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Jan 26, 2006 - biological processes that may influence the Mg/Ca ratio of the shell are suggested and a model that combines ... predominantly determined by biological factors ...... tium, fluorine, and cadmium into benthic foraminiferal.
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Volume 7, Number 1 26 January 2006 Q01P08, doi:10.1029/2005GC001015

AN ELECTRONIC JOURNAL OF THE EARTH SCIENCES Published by AGU and the Geochemical Society

ISSN: 1525-2027

Impact of biomineralization processes on the Mg content of foraminiferal shells: A biological perspective Shmuel Bentov and Jonathan Erez Institute of Earth Sciences, The Hebrew University of Jerusalem, Jerusalem, 91904 Israel ([email protected])

[1] The Mg/Ca ratio in foraminiferal shells is widely used as a proxy for paleotemperatures. Nevertheless, it seems that the basic Mg content of foraminifera is determined by biological factors, as can be concluded from the large inter species and intrashell variability and the frequent deviations from inorganic behavior. This paper discusses three possible ways by which foraminifera can control or modify the Mg content in their shell: (1) involvement of organic matrix in the precipitation process that may alter the partition coefficient of Mg in biogenic calcite, (2) controlled conversion of transient amorphous phases to calcite, and (3) modification of the Mg concentration in the parent solution from which the crystals precipitate. The first two mechanisms are probably responsible for the precipitation of high-Mg calcite phases (whole shell or sublayers), while the third mechanism leads to the formation of low-Mg calcite phases. We propose a model adapted from epithelial cells that allows massive Mg2+ removal from the biomineralization site. This model is especially relevant to the planktonic and deep benthic low-Mg foraminifera that are frequently used for paleotemperature reconstructions. We discuss the possible biological roles of Mg in the shell in terms of the calcite polymorph conservation, the in vivo chemical stability of the shell, the functions of Mg as a stabilizer of transient phases and as a controlling agent of the precipitation process. Several temperature sensitive biological processes that may influence the Mg/Ca ratio of the shell are suggested and a model that combines biogenic and inorganic considerations is presented. The model uses Mg heterogeneity in the shell together with temperature response (biologic and inorganic) of biomineralization processes, to account for the deviation of planktonic foraminifera from inorganic calcite at equilibrium with seawater. Components: 7644 words, 3 figures. Keywords: biomineralization; magnesium; paleotemperature; vital effects. Index Terms: 0419 Biogeosciences: Biomineralization; 0473 Biogeosciences: Paleoclimatology and paleoceanography (3344, 4900); 1050 Geochemistry: Marine geochemistry (4835, 4845, 4850). Received 11 May 2005; Revised 25 September 2005; Accepted 27 October 2005; Published 26 January 2006. Bentov, S., and J. Erez (2006), Impact of biomineralization processes on the Mg content of foraminiferal shells: A biological perspective, Geochem. Geophys. Geosyst., 7, Q01P08, doi:10.1029/2005GC001015. ————————————

Theme: Development of the Foraminiferal Mg/Ca Proxy for Paleoceanography Guest Editor: Pamela Martin

1. Introduction [2] Field and laboratory calibrations demonstrated that Mg/Ca variations within a foraminiferal species are well correlated with temperature changes [e.g., Nurnberg et al., 1996; Rosenthal et al., 1997; Lear et al., 2000; Anand et al., 2003; Russell et al., 2004]. Copyright 2006 by the American Geophysical Union

However, it was recognized that there are major ‘‘vital effects’’ that influence the initial Mg/Ca ratio of a given species and its response to temperature. The physiological processes that govern and/or influence the Mg content in foraminifera shells and their response to temperature are poorly known. We will try to elucidate this issue and consider some 1 of 11

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Figure 1. Mg content of various foraminifera groups based on Blackmon and Todd [1959]. The average value is marked by red squares and the ranges are shown by vertical bars. At the right, the range of values expected for inorganic precipitation of calcite from seawater based on values at 5 and 25C according to Mucci [1987] is shown.

potential physiological-mediated responses of the Mg/Ca ratio to temperature in foraminifera.

1.1. Taxonomic Variability of Mg/Ca Ratio [3] The foraminifera as a group display a very large variability in the Mg content of their carbonate shells, ranging between more than 20 mole% to less than 0.1 mole% of MgCO3 [Blackmon and Todd, 1959] (Figure 1). On the basis of Mg content, the foraminifera have been divided to three major groups: high-Mg (>10%), intermediate-Mg (5 – 10%), and low-Mg (10 mol%) while the secondary layers that cover and thicken the shell wall contain much less Mg (3 mol%). In vivo microscopic observations suggest that these two calcite phases were formed in different biomineralization pathways. When evaluating the final Mg content of a shell, the sublayers and their relative proportion must be taken into consideration. This proportion may change with various environmental factors and can also play an

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important role in determining the apparent temperature dependency of the foraminiferal Mg/Ca ratio. For example, the steep Mg/Ca response to temperature in low-Mg foraminifera compared to inorganic calcite [Nurnberg et al., 1996; Rosenthal et al., 1997] may be caused by a temperature-mediated increase in the proportion of the high-Mg primary calcite. In Figure 3, we simulated this scenario by increasing the ratio between the high–Mg and low-Mg phases as a function of temperature. For both phases we assumed the inorganic dependence of DMg on temperature according to Oomori et al. [1987]. We further assume a four fold difference in the Mg content between the phases, and an increase in the proportion of the high Mg phase by 0.2% per C. The combined curve that was obtained, which represents the total Mg content, is in good fit to the observed Mg/Ca temperature dependence of planktonic foraminifera, a slope that is steeper than that of the inorganic calcite (Figure 3).

7. Summary [32] Foraminifera display large variability in the Mg content of different genera from the same environment, large Mg inhomogeneity within individual shells and significant deviations from equilibrium Mg/Ca ratios expected for inorganic calcite. All these phenomena suggest a strong biological control on the incorporation of Mg into foraminiferal shells. We propose that the high variability of Mg/Ca ratio between species represents different biomineralization pathways that developed during the evolution of foraminifera. The high-Mg calcite shells may precipitate from transient amorphous precursors and/or with the influence of an organic matrix. We propose that the low-Mg calcite shells precipitate from confined seawater parent solution that is depleted in Mg. The Mg ions may be removed from this solution by specific Mg channels into the cytosol and/or other cellular compartments, and later pumped out of the cell against a concentration gradient. Nevertheless, field and laboratory calibrations show that the Mg/Ca in foraminifera is a good recorder of ambient temperature and Mg/Ca ratio in seawater, suggesting a certain inorganic control and possibly a physiological response that may also be sensitive to these ambient conditions. A better understanding of the biomineralization processes in foraminifera and their associated vital effects, will allow us to use Mg/Ca ratios and other 9 of 11

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paleoceanographic proxies with higher degree of confidence.

Acknowledgments [33] The authors would like to thank the reviewers (E. Boyle and P. Dove) and the Associate Editor (P. Martin) for their helpful comments. We thank M. Edelman for critically reading the manuscript and his valuable assistance. Funding for this project was provided by the U.S.-Israel Science Foundation (grant 2000284) and German – Israeli Foundation (grant G-720-145.8/01). This work is part of the Ph.D. thesis of S. Bentov at the Hebrew University of Jerusalem.

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