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Biomineralisation by earthworms: an investigation into the stability and distribution of amorphous calcium carbonate
Background
Many biominerals form from amorphous calcium carbonate (ACC), but this phase is highly unstable when synthesised in its pure form inorganically. Several species of earthworm secrete calcium carbonate granules which contain highly stable ACC. We analysed the milky fluid from which granules form and solid granules for amino acid (by liquid chromatography) and functional group (by Fourier transform infrared (FTIR) spectroscopy) compositions. Granule elemental composition was determined using inductively coupled plasma-optical emission spectroscopy (ICP-OES) and electron microprobe analysis (EMPA). Mass of ACC present in solid granules was quantified using FTIR and compared to granule elemental and amino acid compositions. Bulk analysis of granules was of powdered bulk material. Spatially resolved analysis was of thin sections of granules using synchrotron-based μ-FTIR and EMPA electron microprobe analysis.
Results
The milky fluid from which granules form is amino acid-rich (≤ 136 ± 3 nmol mg−1 (n = 3; ± std dev) per individual amino acid); the CaCO3 phase present is ACC. Even four years after production, granules contain ACC. No correlation exists between mass of ACC present and granule elemental composition. Granule amino acid concentrations correlate well with ACC content (r ≥ 0.7, p ≤ 0.05) consistent with a role for amino acids (or the proteins they make up) in ACC stabilisation. Intra-granule variation in ACC (RSD = 16%) and amino acid concentration (RSD = 22–35%) was high for granules produced by the same earthworm. Maps of ACC distribution produced using synchrotron-based μ-FTIR mapping of granule thin sections and the relative intensity of the ν2: ν4 peak ratio, cluster analysis and component regression using ACC and calcite standards showed similar spatial distributions of likely ACC-rich and calcite-rich areas. We could not identify organic peaks in the μ-FTIR spectra and thus could not determine whether ACC-rich domains also had relatively high amino acid concentrations. No correlation exists between ACC distribution and elemental concentrations determined by EMPA.
Conclusions
ACC present in earthworm CaCO3 granules is highly stable. Our results suggest a role for amino acids (or proteins) in this stability. We see no evidence for stabilisation of ACC by incorporation of inorganic components
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Barium and its Importance as an Indicator of (Paleo)Productivity
Barium (Ba) is a trace element which occurs predominantly as barite mineral (BaSO4) in the marine environment. Previous work suggests that barite concentrations are related to the organic carbon flux and marine biological debris in the water column suggesting a direct or indirect involvement in the marine biological cycling. In addition, barite has a high preservation rate (~30%) in sediments and it is less affected by early diagenesis than other proxies for productivity such as carbonates (~10%) and organic carbon (~1%), for example. Therefore, Ba is considered an excellent proxy for ocean (paleo)productivity. However, correlating barite to productivity involves some caveats. Specifically, the post-depositional formation of barite in oxic sediments can lead to Ba release into porewaters under anoxic conditions, which can form barite again under oxic conditions. This diagenetic formation is not correlated to export production as the seawater authigenic barite formed with decaying organic matter in the water column. Therefore, the main goal of this work is to briefly review the marine Ba cycle and highlight its importance for (paleo)productivity research