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Coccolithophores are biogeochemically and ecologically important phytoplankton that produce a composite calcium carbonate-based exoskeleton – the coccosphere – comprised of individual platelets, known as coccoliths. Coccoliths are stunning examples of biomineralization; their formation featuring exceptional control over both biomineral chemistry and shape. Understanding how coccoliths are formed requires information about minor element distribution and chemical environment. Here, the first high-resolution 3D synchrotron X-ray fluorescence (XRF) mapping of a coccolith is presented, showing that the lopadoliths of Scyphosphaera apsteinii display stripes of different Sr concentration. The presence of Sr stripes is unaffected by elevated Sr in the culture medium, macro-nutrient concentration, and light intensity, indicating that the observed stripiness is an expression of the fundamental coccolith formation process in this species. Current Sr fractionation models, by contrast, predict an even Sr distribution and will have to be modified to account for this stripiness. Additionally, nano-XANES analyses show that Sr resides in a Ca site in the calcite lattice in both high and low Sr stripes, confirming a central assumption of current Sr fractionation models. [ABSTRACT FROM AUTHOR]

Coccolithophores have a distinct haplo-diplontic life cycle, which allows them to grow and divide into two different life cycle phases (haploid and diploid). These life cycle phases vary significantly in inorganic carbon content and morphology and inhabit distinct niches, with haploids generally preferring low-nitrogen and high-temperature and high-light environments in situ. This niche contrast indicates different physiology of the life cycle phases, which is considered here in the context of a trait trade-off framework, in which a particular set of traits comes with both costs and benefits. However, coccolithophore's phase trade-offs are not fully identified, limiting our understanding of the functionality of the coccolithophore life cycle. Here, we investigate the response of the two life cycle phases of the coccolithophore Coccolithus braarudii to key environmental drivers: light, temperature, and nitrogen, using laboratory experiments. With these data, we identify the main trade-offs of each life cycle phase and use models to test the role of such trade-offs under different environmental conditions. The lab experiments show the life cycle phases have similar cell size, minimum nitrogen quotas, uptake rates, and temperature and light optima. However, we find that they have different coccosphere sizes, maximum growth rates, and maximum nitrogen quotas. We also observe a trade-off between maximum growth rate and maximum nitrogen quota, with higher growth rates and low maximum nitrogen quotas in the haploid phase and vice versa in the diploid phase. Testing these phase characteristics in a numerical chemostat model, we find that the growth–quota trade-off allows C. braarudii to exploit variable nitrogen conditions more efficiently. Because the diploid ability to store more nitrogen is advantageous when the nitrogen supply is intermittent, the higher haploid growth rate is advantageous when the nitrogen supply is constant. Although the ecological drivers of C. braarudii life cycle fitness are likely multi-faceted, spanning both top-down and bottom-up trait trade-offs, our results suggest that a trade-off between nitrogen storage and maximum growth rate is an essential bottom-up control on the distribution of C. braarudii life cycle phases. [ABSTRACT FROM AUTHOR]

Coccolithophores have a distinct haplo-diplontic life cycle, which allows them to grow and divide in two different life cycle phases (haploid and diploid). These life cycle phases vary significantly in inorganic carbon content and morphology, and inhabit distinct niches, with haploids generally preferring low-nutrient and high-temperature and light environments. This niche contrast indicates different physiology of the life cycle phases, which is considered here in the context of a trait trade-off framework, in which a particular set of traits comes with both costs and benefits. However, coccolithophore's phase trade-offs are not fully identified, limiting our understanding of the functionality of the coccolithophore life cycle. Here, we investigate the response of the two life cycle phases of the coccolithophore Coccolithus braarudii to key environmental drivers: light, temperature and nutrients, using laboratory experiments. With this data, we identify the main trade-offs of each life cycle phase and use models to test the role of such trade-offs under different environmental conditions. The lab experiments show the life cycle phases have similar cell size, nitrogen requirement, uptake rates, and temperature and light optima. However, we find that they have different coccosphere sizes, maximum growth rates and nitrogen quotas. We also observe a trade-off between maximum growth rate and nitrogen quota, with higher growth rates and small nitrogen storage in the haploid phase and vice versa in the diploid phase. Testing these phase characteristics in the model, we find that the growth-quota trade-off allows C. braarudii to exploit variable nitrogen conditions more efficiently. Because while the diploid ability to store more nitrogen is advantageous when the nitrogen supply is intermittent, the higher haploid growth rate is advantageous when the nitrogen supply is constant. Although the ecological drivers of C. braarudii life cycle fitness are likely multi-faceted, spanning both top-down and bottom-up trait trade-offs, our results suggest that a trade-off between nitrogen storage and maximum growth rate is an essential bottom-up control on the distribution of C. braarudii life cycle phases. [ABSTRACT FROM AUTHOR]

The calcite platelets of coccolithophores (Haptophyta), the coccoliths, are among the most elaborate biomineral structures. How these unicellular algae accomplish the complex morphogenesis of coccoliths is still largely unknown. It has long been proposed that the cytoskeleton plays a central role in shaping the growing coccoliths. Previous studies have indicated that disruption of the microtubule network led to defects in coccolith morphogenesis in Emiliania huxleyi and Coccolithus braarudii. Disruption of the actin network also led to defects in coccolith morphology in E. huxleyi, but its impact on coccolith morphology in C. braarudii was unclear, as coccolith secretion was largely inhibited under the conditions used. A more detailed examination of the role of actin and microtubule networks is therefore required to address the wider role of the cytoskeleton in coccolith morphogenesis. In this study, we have examined coccolith morphology in C. braarudii and Scyphosphaera apsteinii following treatment with the microtubule inhibitors vinblastine and colchicine (S. apsteinii only) and the actin inhibitor cytochalasin B. We found that all cytoskeleton inhibitors induced coccolith malformations, strongly suggesting that both microtubules and actin filaments are instrumental in morphogenesis. By demonstrating the requirement for the microtubule and actin networks in coccolith morphogenesis in diverse species, our results suggest that both of these cytoskeletal elements are likely to play conserved roles in defining coccolith morphology. [ABSTRACT FROM AUTHOR]

Coccolithophores feature a haplo-diplontic life cycle comprised of diploid cells producing heterococcoliths and haploid cells producing morphologically different holococcoliths. These life cycle phases of each species appear to have distinct spatial and temporal distributions in the oceans, with the heavily calcified heterococcolithophores (HET) often more prevalent in winter and at greater depths, whilst the lightly calcified holococcolithophores (HOL) are more abundant in summer and in shallower waters. The haplo-diplontic life cycle may therefore allow coccolithophores to expand their ecological niche, switching between life cycle phases to exploit conditions that are more favourable. However, coccolithophore life cycles remain poorly understood and fundamental information on the physiological differences between life cycle phases is required if we are to better understand the ecophysiology of coccolithophores. In this study, we have examined the physiology of HET and HOL phases of the coccolithophore Coccolithus braarudii in response to changes in light and nutrient availability. We found that the HOL phase was more tolerant to high light than the HET phase, which exhibited defects in calcification at high irradiances. The HET phase exhibited defects in coccolith formation under both nitrate (N) and phosphate (P) limitation, whilst no defects in calcification were detected in the HOL phase. The HOL phase grew to a higher cell density under P-limitation than N-limitation, whereas no difference was observed in the maximum cell density reached by the HET phase at these nutrient concentrations. HET cells grown under a light:dark cycle divided primarily in the dark and early part of the light phase, whereas HOL cells continued to divide throughout the 24 h period. The physiological differences may contribute to the distinct biogeographic distributions observed between life cycle phases, with the HOL phase potentially better adapted to high light, low nutrient regimes, such as those found in seasonally stratified surface waters. Coccolithus braarudii life cycle phases exhibit different physiological responses. The heavily calcified heterococcolithophores (HET) life cycle phase is more sensitive to high light. The lightly calcified holococcolithophores (HOL) life cycle phase may be better suited to growth under low phosphate availability. [ABSTRACT FROM AUTHOR]

Coccolithophores are an important group of calcifying marine phytoplankton. Although coccolithophores are not silicified, some species exhibit a requirement for Si in the calcification process. These species also possess a novel protein (SITL) that resembles the SIT family of Si transporters found in diatoms. However, the nature of Si transport in coccolithophores is not yet known, making it difficult to determine the wider role of Si in coccolithophore biology. Here, we show that coccolithophore SITLs act as Na+‐coupled Si transporters when expressed in heterologous systems and exhibit similar characteristics to diatom SITs. We find that CbSITL from Coccolithus braarudii is transcriptionally regulated by Si availability and is expressed in environmental coccolithophore populations. However, the Si requirement of C. braarudii and other coccolithophores is very low, with transport rates of exogenous Si below the level of detection in sensitive assays of Si transport. As coccoliths contain only low levels of Si, we propose that Si acts to support the calcification process, rather than forming a structural component of the coccolith itself. Si is therefore acting as a micronutrient in coccolithophores and natural populations are only likely to experience Si limitation in circumstances where dissolved silicon (DSi) is depleted to extreme levels. [ABSTRACT FROM AUTHOR]

Coccolithophores are major producers of ocean biogenic calcite, but this process is predicted to be negatively affected by future ocean acidification scenarios. Since coccolithophores calcify intracellularly, the mechanisms through which changes in seawater carbonate chemistry affect calcification remain unclear. Here we show that voltagegated H+ channels in the plasma membrane of Coccolithus braarudii serve to regulate pH and maintain calcification under normal conditions but have greatly reduced activity in cells acclimated to low pH. This disrupts intracellular pH homeostasis and impairs the ability of C. braarudii to remove H+ generated by the calcification process, leading to specific coccolith malformations. These coccolith malformations can be reproduced by pharmacological inhibition of H+ channels. Heavily calcified coccolithophore species such as C. braarudii, which make the major contribution to carbonate export to the deep ocean, have a large intracellular H+ load and are likely to be most vulnerable to future decreases in ocean pH. [ABSTRACT FROM AUTHOR]

The marine calcifying algae Emiliania huxleyi (coccolithophores) was grown in laboratory cultures under varying conditions with respect to the environmental parameters of temperature and carbonate ion concentration [C[O.sub.3.sup.2-]] concentration. The Ca isotope composition of E. huxleyi's coccoliths reveals new insights into fractionation processes during biomineralization. The temperature-dependent Ca isotope fractionation resembles previous calibrations of inorganic and biogenic calcite and aragonite. Unlike inorganically precipitated calcite, the [C[O.sub.3.sup.2-]] concentration of the medium has no significant effect on the Ca isotope composition of the coccoliths. These results indicate a decoupling of the chemical properties of the bulk medium and the calcifying vesicle. Cellular Ca pathways of E. huxleyi indicate that fractionation cannot occur at the crystal surface, as occurs during inorganic precipitation. The dominant processes leading to the observed Ca isotope fractionation pattern in E. huxleyi are most likely the dehydration of the Ca aquocomplex at the plasma membrane and the attachment of dissolved Ca to proteins of Ca channels. The independence of Ca isotope fractionation from [C[O.sub.3.sup.2-]] and the small temperature dependence of E. huxleyi are also important for defining the isotopic signature of the oceanic Ca sink. Since coccolithuphores contribute to about half the global CaC[O.sub.3] production, a relatively uniform isotopic composition of the oceanic Ca sink is further supported. Keywords: Emiliania huxleyi, calcium isotopes, coccolithophores, isotope fractionation.

In recent studies the Sr/Ca ratio of coccolithophore calcite was used as a proxy for past coccolithophore growth and calcification rates. Since Sr and Ca concentrations in seawater have not remained constant through time, interpretation of St/Ca data from the coccolith-dominated sedimentary record requires knowledge about the incorporation of seawater Sr into coccolith calcite during coccolithogenesis. Here we show that Sr/Ca of Emiliania huxleyi coccoliths is linearly related to seawater Sr/Ca, meaning that the Sr exchange coefficient does not change with changing seawater St/Ca. The exchange coefficient for Sr in this study. 0.39, is close to values presented in the literature and is high compared with values obtained by inorganic precipitation experiments. This suggests a strong effect of cell physiology on biogenic calcite precipitation in coccolithophores. We present a conceptual model, based on the translnembrane transport of Sr and Ca, which explains the offset.