calcite also attracts attention in medical science and pharmacy as a primary or intermediate component in biomaterials because it possesses excellent biocompatibility along with suitable physicochemical properties. Calcite blocks have already been used during surgical procedures as a bone substitute for reconstructing bone defects formed by diseases and injury. When producing CaCO3 biomaterials and bioceramics, in particular, in vivo control of the size and polymorphic nature of CaCO3 is required. In this study, we investigated the effects of PO4 on calcite formation during the phase conversion of calcium sulfate anhydrate (CaSO4, CSA), which is sometimes used as a starting material for bone substitutes because of its suitable setting ability. CSA powder was immersed in 2 mol/L Na2CO3 solution containing a range of PO4 concentrations (0–60 mmol/L) at 40 °C for 3 days. The treated samples were investigated by X-ray diffraction, Fourier-transform infrared spectroscopy, X-ray fluorescence spectroscopy, and thermal analysis. In addition, the fine structures of the treated samples were observed by field-emission scanning electron microscopy, and the specific surface area was measured. We found that PO4, which is universally present in vivo, can modulate the calcite crystal size during calcite formation. A fluorescence study and calcite crystal growth experiments indicated that PO4 adsorbs tightly onto the surface of calcite, inhibiting crystal growth. In the presence of high PO4 concentrations, vaterite is formed along with calcite, and the appearance and stability of the CaCO3 polymorphs can be controlled by adjusting the PO4 concentration. These findings have implications for medical science and pharmacology, along with mineralogy and geochemistry.
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6

Rubini, Katia, Elisa Boanini, and Adriana Bigi. "Role of Aspartic and Polyaspartic Acid on the Synthesis and Hydrolysis of Brushite." Journal of Functional Biomaterials 10, no. 1 (February 1, 2019): 11. http://dx.doi.org/10.3390/jfb10010011.

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Dicalcium phosphate dihydrate (DCPD) is one of the mineral phases indicated as possible precursors of biological apatites and it is widely employed in the preparation of calcium phosphate bone cements. Herein, we investigated the possibility to functionalize DCPD with aspartic acid (ASP) and poly-aspartic acid (PASP), as models of the acidic macromolecules of biomineralized tissues, and studied their influence on DCPD hydrolysis. To this aim, the synthesis of DCPD was performed in aqueous solution in the presence of increasing concentrations of PASP and ASP, whereas the hydrolysis reaction was carried out in physiological solution up to three days. The results indicate that it is possible to prepare DCPD functionalized with PASP up to a polyelectrolyte content of about 2.3 wt%. The increase of PASP content induces crystal aggregation, reduction of the yield of the reaction and of the thermal stability of the synthesized DCPD. Moreover, DCPD samples functionalized with PASP display a slower hydrolysis than pure DCPD. On the other hand, in the explored range of concentrations (up to 10 mM) ASP is not incorporated into DCPD and does not influence its crystallization nor its hydrolysis. At variance, when present in the hydrolysis solution, ASP, and even more PASP, delays the conversion into the more stable phases, octacalcium phosphate and/or hydroxyapatite. The greater influence of PASP on the synthesis and hydrolysis of DCPD can be ascribed to the cooperative action of the carboxylate groups and to its good fit with DCPD structure.
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7

Xia, Zhonghui, Xin Zhang, Yujuan Zhou, Liping Yao, Zhen Zhang, Rongqing Zhang, and Xiaojun Liu. "The Matrix Protein Cysrichin, a Galaxin-like Protein from Hyriopsis cumingii, Induces Vaterite Formation In Vitro." Biology 12, no. 3 (March 15, 2023): 447. http://dx.doi.org/10.3390/biology12030447.

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In this study, we cloned a novel matrix protein, cysrichin, with 16.03% homology and a similar protein structure to the coral biomineralized protein galaxin. Tissue expression analysis showed that cysrichin was mainly expressed in mantle and gill tissues. In situ hybridization indicated that cysrichin mRNA was detected in the entire epithelium region of mantle tissue. RNAi analysis and shell notching experiment confirmed that cysrichin participates in the prismatic layer and nacreous layer formation of the shell. An in vitro crystallization experiment showed that the cysrichin protein induced lotus-shaped and round-shaped crystals, which were identified as vaterite crystals. These results may provide new clues for understanding the formation of vaterite in freshwater shellfish.
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8

Perrin, Christine. "Early diagenesis of carbonate biocrystals : isomineralogical changes in aragonite coral skeletons." Bulletin de la Société Géologique de France 175, no. 2 (March 1, 2004): 95–106. http://dx.doi.org/10.2113/175.2.95.

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Abstract Early diagenetic changes occurring in aragonite coral skeletons were characterized at the micro- and ultra-structural scales in living and fossil scleractinian colonies, the latter of Pleistocene age. The skeleton of scleractinian corals, like all biomineralized structures, is a composite material formed by the intimate association of inorganic aragonite crystallites and organic matrices. In addition to its organo-mineral duality, the scleractinian skeleton is formed by the three-dimensional arrangement of two clearly distinct basic structural features, the centers of calcification and the fibers. The latter are typically characterized by a transverse micron-scale zonation revealing their incremental growth process. The size, geometry and three-dimensional arrangement of calcification centers and fibers are taxon-specific. The earliest diagenetic modifications of these skeletons have been clearly recognized in the older parts of living colonies. The first steps of diagenesis therefore take place only a few years after the skeleton had been secreted by the living polyps, and in the same environmental conditions. Comparisons with the uppermost living parts of the coral colonies clearly show that these first diagenetic changes are driven by the biological ultrastructural characteristics of these skeletons and are conditioned by the presence of organic envelopes interbedded with and surrounding aragonite crystallites. These first diagenetic processes induce the development of thin fringes of fibrous aragonite cements growing syntaxially on the aragonitic coral fibers, an alteration of the incremental zonation of coral fibers and also preferential diagenetic changes in the calcification centers, including dissolution of their minute internal crystals. Diagenetic patterns observed in Pleistocene coral colonies typically involve the same processes already recognized in the older skeletal parts of living colonies, suggesting that diagenesis occurs through continuous processes instead of clearly differentiated stages. Selective dissolution affects calcification centers and some growth increments of coral fibers. Alteration of the initial transverse zonation of coral fibers also occur through the development of micro-inclusions clearly seen in ultra-thin sections. Although usually thicker than those observed in the ancient skeletal parts of living colonies, syntaxial aragonite cements commonly occur in these fossil skeletons. These cements are often associated with gradual textural modifications of the underlying coral fibers, in particular the loss of the transverse micron-scale zonation. This suggests that the coral skeleton forming the substratum of diagenetic cements is progressively recrystallized in secondary aragonite. This recrystallization of coral aragonite begins at the external margin of the skeleton, just below the diagenetic cements and gradually moves towards the internal skeletal parts. Recrystallization takes place through concomitant fine-scale dissolution-precipitation processes and occurs with textural changes but no mineralogical change. The process of recrystallization is likely initiated by a biological degradation of organic skeletal matrices and can be also driven by thermodynamical constraints involving the lowering of surface free energies resulting from changes in crystal size. Alteration of skeletal organic matrix, textural changes in coral biocrystals through recrystallization and precipitation of secondary diagenetic aragonite may bias the original geochemical characteristics of coral skeletons. Although more work is needed to establish the influence of these early diagenetic processes on the geochemical signatures, it is already well known that the breakdown of organic skeletal envelopes and early recrystallization of shallow-water carbonates alter the stable isotopic composition. The widespread use of coral skeletons as environmental and climatic proxies makes strongly necessary a better understanding of these early diagenetic mechanisms and a precise characterization of the fine-scale diagenetic patterns of specimens for the optimization of geochemical interpretations. In particular, it cannot be assumed that an entire aragonitic composition can guarantee that there is no or slight diagenetic alteration.
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9

Vargas, Gabriele, Jefferson Cypriano, Tarcisio Correa, Pedro Leão, Dennis Bazylinski, and Fernanda Abreu. "Applications of Magnetotactic Bacteria, Magnetosomes and Magnetosome Crystals in Biotechnology and Nanotechnology: Mini-Review." Molecules 23, no. 10 (September 24, 2018): 2438. http://dx.doi.org/10.3390/molecules23102438.

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Magnetotactic bacteria (MTB) biomineralize magnetosomes, which are defined as intracellular nanocrystals of the magnetic minerals magnetite (Fe3O4) or greigite (Fe3S4) enveloped by a phospholipid bilayer membrane. The synthesis of magnetosomes is controlled by a specific set of genes that encode proteins, some of which are exclusively found in the magnetosome membrane in the cell. Over the past several decades, interest in nanoscale technology (nanotechnology) and biotechnology has increased significantly due to the development and establishment of new commercial, medical and scientific processes and applications that utilize nanomaterials, some of which are biologically derived. One excellent example of a biological nanomaterial that is showing great promise for use in a large number of commercial and medical applications are bacterial magnetite magnetosomes. Unlike chemically-synthesized magnetite nanoparticles, magnetosome magnetite crystals are stable single-magnetic domains and are thus permanently magnetic at ambient temperature, are of high chemical purity, and display a narrow size range and consistent crystal morphology. These physical/chemical features are important in their use in biotechnological and other applications. Applications utilizing magnetite-producing MTB, magnetite magnetosomes and/or magnetosome magnetite crystals include and/or involve bioremediation, cell separation, DNA/antigen recovery or detection, drug delivery, enzyme immobilization, magnetic hyperthermia and contrast enhancement of magnetic resonance imaging. Metric analysis using Scopus and Web of Science databases from 2003 to 2018 showed that applied research involving magnetite from MTB in some form has been focused mainly in biomedical applications, particularly in magnetic hyperthermia and drug delivery.
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Lee, Shichoon, Seung Goo Lee, Myungsun Sim, Donghoon Kwak, Jong Hwan Park, and Kilwon Cho. "Control over the Vertical Growth of Single Calcitic Crystals in Biomineralized Structures." Crystal Growth & Design 11, no. 11 (November 2, 2011): 4920–26. http://dx.doi.org/10.1021/cg200773x.

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11

Monteil, Caroline L., Nicolas Menguy, Sandra Prévéral, Alan Warren, David Pignol, and Christopher T. Lefèvre. "Accumulation and Dissolution of Magnetite Crystals in a Magnetically Responsive Ciliate." Applied and Environmental Microbiology 84, no. 8 (February 9, 2018): e02865-17. http://dx.doi.org/10.1128/aem.02865-17.

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ABSTRACTMagnetotactic bacteria (MTB) represent a group of microorganisms that are widespread in aquatic habitats and thrive at oxic-anoxic interfaces. They are able to scavenge high concentrations of iron thanks to the biomineralization of magnetic crystals in their unique organelles, the so-called magnetosome chains. Although their biodiversity has been intensively studied, their ecology and impact on iron cycling remain largely unexplored. Predation by protozoa was suggested as one of the ecological processes that could be involved in the release of iron back into the ecosystem. Magnetic protozoa were previously observed in aquatic environments, but their diversity and the fate of particulate iron during grazing are poorly documented. In this study, we report the morphological and molecular characterizations of a magnetically responsive MTB-grazing protozoan able to ingest high quantities of MTB. This protozoan is tentatively identified asUronema marinum, a ciliate known to be a predator of bacteria. Using light and electron microscopy, we investigated in detail the vacuoles in which the lysis of phagocytized prokaryotes occurs. We carried out high-resolution observations of aligned magnetosome chains and ongoing dissolution of crystals. Particulate iron in the ciliate represented approximately 0.01% of its total volume. We show the ubiquity of this interaction in other types of environments and describe different grazing strategies. These data contribute to the mounting evidence that the interactions between MTB and protozoa might play a significant role in iron turnover in microaerophilic habitats.IMPORTANCEIdentifying participants of each biogeochemical cycle is a prerequisite to our understanding of ecosystem functioning. Magnetotactic bacteria (MTB) participate in iron cycling by concentrating large amounts of biomineralized iron minerals in their cells, which impacts their chemical environment at, or below, the oxic-anoxic transition zone in aquatic habitats. It was shown that some protozoa inhabiting this niche could become magnetic by the ingestion of magnetic crystals biomineralized by grazed MTB. In this study, we show that magnetic MTB grazers are commonly observed in marine and freshwater sediments and can sometimes accumulate very large amounts of particulate iron. We describe here different phagocytosis strategies, determined using magnetic particles from MTB as tracers after their ingestion by the protozoa. This study paves the way for potential scientific or medical applications using MTB grazers as magnetosome hyperaccumulators.
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Baaziz, Walid, Corneliu Ghica, Jefferson Cypriano, Fernanda Abreu, Karine Anselme, Ovidiu Ersen, Marcos Farina, and Jacques Werckmann. "New Phenotype and Mineralization of Biogenic Iron Oxide in Magnetotactic Bacteria." Nanomaterials 11, no. 12 (November 25, 2021): 3189. http://dx.doi.org/10.3390/nano11123189.

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Many magnetotactic bacteria (MTB) biomineralize magnetite crystals that nucleate and grow inside intracellular membranous vesicles originating from invaginations of the cytoplasmic membrane. The crystals together with their surrounding membranes are referred to as magnetosomes. Magnetosome magnetite crystals nucleate and grow using iron transported inside the vesicle by specific proteins. Here, we tackle the question of the organization of magnetosomes, which are always described as constituted by linear chains of nanocrystals. In addition, it is commonly accepted that the iron oxide nanocrystals are in the magnetite-based phase. We show, in the case of a wild species of coccus-type bacterium, that there is a double organization of the magnetosomes, relatively perpendicular to each other, and that the nanocrystals are in fact maghemite. These findings were obtained, respectively, by using electron tomography of whole mounts of cells directly from the environment and high-resolution transmission electron microscopy and diffraction. Structure simulations were performed with the MacTempas software. This study opens new perspectives on the diversity of phenotypes within MTBs and allows to envisage other mechanisms of nucleation and formation of biogenic iron oxide crystals.
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Ferreira Brasileiro, Pedro Pinto, Bruno Augusto Cabral Roque, Yana Batista Brandão, Alessandro Alberto Casazza, Attilio Converti, Mohand Benachour, and Leonie Asfora Sarubbo. "Cascade System for Biomineralization in Cement: Project, Construction and Operationalization to Enhance Building Energy Efficiency." Energies 15, no. 14 (July 20, 2022): 5262. http://dx.doi.org/10.3390/en15145262.

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Anthropogenic and natural actions cause internal and external fractures in concrete. To recover these structures, bio-concretes have been developed with bacteria of the genus Bacillus. These microorganisms consume calcium lactate, synthesize calcium carbonate and biomineralize CaCO3 crystals within the structures of concrete. The aim of the present study was to construct equipment, denominated “Cascade System for Biomineralization in Cement” (CSBC), to determine the limiting velocity of the biomineralization of CaCO3. The construction of the equipment took into consideration chemical and biochemical phenomena responsible for biomineralization. Parts made with 3D printing and a circuit with Arduino UNO R3 board were used in the assembly of the system. The prototype proved to be stable and can be considered a promising tool for future application in research of the regeneration of reinforced concreted in a practical, fast and economical way, especially to the energy sector.
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Kawamata, Seiichi. "Localization of pyroantimonate-precipitable calcium in biomineralization with special reference to the calcium carbonate crystals (otoconia)." Proceedings, annual meeting, Electron Microscopy Society of America 48, no. 3 (August 12, 1990): 562–63. http://dx.doi.org/10.1017/s0424820100160364.

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Biomineralized tissues (otoconia, bones and teeth) incorporate and/or liberate a lot of Ca ion. However, the direct visualization of ionic and loosely-bound Ca has not been successful because of the hardness of these tissues. This problem was partly overcome in the present study. The pyroantimonate technique and a new demineralization method were successfully combined to visualize pyroantimonateprecipi table Ca in the organs where CaCO3 crystals are formed.Tree frogs, with or without 0.8% CaCl2 loading, were decapitated. Their spinal columns containing the endolymphatic sac were sliced and immersed in a fixative that contained 2% glutaraldehyde and 2% K-pyroantimonate in 0.05M K-phosphate buffer (pH 7.4). Rats (2 and 8 weeks of age) anesthesized with ether were perfused via left ventricle first with saline, followed by a fixative containing 3% glutaraldehyde and 2% K-pyroantimonate in 0.1M K-phosphate buffer (pH 7.4). The otolithic organs were removed and immersed in the same fixative. Specimens of the tree frogs and rats were postfixed in 1% OsO4 and 2% K-pyroantimonate.
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Melchiorre, Erik B., Bryan H. Seymour, and Robert D. Evans. "The Interpretation of Biogeochemical Growths on Gold Coins from the SS Central America Shipwreck: Applications for Biogeochemistry and Geoarchaeology." Journal of Marine Science and Engineering 7, no. 7 (July 5, 2019): 209. http://dx.doi.org/10.3390/jmse7070209.

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Black crusts that formed on gold coins recovered from the 1857 shipwreck of the SS Central America played a key role in their preservation in a near original state. Within a few years of the sinking, the significant quantities of iron and steel on the shipwreck produced laminar geochemical precipitates of fine-grained iron minerals on the coins. This coating served to armor the coins from future chemical or biological attacks. Once coated, the coins were colonized by at least two distinct populations of gold-tolerant bacteria that precipitated abundant nanoparticulate gold in the black crust material and produced biomineralized bacteria in a web-like mat. Above this middle layer of black crust, the outer layer consisted of a geochemical reaction front of euhedral crystals of iron sulfate and iron oxy-hydroxide species, formed by the interaction of seawater with the chemical wastes of the bacterial mat. Understanding this process has application for assessing the diverse and extreme conditions under which nano-particulate gold may form through biological processes, as well as understanding the conditions that contribute to the preservation or degradation of marine archaeological materials.
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Uebe, René, Birgit Voigt, Thomas Schweder, Dirk Albrecht, Emanuel Katzmann, Claus Lang, Lars Böttger, Berthold Matzanke, and Dirk Schüler. "Deletion of a fur-Like Gene Affects Iron Homeostasis and Magnetosome Formation in Magnetospirillum gryphiswaldense." Journal of Bacteriology 192, no. 16 (June 18, 2010): 4192–204. http://dx.doi.org/10.1128/jb.00319-10.

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ABSTRACT Magnetotactic bacteria synthesize specific organelles, the magnetosomes, which are membrane-enveloped crystals of the magnetic mineral magnetite (Fe3O4). The biomineralization of magnetite involves the uptake and intracellular accumulation of large amounts of iron. However, it is not clear how iron uptake and biomineralization are regulated and balanced with the biochemical iron requirement and intracellular homeostasis. In this study, we identified and analyzed a homologue of the ferric uptake regulator Fur in Magnetospirillum gryphiswaldense, which was able to complement a fur mutant of Escherichia coli. A fur deletion mutant of M. gryphiswaldense biomineralized fewer and slightly smaller magnetite crystals than did the wild type. Although the total cellular iron accumulation of the mutant was decreased due to reduced magnetite biomineralization, it exhibited an increased level of free intracellular iron, which was bound mostly to a ferritin-like metabolite that was found significantly increased in Mössbauer spectra of the mutant. Compared to that of the wild type, growth of the fur mutant was impaired in the presence of paraquat and under aerobic conditions. Using a Fur titration assay and proteomic analysis, we identified constituents of the Fur regulon. Whereas the expression of most known magnetosome genes was unaffected in the fur mutant, we identified 14 proteins whose expression was altered between the mutant and the wild type, including five proteins whose genes constitute putative iron uptake systems. Our data demonstrate that Fur is a regulator involved in global iron homeostasis, which also affects magnetite biomineralization, probably by balancing the competing demands for biochemical iron supply and magnetite biomineralization.
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Wendt, Jobst. "A rare case of an evolutionary late and ephemeral biomineralization: tunicates with composite calcareous skeletons." Journal of Paleontology 94, no. 4 (January 24, 2020): 748–57. http://dx.doi.org/10.1017/jpa.2019.109.

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AbstractIn contrast to almost all other invertebrate phyla that constructed biomineralized skeletons during the “Cambrian explosion” and maintained them during the entire fossil record, ascidian tunicates evolved this protective and stabilizing advantage only during the Permian, although soft-bodied representatives of this subphylum made their first appearance already in the early Cambrian. It remains enigmatic why these compound calcareous skeletons persisted only until the Late Triassic, subsequently followed by less-rigid internal skeletons from the Lower Jurassic onwards, which consist of scattered isolated spicules only. In addition to recently described aragonitic ascidian exoskeletons from the Permian and Triassic, new discoveries of similar, but colonial ascidian compound endoskeletons in the lower Carnian exhibit a short-living branch of this group, which moreover contain the first indubitable calcareous spicules. The latter are embedded in the solid endoskeleton, which is composed of polygonal aragonitic plates with smooth outer and zigzag lined inner boundaries. They consist of irregular, parallel (orthogonal), or fan-shaped (clinogonal) arrangements of acicular aragonite crystals. The following taxa are described as new: order Cassianomorpha new order with the family Cassianosomidae new family and the genus Toscanisoma new genus with the species T. multipartitum new species and T. triplicatum new species.UUID: http://zoobank.org/03555353-cdab-42e8-8e99-9bfce15fa249
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Bazylinski, Dennis A., Timothy J. Williams, Christopher T. Lefèvre, Ryan J. Berg, Chuanlun L. Zhang, Samuel S. Bowser, Annette J. Dean, and Terrence J. Beveridge. "Magnetococcus marinus gen. nov., sp. nov., a marine, magnetotactic bacterium that represents a novel lineage (Magnetococcaceae fam. nov., Magnetococcales ord. nov.) at the base of the Alphaproteobacteria." International Journal of Systematic and Evolutionary Microbiology 63, Pt_3 (March 1, 2013): 801–8. http://dx.doi.org/10.1099/ijs.0.038927-0.

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Magnetotactic bacteria are a morphologically, metabolically and phylogenetically disparate array of bacteria united by the ability to biomineralize membrane-encased, single-magnetic-domain mineral crystals (magnetosomes) that cause the cell to orientate along the Earth’s geomagnetic field. The most commonly observed type of magnetotactic bacteria is the ubiquitous magnetotactic cocci, which comprise their own phylogenetic group. Strain MC-1T, a member of this group, was isolated from water collected from the oxic–anoxic interface of the Pettaquamscutt Estuary in Rhode Island, USA, and cultivated in axenic culture. Cells of strain MC-1T are roughly spherical, with two sheathed bundles of flagella at a single pole (bilophotrichous). Strain MC-1T uses polar magnetotaxis, and has a single chain of magnetite crystals per cell. Cells grow chemolithoautotrophically with thiosulfate or sulfide as the electron donors, and chemo-organoheterotrophically on acetate. During autotrophic growth, strain MC-1T relies on the reductive tricarboxylic acid cycle for CO2 fixation. The DNA G+C content is 54.2 mol%. The new genus and species Magnetococcus marinus gen. nov., sp. nov. are proposed to accommodate strain MC-1T ( = ATCC BAA-1437T = JCM 17883T), which is nominated as the type strain of Magnetococcus marinus. A new order (Magnetococcales ord. nov.) and family (Magnetococcaceae fam. nov.) are proposed for the reception of Magnetococcus and related magnetotactic cocci, which are provisionally included in the Alphaproteobacteria as the most basal known lineage of this class.
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de Oliveira, Jandira Ferreira, Eliane Wajnberg, Darci Motta de Souza Esquivel, Sevil Weinkauf, Michael Winklhofer, and Marianne Hanzlik. "Ant antennae: are they sites for magnetoreception?" Journal of The Royal Society Interface 7, no. 42 (May 27, 2009): 143–52. http://dx.doi.org/10.1098/rsif.2009.0102.

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Migration of the Pachycondyla marginata ant is significantly oriented at 13° with respect to the geomagnetic north–south axis. On the basis of previous magnetic measurements of individual parts of the body (antennae, head, thorax and abdomen), the antennae were suggested to host a magnetoreceptor. In order to identify Fe 3+ /Fe 2+ sites in antennae tissue, we used light microscopy on Prussian/Turnbull's blue-stained tissue. Further analysis using transmission electron microscopy imaging and diffraction, combined with elemental analysis, revealed the presence of ultra-fine-grained crystals (20–100 nm) of magnetite/maghaemite (Fe 3 O 4 /γ-Fe 2 O 3 ), haematite (α-Fe 2 O 3 ), goethite (α-FeOOH) besides (alumo)silicates and Fe/Ti/O compounds in different parts of the antennae, that is, in the joints between the third segment/pedicel, pedicel/scape and scape/head, respectively. The presence of (alumo)silicates and Fe/Ti/O compounds suggests that most, if not all, of the minerals in the tissue are incorporated soil particles rather than biomineralized by the ants. However, as the particles were observed within the tissue, they do not represent contamination. The amount of magnetic material associated with Johnston's organ and other joints appears to be sufficient to produce a magnetic-field-modulated mechanosensory output, which may therefore underlie the magnetic sense of the migratory ant.
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Li, Luoyang, Marissa J. Betts, Hao Yun, Bing Pan, Timothy P. Topper, Guoxiang Li, Xingliang Zhang, and Christian B. Skovsted. "Fibrous or Prismatic? A Comparison of the Lamello-Fibrillar Nacre in Early Cambrian and Modern Lophotrochozoans." Biology 12, no. 1 (January 11, 2023): 113. http://dx.doi.org/10.3390/biology12010113.

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The Precambrian–Cambrian interval saw the first appearance of disparate modern metazoan phyla equipped with a wide array of mineralized exo- and endo-skeletons. However, the current knowledge of this remarkable metazoan skeletonization bio-event and its environmental interactions is limited because uncertainties have persisted in determining the mineralogy, microstructure, and hierarchical complexity of these earliest animal skeletons. This study characterizes in detail a previously poorly understood fibrous microstructure—the lamello-fibrillar (LF) nacre—in early Cambrian mollusk and hyolith shells and compares it with shell microstructures in modern counterparts (coleoid cuttlebones and serpulid tubes). This comparative study highlights key differences in the LF nacre amongst different lophotrochozoan groups in terms of mineralogical compositions and architectural organization of crystals. The results demonstrate that the LF nacre is a microstructural motif confined to the Mollusca. This study demonstrates that similar fibrous microstructure in Cambrian mollusks and hyoliths actually represent a primitive type of prismatic microstructure constituted of calcitic prisms. Revision of these fibrous microstructures in Cambrian fossils demonstrates that calcitic shells are prevalent in the so-called aragonite sea of the earliest Cambrian. This has important implications for understanding the relationship between seawater chemistry and skeletal mineralogy at the time when skeletons were first acquired by early lophotrochozoan biomineralizers.
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Oestreicher, Zachery, Carmen Valverde-Tercedor, Eric Mumper, Lumarie Pérez-Guzmán, Nadia N. Casillas-Ituarte, Concepcion Jimenez-Lopez, Dennis A. Bazylinski, Steven K. Lower, and Brian H. Lower. "Localization of Native Mms13 to the Magnetosome Chain of Magnetospirillum magneticum AMB-1 Using Immunogold Electron Microscopy, Immunofluorescence Microscopy and Biochemical Analysis." Crystals 11, no. 8 (July 28, 2021): 874. http://dx.doi.org/10.3390/cryst11080874.

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Magnetotactic bacteria (MTB) biomineralize intracellular magnetite (Fe3O4) crystals surrounded by a magnetosome membrane (MM). The MM contains membrane-specific proteins that control Fe3O4 mineralization in MTB. Previous studies have demonstrated that Mms13 is a critical protein within the MM. Mms13 can be isolated from the MM fraction of Magnetospirillum magneticum AMB-1 and a Mms13 homolog, MamC, has been shown to control the size and shape of magnetite nanocrystals synthesized in-vitro. The objective of this study was to use several independent methods to definitively determine the localization of native Mms13 in M. magneticum AMB-1. Using Mms13-immunogold labeling and transmission electron microscopy (TEM), we found that Mms13 is localized to the magnetosome chain of M. magneticum AMB-1 cells. Mms13 was detected in direct contact with magnetite crystals or within the MM. Immunofluorescence detection of Mms13 in M. magneticum AMB-1 cells by confocal laser scanning microscopy (CLSM) showed Mms13 localization along the length of the magnetosome chain. Proteins contained within the MM were resolved by SDS-PAGE for Western blot analysis and LC-MS/MS (liquid chromatography with tandem mass spectrometry) protein sequencing. Using Anti-Mms13 antibody, a protein band with a molecular mass of ~14 kDa was detected in the MM fraction only. This polypeptide was digested with trypsin, sequenced by LC-MS/MS and identified as magnetosome protein Mms13. Peptides corresponding to the protein’s putative MM domain and catalytic domain were both identified by LC-MS/MS. Our results (Immunogold TEM, Immunofluorescence CLSM, Western blot, LC-MS/MS), combined with results from previous studies, demonstrate that Mms13 and homolog proteins MamC and Mam12, are localized to the magnetosome chain in MTB belonging to the class Alphaproteobacteria. Because of their shared localization in the MM and highly conserved amino acid sequences, it is likely that MamC, Mam12, and Mms13 share similar roles in the biomineralization of Fe3O4 nanocrystals.
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Casella, Laura A., Erika Griesshaber, Xiaofei Yin, Andreas Ziegler, Vasileios Mavromatis, Dirk Müller, Ann-Christine Ritter, et al. "Experimental diagenesis: insights into aragonite to calcite transformation of <i>Arctica islandica</i> shells by hydrothermal treatment." Biogeosciences 14, no. 6 (March 24, 2017): 1461–92. http://dx.doi.org/10.5194/bg-14-1461-2017.

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Abstract. Biomineralised hard parts form the most important physical fossil record of past environmental conditions. However, living organisms are not in thermodynamic equilibrium with their environment and create local chemical compartments within their bodies where physiologic processes such as biomineralisation take place. In generating their mineralised hard parts, most marine invertebrates produce metastable aragonite rather than the stable polymorph of CaCO3, calcite. After death of the organism the physiological conditions, which were present during biomineralisation, are not sustained any further and the system moves toward inorganic equilibrium with the surrounding inorganic geological system. Thus, during diagenesis the original biogenic structure of aragonitic tissue disappears and is replaced by inorganic structural features. In order to understand the diagenetic replacement of biogenic aragonite to non-biogenic calcite, we subjected Arctica islandica mollusc shells to hydrothermal alteration experiments. Experimental conditions were between 100 and 175 °C, with the main focus on 100 and 175 °C, reaction durations between 1 and 84 days, and alteration fluids simulating meteoric and burial waters, respectively. Detailed microstructural and geochemical data were collected for samples altered at 100 °C (and at 0.1 MPa pressure) for 28 days and for samples altered at 175 °C (and at 0.9 MPa pressure) for 7 and 84 days. During hydrothermal alteration at 100 °C for 28 days most but not the entire biopolymer matrix was destroyed, while shell aragonite and its characteristic microstructure was largely preserved. In all experiments up to 174 °C, there are no signs of a replacement reaction of shell aragonite to calcite in X-ray diffraction bulk analysis. At 175 °C the replacement reaction started after a dormant time of 4 days, and the original shell microstructure was almost completely overprinted by the aragonite to calcite replacement reaction after 10 days. Newly formed calcite nucleated at locations which were in contact with the fluid, at the shell surface, in the open pore system, and along growth lines. In the experiments with fluids simulating meteoric water, calcite crystals reached sizes up to 200 µm, while in the experiments with Mg-containing fluids the calcite crystals reached sizes up to 1 mm after 7 days of alteration. Aragonite is metastable at all applied conditions. Only a small bulk thermodynamic driving force exists for the transition to calcite. We attribute the sluggish replacement reaction to the inhibition of calcite nucleation in the temperature window from ca. 50 to ca. 170 °C or, additionally, to the presence of magnesium. Correspondingly, in Mg2+-bearing solutions the newly formed calcite crystals are larger than in Mg2+-free solutions. Overall, the aragonite–calcite transition occurs via an interface-coupled dissolution–reprecipitation mechanism, which preserves morphologies down to the sub-micrometre scale and induces porosity in the newly formed phase. The absence of aragonite replacement by calcite at temperatures lower than 175 °C contributes to explaining why aragonitic or bimineralic shells and skeletons have a good potential of preservation and a complete fossil record.
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Zhan, Yi, Bing Deng, Huixian Wu, Changpeng Xu, Ruiying Wang, Wenqiang Li, and Zhixiong Pan. "Biomineralized Composite Liquid Crystal Fiber Scaffold Promotes Bone Regeneration by Enhancement of Osteogenesis and Angiogenesis." Frontiers in Pharmacology 12 (November 8, 2021). http://dx.doi.org/10.3389/fphar.2021.736301.

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Liquid crystals (LCs) are appealing biomaterials for applications in bone regenerative medicine due to their tunable physical properties and anisotropic viscoelastic behavior. This study reports a novel composite poly (L-lactide) (PLLA) scaffold that is manufactured by a simple electrospinning and biomineralization technique that precisely controls the fibrous structure in liquid LC phase. The enriched-LC composites have superior mineralization ability than neat PLLA; furthermore BMSC cells were inoculated onto the HAP-PLLA/LC with hydroxyapatite (HAP) composite scaffold to test the capability for osteogenesis in vitro. The results show that the PLLA/LC with HAP produced by mineralization leads to better cell compatibility, which is beneficial to cell proliferation, osteogenic differentiation, and expression of the angiogenic CD31 gene. Moreover, in vivo studies showed that the HAP-PLLA/LC scaffold with a bone-like environment significantly accelerates new and mature lamellar bone formation by development of a microenvironment for vascularized bone regeneration. Thus, this bionic composite scaffold in an LC state combining osteogenesis with vascularized activities is a promising biomaterial for bone regeneration in defective areas.
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24

Yun, Hao, Xingliang Zhang, Glenn A. Brock, Luoyang Li, and Guoxiang Li. "Biomineralization of the Cambrian chancelloriids." Geology, February 5, 2021. http://dx.doi.org/10.1130/g48428.1.

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As extinct animals that flourished during the Cambrian explosion, chancelloriids have a unique body plan lacking guts but with a flexible integument and a suite of star-shaped, hollow sclerites. Due to this body plan, along with the paucity of knowledge on sclerite biomineralization, the phylogenetic position of chancelloriids within the Metazoa is still controversial. Integration of analyses of diverse fossils from Cambrian stage 2 to the Wuliuan Stage of China and Australia indicates that chancelloriid sclerites possess an encasement-like organic layer and a fibrous aragonitic layer. The organic layer is inferred to be a specialized trait derived from the epidermal integument of the animal body. The sclerites were likely biomineralized by using the outer organic layer as a template to absorb cations and precipitate crystal nuclei, reflecting a strategy adopted by a range of eumetazoans with a developed epidermis. Therefore, the hypothesis that chancelloriids represent an epitheliozoan-grade animal and an early explorer of template-based biomineralization is supported.
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Paramasivan, Mareeswari, T. S. Sampath Kumar, Hemalatha Kanniyappan, Vignesh Muthuvijayan, and T. S. Chandra. "Biomimetic ion substituted and Co-substituted hydroxyapatite nanoparticle synthesis using Serratia Marcescens." Scientific Reports 13, no. 1 (March 18, 2023). http://dx.doi.org/10.1038/s41598-023-30996-z.

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AbstractBiomimicry is becoming deep-rooted as part of bioceramics owing to its numerous functional advantages. Naturally occurring hydroxyapatite (HA) apart from primary nano structures are also characterised by various ionic substitutions. The ease of accommodating such key elements into the HA lattice is known to enhance bone healing properties of bioceramics. In this work, hydroxyapatite synthesized via biomimetic approach was substituted with individual as well as multiple cations for potential applications in bone repair. Ion substitutions of Sr, Mg and Zn was carried out on HA for the first time by using Serratia grown in a defined biomineralization medium. The individual ions of varying concentration substituted in Serratia HA (SHA) (Sr SHA, Mg SHA and Zn SHA) were analysed for crystallinity, functional groups, morphology and crystal size. All three showed decreased crystallinity, phase purity, large agglomerated aggregates and needle-shaped morphologies. Fourier transform infrared spectroscopy (FTIR) spectra indicated increased carbonate content of 5.8% resembling that of natural bone. Additionally, the reduced O–H intensities clearly portrayed disruption of HA lattice and subsequent ion-substitution. The novelty of this study lies primarily in investigating the co-substitution of a combination of 1% Sr, Zn and Mg in SHA and establishing the associated change in bone parameters. Scanning electron microscope (SEM) and transmission electron microscope (TEM) images clearly illustrated uniform nano-sized agglomerates of average dimensions of 20–50 nm length and 8–15 nm width for Sr SHA; 10–40 nm length and 8–10 nm width for both Zn SHA and Mg SHA and 40–70 nm length and 4–10 nm width in the case of 1% Sr, Zn, Mg SHA. In both individual as well as co-substitutions, significant peak shifts were not observed possibly due to the lower concentrations. However, cell volumes increased in both cases due to presence of Sr2+ validating its dominant integration into the SHA lattice. Rich trace ion deposition was presented by energy dispersive X-ray spectroscopy (EDS) and quantified using inductively coupled plasma optical emission spectrometer (ICP-OES). In vitro cytotoxicity studies in three cell lines viz. NIH/3T3 fibroblast cells, MG-63 osteosarcoma cells and RAW 264.7 macrophages showed more than 90% cell viability proving the biocompatible nature of 1% Sr, Zn and Mg in SHA. Microbial biomineralization by Serratia produced nanocrystals of HA that mimicked “bone-like apatite” as evidenced by pure phase, carbonated groups, reduced crystallinity, nano agglomerates, variations in cell parameters, rich ion deposition and non-toxic nature. Therefore ion-substituted and co-substituted biomineralized nano SHA appears to be a suitable candidate for applications in biomedicine addressing bone injuries and aiding regeneration as a result of its characteristics close to that of the human bone.
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Mohapatra, Adityanarayan, Ayeskanta Mohanty, Sathiyamoorthy Padmanaban, Sahil Chahal, Veena Vijayan, Santhosh Kalash Rajendrakumar, and In-Kyu Park. "Targeted treatment of gouty arthritis by biomineralized metallic nanozymes mediated oxidative stress mitigating nanotherapy." Journal of Materials Chemistry B, 2023. http://dx.doi.org/10.1039/d3tb00669g.

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Gouty arthritis is characterized by chronic deposition of monosodium urate (MSU) crystals in the joints and other tissues, resulting in the production of excess reactive oxygen species (ROS) and proinflammatory...
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27

Gómez-Martínez, Octavio, Daniel H. Aguilar, Juan J. Alvarado-Gil, Patricia Quintana, and Dalila Aldana-Aranda. "Texturization Analysis by X-ray Diffraction of Shells of the Mussel Ischadium recurvum (Rafinesque, 1820) (Mollusca Bivalvia)." MRS Proceedings 711 (2001). http://dx.doi.org/10.1557/proc-711-hh3.45.1.

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ABSTRACTMost of the inorganic biomineralized materials are deposited on an organic matrix that controls the orientation and structure of the crystals. It is thought that chemical groups at the surface of the matrix may act as a template for the nucleation and growth of the mineral. A x-ray diffraction study of the texturization development of the bivalve mollusk shells is presented; specifically, the mussel Ischadium recurvum (Rafinesque, 1820), in different growing stages. The x-ray reflections show a preferred orientation that changes as the mollusk grows, and at the final stages only two crystallographic planes prevail.
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Awal, Ram Prasad, Christopher T. Lefevre, and Dirk Schüler. "Functional expression of foreign magnetosome genes in the alphaproteobacterium Magnetospirillum gryphiswaldense." mBio, June 15, 2023. http://dx.doi.org/10.1128/mbio.03282-22.

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ABSTRACT Magnetosomes of magnetotactic bacteria (MTB) consist of structurally perfect, nano-sized magnetic crystals enclosed within vesicles of a proteo-lipid membrane. In species of Magnetospirillum, biosynthesis of their cubo-octahedral-shaped magnetosomes was recently demonstrated to be a complex process, governed by about 30 specific genes that are comprised within compact magnetosome gene clusters (MGCs). Similar, yet distinct gene clusters were also identified in diverse MTB that biomineralize magnetosome crystals with different, genetically encoded morphologies. However, since most representatives of these groups are inaccessible by genetic and biochemical approaches, their analysis will require the functional expression of magnetosome genes in foreign hosts. Here, we studied whether conserved essential magnetosome genes from closely and remotely related MTB can be functionally expressed by rescue of their respective mutants in the tractable model Magnetospirillum gryphiswaldense of the Alphaproteobacteria . Upon chromosomal integration, single orthologues from other magnetotactic Alphaproteobacteria restored magnetosome biosynthesis to different degrees, while orthologues from distantly related Magnetococcia and Deltaproteobacteria were found to be expressed but failed to re-induce magnetosome biosynthesis, possibly due to poor interaction with their cognate partners within multiprotein magnetosome organelle of the host. Indeed, co-expression of the known interactors MamB and MamM from the alphaproteobacterium Magnetovibrio blakemorei increased functional complementation. Furthermore, a compact and portable version of the entire MGCs of M. magneticum was assembled by transformation-associated recombination cloning, and it restored the ability to biomineralize magnetite both in deletion mutants of the native donor and M. gryphiswaldense , while co-expression of gene clusters from both M. gryphiswaldense and M. magneticum resulted in overproduction of magnetosomes. IMPORTANCE We provide proof of principle that Magnetospirillum gryphiswaldense is a suitable surrogate host for the functional expression of foreign magnetosome genes and extended the transformation-associated recombination cloning platform for the assembly of entire large magnetosome gene cluster, which could then be transplanted to different magnetotactic bacteria. The reconstruction, transfer, and analysis of gene sets or entire magnetosome clusters will be also promising for engineering the biomineralization of magnetite crystals with different morphologies that would be valuable for biotechnical applications.
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Liu, Peiyu, Yan Liu, Xinyi Ren, Zhifei Zhang, Xiang Zhao, Andrew P. Roberts, Yongxin Pan, and Jinhua Li. "A novel magnetotactic Alphaproteobacterium producing intracellular magnetite and calcium-bearing minerals." Applied and Environmental Microbiology, September 22, 2021. http://dx.doi.org/10.1128/aem.01556-21.

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Magnetotactic bacteria (MTB) are prokaryotes that form intracellular magnetite (Fe 3 O 4 ) or greigite (Fe 3 S 4 ) nanocrystals with tailored sizes, often in chain configurations. Such magnetic particles are each surrounded by a lipid bilayer membrane, called a magnetosome, and provide a model system for studying the formation and function of specialized internal structures in prokaryotes. Using fluorescence-coupled scanning electron microscopy, we identified a novel magnetotactic spirillum, XQGS-1, from freshwater Xingqinggong Lake, Xi’an City, Shaanxi Province, China. Phylogenetic analyses based on 16S rRNA gene sequences indicate that strain XQGS-1 represents a novel genus of the Alphaproteobacteria class in the Proteobacteria phylum. Transmission electron microscope analyses reveal that strain XQGS-1 forms on average 17 ± 3 magnetite magnetosome particles with ideal truncated octahedral morphology with average length and width of 88.3 ± 11.7 nm and 83.3 ± 11.0 nm, respectively. They are tightly organized into a single chain along the cell long axis close to the concave side of the cell. Intra-chain magnetic interactions likely result in these large equidimensional magnetite crystals behaving as magnetically stable single domain particles that enable bacterial magnetotaxis. Combined structural and chemical analyses demonstrate that XQGS-1 cells also biomineralize intracellular amorphous calcium phosphate (2-3 granules per cell; 90.5 ± 19.3 nm average size) and weakly crystalline calcium carbonate (2-3 granules per cell; 100.4 ± 21.4 nm average size) in addition to magnetite. Our results expand the taxonomic diversity of MTB and provide evidence for intracellular calcium phosphate biomineralization in MTB. IMPORTANCE Biomineralization is a widespread process in eukaryotes that form shells, teeth, or bones. It also occurs commonly in prokaryotes, resulting in more than sixty known minerals formed by different bacteria under wide-ranging conditions. Among them, magnetotactic bacteria (MTB) are remarkable because they might represent the earliest organisms that biomineralize intracellular magnetic iron minerals (i.e., magnetite (Fe 3 O 4 ) or greigite (Fe 3 S 4 )). Here we report a novel magnetotactic spirillum (XQGS-1) that is phylogenetically affiliated with the Alphaproteobacteria class. In addition to magnetite crystals, XQGS-1 cells form intracellular sub-micron calcium carbonate and calcium phosphate granules. This finding supports the view that MTB are also an important microbial group for intracellular calcium carbonate and calcium phosphate biomineralization.
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30

Pellegrino, Luca, Marcello Natalicchio, Kenta Abe, Richard W. Jordan, Sergio E. Favero Longo, Simona Ferrando, Giorgio Carnevale, and Francesco Dela Pierre. "Tiny, glassy, and rapidly trapped: The nano-sized planktic diatoms in Messinian (late Miocene) gypsum." Geology, July 20, 2021. http://dx.doi.org/10.1130/g49342.1.

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Primary gypsum represents an excellent paleobiological archive due to its early and fast growth, favoring the preservation of delicate biomineralized structures. The Mediterranean region is renowned for evaporite deposits that formed during the Messinian salinity crisis (MSC), an event that supposedly annihilated most of the marine biota. However, the Messinian evaporites have been scarcely studied for their fossil content. Abundant nano-sized planktic diatoms and associated organic matter are observed for the first time in bottom-grown gypsum crystals that formed during the early stage of the MSC in different marginal basins of the western Mediterranean. This discovery increases our knowledge of the Messinian biota and reveals that nano-sized planktic diatoms played a prominent role in carbon and silicon export during gypsum deposition. The co-occurrence of these diatoms with larger diatoms, possibly associated with a deep chlorophyll maximum, suggests that Messinian gypsum formed in stratified and relatively deep basins (far below the photic zone), typified by marine conditions in the upper water column. The nano-sized planktic diatoms may have taken advantage of the hydrological reconfigurations experienced by the Mediterranean since the onset of the MSC. This study confirms that primary gypsum represents a promising archive of information for elucidating the marine biotic response to an ancient environmental crisis.
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Zhang, Wensi, Yinzhao Wang, Li Liu, Yongxin Pan, and Wei Lin. "Identification and Genomic Characterization of Two Previously Unknown Magnetotactic Nitrospirae." Frontiers in Microbiology 12 (July 27, 2021). http://dx.doi.org/10.3389/fmicb.2021.690052.

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Magnetotactic bacteria (MTB) are a group of microbes that biomineralize membrane-bound, nanosized magnetite (Fe3O4), and/or greigite (Fe3S4) crystals in intracellular magnetic organelle magnetosomes. MTB belonging to the Nitrospirae phylum can form up to several hundreds of Fe3O4 magnetosome crystals and dozens of sulfur globules in a single cell. These MTB are widespread in aquatic environments and sometimes account for a significant proportion of microbial biomass near the oxycline, linking these lineages to the key steps of global iron and sulfur cycling. Despite their ecological and biogeochemical importance, our understanding of the diversity and ecophysiology of magnetotactic Nitrospirae is still very limited because this group of MTB remains unculturable. Here, we identify and characterize two previously unknown MTB populations within the Nitrospirae phylum through a combination of 16S rRNA gene-based and genome-resolved metagenomic analyses. These two MTB populations represent distinct morphotypes (rod-shaped and coccoid, designated as XYR, and XYC, respectively), and both form more than 100 bullet-shaped magnetosomal crystals per cell. High-quality draft genomes of XYR and XYC have been reconstructed, and they represent a novel species and a novel genus, respectively, according to their average amino-acid identity values with respect to available genomes. Accordingly, the names Candidatus Magnetobacterium cryptolimnobacter and Candidatus Magnetomicrobium cryptolimnococcus for XYR and XYC, respectively, were proposed. Further comparative genomic analyses of XYR, XYC, and previously reported magnetotactic Nitrospirae reveal the general metabolic potential of this MTB group in distinct microenvironments, including CO2 fixation, dissimilatory sulfate reduction, sulfide oxidation, nitrogen fixation, or denitrification processes. A remarkably conserved magnetosome gene cluster has been identified across Nitrospirae MTB genomes, indicating its putative important adaptive roles in these bacteria. Taken together, the present study provides novel insights into the phylogenomic diversity and ecophysiology of this intriguing, yet poorly understood MTB group.
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Li, Jinhua, Heng Zhang, Peiyu Liu, Nicolas Menguy, Andrew P. Roberts, Haitao Chen, Yinzhao Wang, and Yongxin Pan. "Phylogenetic and Structural Identification of a Novel Magnetotactic Deltaproteobacteria Strain, WYHR-1, from a Freshwater Lake." Applied and Environmental Microbiology 85, no. 14 (May 3, 2019). http://dx.doi.org/10.1128/aem.00731-19.

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ABSTRACT Magnetotactic bacteria (MTB) are phylogenetically diverse prokaryotes that are able to biomineralize intracellular, magnetic chains of magnetite or greigite nanocrystals called magnetosomes. Simultaneous characterization of MTB phylogeny and biomineralization is crucial but challenging because most MTB are extremely difficult to culture. We identify a large rod, bean-like MTB (tentatively named WYHR-1) from freshwater sediments of Weiyang Lake, Xi’an, China, using a coupled fluorescence and scanning electron microscopy approach at the single-cell scale. Phylogenetic analysis of 16S rRNA gene sequences indicates that WYHR-1 is a novel genus from the Deltaproteobacteria class. Transmission electron microscope observations reveal that WYHR-1 cells contain tens of magnetite magnetosomes that are organized into a single chain bundle along the cell long axis. Mature WYHR-1 magnetosomes are bullet-shaped, straight, and elongated along the [001] direction, with a large flat end terminated by a {100} face at the base and a conical top. This crystal morphology is distinctively different from bullet-shaped magnetosomes produced by other MTB in the Deltaproteobacteria class and the Nitrospirae phylum. This indicates that WYHR-1 may have a different crystal growth process and mechanism from other species, which results from species-specific magnetosome biomineralization in MTB. IMPORTANCE Magnetotactic bacteria (MTB) represent a model system for understanding biomineralization and are also studied intensively in biogeomagnetic and paleomagnetic research. However, many uncultured MTB strains have not been identified phylogenetically or investigated structurally at the single-cell level, which limits comprehensive understanding of MTB diversity and their role in biomineralization. We have identified a novel MTB strain, WYHR-1, from a freshwater lake using a coupled fluorescence and scanning electron microscopy approach at the single-cell scale. Our analyses further indicate that strain WYHR-1 represents a novel genus from the Deltaproteobacteria class. In contrast to bullet-shaped magnetosomes produced by other MTB in the Deltaproteobacteria class and the Nitrospirae phylum, WYHR-1 magnetosomes are bullet-shaped, straight, and highly elongated along the [001] direction, are terminated by a large {100} face at their base, and have a conical top. Our findings imply that, consistent with phylogenetic diversity of MTB, bullet-shaped magnetosomes have diverse crystal habits and growth patterns.
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33

Wittig, Nina K., and Henrik Birkedal. "Bone hierarchical structure: spatial variation across length scales." Acta Crystallographica Section B Structural Science, Crystal Engineering and Materials 78, no. 3 (March 31, 2022). http://dx.doi.org/10.1107/s2052520622001524.

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Bone is a complex hierarchical biomineralized material, which is special amongst biominerals because it is replete with cells, namely, osteocytes. While bone has been scrutinized for centuries, many questions remain open and new research hints that the ultrastructure of bone, encompassing both the bone matrix itself and the embedded cell network, is much more heterogeneous than hitherto realized. A number of these new findings have been made thanks to the enormous developments in X-ray imaging that have occurred in recent decades, and there is promise that they will also allow many of the remaining open questions to be addressed. X-ray absorption or phase imaging affords high three-dimensional (3D) resolution and allows traversing the length scales of bone all the way down to the fine details of the lacuno-canalicular network housing the osteocytes. Multimodal X-ray imaging provides combined information covering both the length scales defined by the size of the measured volume and tomographic resolution, as well as those probed by the signal that is measured. In X-ray diffraction computed tomography (XRD-CT), for example, diffraction signals can be reconstructed tomographically, which offers detailed information about the spatial variations in the crystallographic properties of the bone biomineral. Orientational information can be obtained by tensor tomography. The combination of both small-angle X-ray scattering (SAXS) and wide-angle X-ray scattering (WAXS) tensor tomography gives information on the orientation of bone nanostructure and crystals, respectively. These new technical developments promise that great strides towards understanding bone structure can be expected in the near future. In this review, recent findings that have resulted from X-ray imaging are highlighted and speculation is given on what can be expected to follow.
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34

Riese, Cornelius N., Manuel Wittchen, Valérie Jérôme, Ruth Freitag, Tobias Busche, Jörn Kalinowski, and Dirk Schüler. "The transcriptomic landscape of Magnetospirillum gryphiswaldense during magnetosome biomineralization." BMC Genomics 23, no. 1 (October 10, 2022). http://dx.doi.org/10.1186/s12864-022-08913-x.

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Abstract Background One of the most complex prokaryotic organelles are magnetosomes, which are formed by magnetotactic bacteria as sensors for navigation in the Earth’s magnetic field. In the alphaproteobacterium Magnetospirillum gryphiswaldense magnetosomes consist of chains of magnetite crystals (Fe3O4) that under microoxic to anoxic conditions are biomineralized within membrane vesicles. To form such an intricate structure, the transcription of > 30 specific structural genes clustered within the genomic magnetosome island (MAI) has to be coordinated with the expression of an as-yet unknown number of auxiliary genes encoding several generic metabolic functions. However, their global regulation and transcriptional organization in response to anoxic conditions most favorable for magnetite biomineralization are still unclear. Results Here, we compared transcriptional profiles of anaerobically grown magnetosome forming cells with those in which magnetosome biosynthesis has been suppressed by aerobic condition. Using whole transcriptome shotgun sequencing, we found that transcription of about 300 of the > 4300 genes was significantly enhanced during magnetosome formation. About 40 of the top upregulated genes are directly or indirectly linked to aerobic and anaerobic respiration (denitrification) or unknown functions. The mam and mms gene clusters, specifically controlling magnetosome biosynthesis, were highly transcribed, but constitutively expressed irrespective of the growth condition. By Cappable-sequencing, we show that the transcriptional complexity of both the MAI and the entire genome decreased under anaerobic conditions optimal for magnetosome formation. In addition, predominant promoter structures were highly similar to sigma factor σ70 dependent promoters in other Alphaproteobacteria. Conclusions Our transcriptome-wide analysis revealed that magnetite biomineralization relies on a complex interplay between generic metabolic processes such as aerobic and anaerobic respiration, cellular redox control, and the biosynthesis of specific magnetosome structures. In addition, we provide insights into global regulatory features that have remained uncharacterized in the widely studied model organism M. gryphiswaldense, including a comprehensive dataset of newly annotated transcription start sites and genome-wide operon detection as a community resource (GEO Series accession number GSE197098).
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35

Li, Jinhua, Heng Zhang, Nicolas Menguy, Karim Benzerara, Fuxian Wang, Xiaoting Lin, Zhibao Chen, and Yongxin Pan. "Single-Cell Resolution of Uncultured Magnetotactic Bacteria via Fluorescence-Coupled Electron Microscopy." Applied and Environmental Microbiology 83, no. 12 (April 7, 2017). http://dx.doi.org/10.1128/aem.00409-17.

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ABSTRACTMagnetotactic bacteria (MTB) form intracellular chain-assembled nanocrystals of magnetite or greigite termed magnetosomes. The characterization of magnetosome crystals requires electron microscopy due to their nanoscopic sizes. However, electron microscopy does not provide phylogenetic information for MTB. We have developed a strategy for the simultaneous and rapid phylogenetic and biomineralogical characterization of uncultured MTB at the single-cell level. It consists of four steps: (i) enrichment of MTB cells from an environmental sample, (ii) 16S rRNA gene sequencing of MTB, and (iii) fluorescencein situhybridization analyses coordinated with (iv) transmission or scanning electron microscopy of the probe-hybridized cells. The application of this strategy identified a magnetotacticGammaproteobacteriastrain, SHHR-1, from brackish sediments collected from the Shihe River estuary in Qinhuangdao City, China. SHHR-1 magnetosomes are elongated prismatic magnetites which can be idealized as hexagonal prisms. Taxonomic groups of uncultured MTB were also identified in freshwater sediments from Lake Miyun in northern Beijing via this novel coordinated fluorescence and scanning electron microscopy method based on four group-specific rRNA-targeted probes. Our analyses revealed that major magnetotactic taxonomic groups can be accurately determined only with coordinated scanning electron microscopy observations on fluorescently labeled single cells due to limited group coverage and specificity for existing group-specific MTB fluorescencein situhybridization (FISH) probes. Our reported strategy is simple and efficient, offers great promise toward investigating the diversity and biomineralization of MTB, and may also be applied to other functional groups of microorganisms.IMPORTANCEMagnetotactic bacteria (MTB) are phylogenetically diverse and biomineralize morphologically diverse magnetic nanocrystals of magnetite or greigite in intracellular structures termed magnetosomes. However, many uncultured MTB strains have not been phylogenetically identified or structurally investigated at the single-cell level, which limits our comprehensive understanding of the diversity of MTB and their role in biomineralization. We developed a fluorescence-coupled electron microscopy method for the rapid phylogenetic and biomineralogical characterization of uncultured MTB at the single-cell level. Using this novel method, we successfully identified taxonomic groups of several uncultured MTB and one novel magnetotacticGammaproteobacteriastrain, SHHR-1, from natural environments. Our analyses further indicate that strain SHHR-1 forms elongated prismatic magnetites. Our findings provide a promising strategy for the rapid characterization of phylogenetic and biomineralogical properties of uncultured MTB at the single-cell level. Furthermore, due to its simplicity and generalized methodology, this strategy can also be useful in the study of the diversity and biomineralization properties of microbial taxa involved in other mineralization processes.
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