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  • 1
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    Formation and mosaicity of coccolith segment calcite of the marine algae Emiliania huxleyi (2018)
    Wiley
    In:  Journal of Phycology, 54 (1). pp. 85-104.
    Details
    Publication Date: 2021-02-08
    Description: Coccolithophores belong to the most abundant calcium carbonate mineralizing organisms. Coccolithophore biomineralization is a complex and highly regulated process, resulting in a product that strongly differs in its intricate morphology from the abiogenically produced mineral equivalent. Moreover, unlike extracellularly formed biological carbonate hard tissues, coccolith calcite is neither a hybrid composite, nor is it distinguished by a hierarchical microstructure. This is remarkable as the key to optimizing crystalline biomaterials for mechanical strength and toughness lies in the composite nature of the biological hard tissue and the utilization of specific microstructures. To obtain insight into the pathway of biomineralization of Emiliania huxleyi coccoliths, we examine intracrystalline nanostructural features of the coccolith calcite in combination with cell ultrastructural observations related to the formation of the calcite in the coccolith vesicle within the cell. With TEM diffraction and annular dark-field imaging, we prove the presence of planar imperfections in the calcite crystals such as planar mosaic block boundaries. As only minor misorientations occur, we attribute them to dislocation networks creating small-angle boundaries. Intracrystalline occluded biopolymers are not observed. Hence, in E. huxleyi calcite mosaicity is not caused by occluded biopolymers, as it is the case in extracellularly formed hard tissues of marine invertebrates, but by planar defects and dislocations which are typical for crystals formed by classical ion-by-ion growth mechanisms. Using cryo-preparation techniques for SEM and TEM, we found that the membrane of the coccolith vesicle and the outer membrane of the nuclear envelope are in tight proximity, with a well-controlled constant gap of ~4 nm between them. We describe this conspicuous connection as a not yet described interorganelle junction, the “nuclear envelope junction”. The narrow gap of this junction likely facilitates transport of Ca2+ ions from the nuclear envelope to the coccolith vesicle. On the basis of our observations, we propose that formation of the coccolith utilizes the nuclear envelope–endoplasmic reticulum Ca2+-store of the cell for the transport of Ca2+ ions from the external medium to the coccolith vesicle and that E. huxleyi calcite forms by ion-by-ion growth rather than by a nanoparticle accretion mechanism.
    Type: Article , PeerReviewed
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  • 2
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    Mapping of recent brachiopod microstructure: A tool for environmental studies (2018)
    Elsevier
    In:  Journal of Structural Biology, 201 (3). pp. 221-236.
    Details
    Publication Date: 2019-02-01
    Description: Shells of brachiopods are excellent archives for environmental reconstructions in the recent and distant past as their microstructure and geochemistry respond to climate and environmental forcings. We studied the morphology and size of the basic structural unit, the secondary layer fibre, of the shells of several extant brachiopod taxa to derive a model correlating microstructural patterns to environmental conditions. Twenty-one adult specimens of six recent brachiopod species adapted to different environmental conditions, from Antarctica, to New Zealand, to the Mediterranean Sea, were chosen for microstructural analysis using SEM, TEM and EBSD. We conclude that: 1) there is no significant difference in the shape and size of the fibres between ventral and dorsal valves, 2) there is an ontogenetic trend in the shape and size of the fibres, as they become larger, wider, and flatter with increasing age. This indicates that the fibrous layer produced in the later stages of growth, which is recommended by the literature to be the best material for geochemical analyses, has a different morphostructure and probably a lower organic content than that produced earlier in life. In two species of the same genus living in seawater with different temperature and carbonate saturation state, a relationship emerged between the microstructure and environmental conditions. Fibres of the polar Liothyrella uva tend to be smaller, rounder and less convex than those of the temperate Liothyrella neozelanica, suggesting a relationship between microstructural size, shell organic matter content, ambient seawater temperature and calcite saturation state.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 3
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    Archival biogenic micro- and nanostructure data analysis: Signatures of diagenetic systems (2018)
    Elsevier
    In:  Data in Brief, 19 . pp. 299-311.
    Details
    Publication Date: 2021-02-08
    Description: The present data in brief article provides additional data and information to our research article “Micro- and nanostructures reflect the degree of diagenetic alteration in modern and fossil brachiopod shell calcite: a multi-analytical screening approach (CL, FE-SEM, AFM, EBSD)” [1] (Casella et al.). We present fibre morphology, nano- and microstructure, as well as calcite crystal orientations and textures found in pristine, experimentally altered (hydrothermal and thermal), and diagenetically overprinted brachiopod shells. Combination of the screening tools AFM, FE-SEM, and EBSD allows to observe a significant change in microstructural and textural features with an increasing degree of laboratory-based and naturally occurring diagenetic alteration. Amalgamation of neighbouring fibres was observed on the micrometre scale level, whereas progressive decomposition of biopolymers in the shells and fusion of nanoparticulate calcite crystals was detected on the nanometre scale. The presented data in this article and the study described in [1] allows for qualitative information on the degree of diagenetic alteration of fossil archives used for palaeoclimate reconstruction.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 4
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    Hydrothermal alteration of aragonitic biocarbonates: assessment of micro- and nanostructural dissolution–reprecipitation and constraints of diagenetic overprint from quantitative statistical grain-area analysis (2018)
    Copernicus Publications (EGU)
    In:  Biogeosciences (BG), 15 (24). pp. 7451-7484.
    Details
    Publication Date: 2021-02-08
    Description: The assessment of diagenetic overprint on microstructural and geochemical data gained from fossil archives is of fundamental importance for understanding palaeoenvironments. The correct reconstruction of past environmental dynamics is only possible when pristine skeletons are unequivocally distinguished from altered skeletal elements. Our previous studies show (i) that replacement of biogenic carbonate by inorganic calcite occurs via an interface-coupled dissolution–reprecipitation mechanism. (ii) A comprehensive understanding of alteration of the biogenic skeleton is only given when structural changes are assessed on both, the micrometre as well as on the nanometre scale. In the present contribution we investigate experimental hydrothermal alteration of six different modern biogenic carbonate materials to (i) assess their potential for withstanding diagenetic overprint and to (ii) find characteristics for the preservation of their microstructure in the fossil record. Experiments were performed at 175°C with a 100 mM NaCl + 10 mM MgCl2 alteration solution and lasted for up to 35 days. For each type of microstructure we (i) examine the evolution of biogenic carbonate replacement by inorganic calcite, (ii) highlight different stages of inorganic carbonate formation, (iii) explore microstructural changes at different degrees of alteration, and (iv) perform a statistical evaluation of microstructural data to highlight changes in crystallite size between the pristine and the altered skeletons. We find that alteration from biogenic aragonite to inorganic calcite proceeds along pathways where the fluid enters the material. It is fastest in hard tissues with an existing primary porosity and a biopolymer fabric within the skeleton that consists of a network of fibrils. The slowest alteration kinetics occurs when biogenic nacreous aragonite is replaced by inorganic calcite, irrespective of the mode of assembly of nacre tablets. For all investigated biogenic carbonates we distinguish the following intermediate stages of alteration: (i) decomposition of biopolymers and the associated formation of secondary porosity, (ii) homoepitactic overgrowth with preservation of the original phase leading to amalgamation of neighbouring mineral units (i.e. recrystallization by grain growth eliminating grain boundaries), (iii) deletion of the original microstructure, however, at first, under retention of the original mineralogical phase, and (iv) replacement of both, the pristine microstructure and original phase with the newly formed abiogenic product. At the alteration front we find between newly formed calcite and reworked biogenic aragonite the formation of metastable Mg-rich carbonates with a calcite-type structure and compositions ranging from dolomitic to about 80mol % magnesite. This high-Mg calcite seam shifts with the alteration front when the latter is displaced within the unaltered biogenic aragonite. For all investigated biocarbonate hard tissues we observe the destruction of the microstructure first, and, in a second step, the replacement of the original with the newly formed phase.
    Type: Article , PeerReviewed
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  • 5
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    Calcite fibre formation in modern brachiopod shells (2019)
    Nature Research
    In:  Scientific Reports, 9 (598).
    Details
    Publication Date: 2022-01-31
    Description: The fibrous calcite layer of modern brachiopod shells is a hybrid composite material and forms a substantial part of the hard tissue. We investigated how cells of the outer mantle epithelium (OME) secrete calcite material and generate the characteristic fibre morphology and composite microstructure of the shell. We employed AFM, FE-SEM, and TEM imaging of embedded/etched, chemically fixed/decalcified and high-pressure frozen/freeze substituted samples. Calcite fibres are secreted by outer mantle epithelium (OME) cells. Biometric analysis of TEM micrographs indicates that about 50% of these cells are attached via hemidesmosomes to an extracellular organic membrane present at the proximal, convex surface of the fibres. At these sites, mineral secretion is not active. Instead, ion transport from OME cells to developing fibres occurs at regions of closest contact between cells and fibres, however only at sites where the extracellular membrane at the proximal fibre surface is not developed yet. Fibre formation requires the cooperation of several adjacent OME cells. It is a spatially and temporally changing process comprising of detachment of OME cells from the extracellular organic membrane, mineral secretion at detachment sites, termination of secretion with formation of the extracellular organic membrane, and attachment of cells via hemidesmosomes to this membrane.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 6
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    The evolution of thecideide microstructures and textures: traced from Triassic to Holocene (2021)
    Wiley
    Details
    Publication Date: 2024-02-07
    Description: Thecideide brachiopods are an anomalous group of invertebrates. In this study, we discuss the evolution of thecideide brachiopods from the Triassic to the Holocene and base our results and conclusions on microstructure and texture measurements gained from electron backscatter diffraction (EBSD). In fossil and Recent thecideide shells, we observe the following mineral units: (1) nanometric to small granules; (2) acicles; (3) fibres; (4) polygonal crystals; and (5) large roundish crystals. We trace for thecideide shells the change of mineral unit characteristics such as morphology, size, orientation, arrangement and distribution pattern. Triassic thecideide shells contain extensive sections formed of fibres interspersed with large, roundish crystals. Upper Cretaceous to Pleistocene thecideide hard tissues consist of a matrix of minute to small grains reinforced by acicles and small polygonal crystals. Recent thecideide species form their shell of mineral units that show a wide range of shapes, sizes and arrangements. We find from Late Triassic to Recent a gradual decrease in mineral unit size, regularity of mineral unit morphology and orientation and the degree of calcite co‐orientation. While crystallite co‐orientation is the highest for fibrous microstructures, it is strikingly low for taxa that form their shell out of nanogranular to acicular mineral units. Our results indicate that Upper Jurassic species represent transitional forms between ancient taxa with fibrous shells and Recent forms that construct their shells of acicles and granules. We attribute the observed changes in microstructure and texture to be an adaptation to a different habitat and lifestyle associated with cementation to hard substrates.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 7
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    The architecture of Recent brachiopod shells: diversity of biocrystal and biopolymer assemblages in rhynchonellide, terebratulide, thecideide and craniide shells (2022)
    Springer
    Details
    Publication Date: 2024-02-07
    Description: Biological hard tissues are a rich source of design concepts for the generation of advanced materials. They represent the most important library of information on the evolution of life and its environmental conditions. Organisms produce soft and hard tissues in a bottom-up process, a construction principle that is intrinsic to biologically secreted materials. This process emerged early on in the geological record, with the onset of biological mineralization. The phylum Brachiopoda is a marine animal group that has an excellent and continuous fossil record from the early Cambrian to the Recent. Throughout this time interval, the Brachiopoda secreted phosphate and carbonate shells and populated many and highly diverse marine habitats. This required great flexibility in the adaptation of soft and hard tissues to the different marine environments and living conditions. This review presents, juxtaposes and discusses the main modes of mineral and biopolymer organization in Recent, carbonate shell-producing, brachiopods. We describe shell tissue characteristics for taxa of the orders Rhynchonellida, Terebratulida, Thecideida and Craniida. We highlight modes of calcite and organic matrix assembly at the macro-, micro-, and nano-scales based on results obtained by Electron Backscatter Diffraction, Atomic Force Microscopy, Field Emission Scanning Electron Microscopy and Scanning Transmission Electron Microscopy. We show variation in composite hard tissue organization for taxa with different lifestyles, visualize nanometer-scale calcite assemblies for rhynchonellide and terebratulide fibers, highlight thecideide shell microstructure, texture and chemistry characteristics, and discuss the feasibility to use thecideide shells as archives of proxies for paleoenvironment and paleoclimate reconstructions.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 8
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    Isotope ratios, trace elements and ultrastructure results of Mytilus galloprovincialis shells and experimental site data off Ischia (Italy)
    PANGAEA
    In:  Supplement to: Hahn, Sabine; Rodolfo-Metalpa, Riccardo; Griesshaber, Erika; Schmahl, Wolfgang W; Buhl, Dieter; Hall-Spencer, Jason M; Baggini, Cecilia; Fehr, Karl T; Immenhauser, Adrian (2012): Marine bivalve shell geochemistry and ultrastructure from modern low pH environments: environmental effect versus experimental bias. Biogeosciences, 9, 1897-1914, https://doi.org/10.5194/bg-9-1897-2012
    Details
    Publication Date: 2023-05-12
    Description: Bivalve shells can provide excellent archives of past environmental change but have not been used to interpret ocean acidification events. We investigated carbon, oxygen and trace element records from different shell layers in the mussels Mytilus galloprovincialis combined with detailed investigations of the shell ultrastructure. Mussels from the harbour of Ischia (Mediterranean, Italy) were transplanted and grown in water with mean pHT 7.3 and mean pHT 8.1 near CO2 vents on the east coast of the island. Most prominently, the shells recorded the shock of transplantation, both in their shell ultrastructure, textural and geochemical record. Shell calcite, precipitated subsequently under acidified seawater responded to the pH gradient by an in part disturbed ultrastructure. Geochemical data from all test sites show a strong metabolic effect that exceeds the influence of the low-pH environment. These field experiments showed that care is needed when interpreting potential ocean acidification signals because various parameters affect shell chemistry and ultrastructure. Besides metabolic processes, seawater pH, factors such as salinity, water temperature, food availability and population density all affect the biogenic carbonate shell archive.
    Keywords: BIOACID; Biological Impacts of Ocean Acidification; Mediterranean Sea Acidification in a Changing Climate; MedSeA
    Type: Dataset
    Format: application/zip, 4 datasets
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  • 9
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    (Table S1) Results of isotope analysis from shells of Mytilus galloprovincialis at the experimental site B1 and control site C
    PANGAEA
    Details
    Publication Date: 2023-05-12
    Keywords: East coast of Ischia, Italia; ES; Event label; HAND; Ischia_B1; Ischia_C; Mediterranean Sea Acidification in a Changing Climate; MedSeA; Mytilus galloprovincialis, distance from commissure; OS; Sample code/label; Sample type; Sampling by hand; δ13C; δ13C, standard deviation; δ18O, carbonate; δ18O, standard deviation
    Type: Dataset
    Format: text/tab-separated-values, 1190 data points
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  • 10
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    (Table 1) Seawater chemistry calculated over the experiment period at the experimental site B1 and control site C
    PANGAEA
    Details
    Publication Date: 2023-05-12
    Keywords: Aragonite saturation state; Aragonite saturation state, standard deviation; Bicarbonate, standard deviation; Bicarbonate ion; Calcite saturation state; Calcite saturation state, standard deviation; Carbon, inorganic, dissolved; Carbon, inorganic, dissolved, standard deviation; Carbonate ion; Carbonate ion, standard deviation; Carbon dioxide; Carbon dioxide, partial pressure; Carbon dioxide, partial pressure, standard deviation; Carbon dioxide, standard deviation; DEPTH, water; East coast of Ischia, Italia; ES; Event label; HAND; IP; Ischia_B1; Ischia_C; Ischia_harbour; Mediterranean Sea Acidification in a Changing Climate; MedSeA; OS; pH; pH, standard deviation; Sampling by hand; Temperature, standard deviation; Temperature, water; δ13C; δ13C, standard deviation; δ18O, maximum; δ18O, minimum; δ18O, standard deviation; δ18O, water
    Type: Dataset
    Format: text/tab-separated-values, 72 data points
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