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  • 1
    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
    Format: text
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  • 2
    Publication Date: 2018-06-18
    Description: The control exerted by some invertebrates on the calcium carbonate polymorph produced is intriguing but not understood. Mytilus edulis shells, with the abrupt polymorph switch within their valves from an outer calcite to inner aragonite layer, are excellent examples of this phenomenon. Detailed crystallography of intact valves using Electron Backscatter Diffraction (EBSD) is considered in the context of quantitative chemical analyses by electron microprobe. Apart from the outer 40 μm, individual crystals that comprise the calcite layer of M. edulis differ from each other in terms of misorientation by less than 10°. Similar uniformity occurs in the inner aragonite layer with notable 'mineral bridging' between tablets of aragonite nacre. The first-formed aragonite laminae are submicron thickness and the subsequent laminae of uniform 1 μm thickness.Variations in chemical composition through the two valves correspond in part with the distribution of the two polymorphs. Magnesium is present in notably higher concentrations within calcite than aragonite. However, the Mg2+ concentration in calcite is not uniform and increases with growth before decreasing at the polymorph switch. Sodium concentrations decrease steadily through the calcite layer. The aragonite layer is compositionally more uniform. Sulphur is not a good proxy for organic content in this system because it does not reflect the higher organic content of the aragonite. Sector zoning is not responsible for the element distribution seen here while differences in crystal size and association with organic components remain as possible explanations.
    Type: Article , PeerReviewed
    Format: text
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  • 3
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    PANGAEA
    In:  Supplement to: Fitzer, Susan C; Cusack, Maggie; Phoenix, Vernon R; Kamenos, N A (2014): Ocean acidification reduces the crystallographic control in juvenile mussel shells. Journal of Structural Biology, 188(1), 39-45, https://doi.org/10.1016/j.jsb.2014.08.007
    Publication Date: 2019-09-18
    Description: Global climate change threatens the oceans as anthropogenic carbon dioxide causes ocean acidification and reduced carbonate saturation. Future projections indicate under saturation of aragonite, and potentially calcite, in the oceans by 2100. Calcifying organisms are those most at risk from such ocean acidification, as carbonate is vital in the biomineralisation of their calcium carbonate protective shells. This study highlights the importance of multi-generational studies to investigate how marine organisms can potentially adapt to future projected global climate change. Mytilus edulis is an economically important marine calcifier vulnerable to decreasing carbonate saturation as their shells comprise two calcium carbonate polymorphs: aragonite and calcite. M. edulis specimens were cultured under current and projected pCO2 (380, 550, 750 and 1000 µatm), following 6 months of experimental culture, adults produced second generation juvenile mussels. Juvenile mussel shells were examined for structural and crystallographic orientation of aragonite and calcite. At 1000 µatm pCO2, juvenile mussels spawned and grown under this high pCO2 do not produce aragonite which is more vulnerable to carbonate under-saturation than calcite. Calcite and aragonite were produced at 380, 550 and 750 µatm pCO2. Electron back scatter diffraction analyses reveal less constraint in crystallographic orientation with increased pCO2. Shell formation is maintained, although the nacre crystals appear corroded and crystals are not so closely layered together. The differences in ultrastructure and crystallography in shells formed by juveniles spawned from adults in high pCO2 conditions may prove instrumental in their ability to survive ocean acidification.
    Type: Dataset
    Format: text/tab-separated-values, 196 data points
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  • 4
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    PANGAEA
    In:  Supplement to: Fitzer, Susan C; Phoenix, Vernon R; Cusack, Maggie; Kamenos, N A (2014): Ocean acidification impacts mussel control on biomineralisation. Scientific Reports, 4, 6218, https://doi.org/10.1038/srep06218
    Publication Date: 2019-09-27
    Description: Ocean acidification is altering the oceanic carbonate saturation state and threatening the survival of marine calcifying organisms. Production of their calcium carbonate exoskeletons is dependent not only on the environmental seawater carbonate chemistry but also the ability to produce biominerals through proteins. We present shell growth and structural responses by the economically important marine calcifier Mytilus edulis to ocean acidification scenarios (380, 550, 750, 1000 µatm pCO2). After six months of incubation at 750 µatm pCO2, reduced carbonic anhydrase protein activity and shell growth occurs in M. edulis. Beyond that, at 1000 µatm pCO2, biomineralisation continued but with compensated metabolism of proteins and increased calcite growth. Mussel growth occurs at a cost to the structural integrity of the shell due to structural disorientation of calcite crystals. This loss of structural integrity could impact mussel shell strength and reduce protection from predators and changing environments.
    Type: Dataset
    Format: text/tab-separated-values, 2516 data points
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  • 5
    Publication Date: 2019-10-02
    Description: Biomineral production in marine organisms employs transient phases of amorphous calcium carbonate (ACC) in the construction of crystalline shells. Increasing seawater pCO2 leads to ocean acidification (OA) with a reduction in oceanic carbonate concentration which could have a negative impact on shell formation and therefore survival. We demonstrate significant changes in the hydrated and dehydrated forms of ACC in the aragonite and calcite layers of Mytilus edulis shells cultured under acidification conditions (1000 µatm pCO2) compared to present day conditions (380 µatm pCO2). In OA conditions, Mytilus edulis has more ACC at crystalisation sites. Here, we use the high-spatial resolution of synchrotron X-ray Photo Emission Electron Microscopy (XPEEM) combined with X-ray Absorption Spectroscopy (XAS) to investigate the influence of OA on the ACC formation in the shells of adult Mytilus edulis. Electron Backscatter Diffraction (EBSD) confirms that OA reduces crystallographic control of shell formation. The results demonstrate that OA induces more ACC formation and less crystallographic control in mussels suggesting that ACC is used as a repair mechanism to combat shell damage under OA. However, the resultant reduced crystallographic control in mussels raises concerns for shell protective function under predation and changing environments.
    Type: Dataset
    Format: text/tab-separated-values, 24476 data points
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  • 6
    Publication Date: 2019-10-02
    Description: Ocean acidification threatens organisms that produce calcium carbonate shells by potentially generating an under-saturated carbonate environment. Resultant reduced calcification and growth, and subsequent dissolution of exoskeletons, would raise concerns over the ability of the shell to provide protection for the marine organism under ocean acidification and increased temperatures. We examined the impact of combined ocean acidification and temperature increase on shell formation of the economically important edible mussel Mytilus edulis. Shell growth and thickness along with a shell thickness index and shape analysis were determined. The ability of M. edulis to produce a functional protective shell after 9 months of experimental culture under ocean acidification and increasing temperatures (380, 550, 750, 1000 µatm pCO2, and 750, 1000 µatm pCO2 + 2°C) was assessed. Mussel shells grown under ocean acidification conditions displayed significant reductions in shell aragonite thickness, shell thickness index, and changes to shell shape (750, 1000 ?atm pCO2) compared to those shells grown under ambient conditions (380 ?atm pCO2). Ocean acidification resulted in rounder, flatter mussel shells with thinner aragonite layers likely to be more vulnerable to fracture under changing environments and predation. The changes in shape presented here could present a compensatory mechanism to enhance protection against predators and changing environments under ocean acidification when mussels are unable to grow thicker shells. Here, we present the first assessment of mussel shell shape to determine implications for functional protection under ocean acidification.
    Type: Dataset
    Format: text/tab-separated-values, 1680 data points
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  • 7
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    PANGAEA
    In:  Supplement to: Fitzer, Susan C; Zhu, Wenzhong; Tanner, K Elizabeth; Phoenix, Vernon R; Kamenos, N A; Cusack, Maggie (2014): Ocean acidification alters the material properties of Mytilus edulis shells. Journal of The Royal Society Interface, 12(103), 20141227-20141227, https://doi.org/10.1098/rsif.2014.1227
    Publication Date: 2019-11-30
    Description: Ocean acidification (OA) and the resultant changing carbonate saturation states is threatening the formation of calcium carbonate shells and exoskeletons of marine organisms. The production of biominerals in such organisms relies on the availability of carbonate and the ability of the organism to biomineralize in changing environments. To understand how biomineralizers will respond to OA the common blue mussel, Mytilus edulis, was cultured at projected levels of pCO2 (380, 550, 750, 1000 µatm) and increased temperatures (ambient, ambient plus 2°C). Nanoindentation (a single mussel shell) and microhardness testing were used to assess the material properties of the shells. Young's modulus (E), hardness (H) and toughness (KIC) were measured in mussel shells grown in multiple stressor conditions. OA caused mussels to produce shell calcite that is stiffer (higher modulus of elasticity) and harder than shells grown in control conditions. The outer shell (calcite) is more brittle in OA conditions while the inner shell (aragonite) is softer and less stiff in shells grown under OA conditions. Combining increasing ocean pCO2 and temperatures as projected for future global ocean appears to reduce the impact of increasing pCO2 on the material properties of the mussel shell. OA may cause changes in shell material properties that could prove problematic under predation scenarios for the mussels; however, this may be partially mitigated by increasing temperature.
    Type: Dataset
    Format: text/tab-separated-values, 22000 data points
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  • 8
    Publication Date: 2016-02-15
    Electronic ISSN: 2045-2322
    Topics: Natural Sciences in General
    Published by Springer Nature
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  • 9
    Publication Date: 2007-09-01
    Description: With their extensive fossil record and shells of stable low-Mg calcite, rhynchonelliform brachiopods are attractive sources of climate information via seawater temperature proxies such as stable oxygen isotope composition. In Terebratalia transversa (Sowerby) there is a progression towards oxygen isotope equilibrium in the calcite of the innermost secondary layer. This study confirms the lack of any vital effects influencing oxygen isotope composition of T. transversa, even in specialised areas of the innermost secondary layer. Calcite Mg/Ca ratio is another potential seawater temperature proxy, that has the advantage of not being influenced by salinity. Mg concentrations measured by electron microprobe analyses indicate that there is no concomitant decrease in Mg concentration towards the inner secondary layer, associated with the progressive shift towards oxygen isotope equilibrium. Mg distribution is heterogeneous throughout the shell and correlates with that of sulphur, which may be a proxy for organic components, suggesting that some of the Mg may not be in the calcite lattice. It is essential therefore, to determine the chemical environment of the magnesium ions to avoid any erroneous temperature extrapolations in brachiopods or any other calcite biomineral.
    Print ISSN: 1755-6910
    Electronic ISSN: 1755-6929
    Topics: Geosciences
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  • 10
    Publication Date: 2007-09-01
    Description: Electron backscatter diffraction (EBSD) is used to determine the detailed crystallographic orientation of calcite crystals of craniid brachiopods in the context of shell ultrastructure. Sections of shells of two Recent species, Novocrania anomala and Novocrania huttoni, are analysed to provide 3D crystallographic patterns at high spatial resolution. The c-axis of semi-nacre calcite crystals is oriented parallel to the laminae that define the ultrastructure of the secondary layer. This orientation differs from that of rhynchonelliform calcitic brachiopods where the c-axis is perpendicular to the length of morphological fibres and to the shell exterior.
    Print ISSN: 1755-6910
    Electronic ISSN: 1755-6929
    Topics: Geosciences
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