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Coccolithophore is one type of phytoplankton that forms calcium carbonate plates called coccoliths, and is relatively diverse with about 200 species in the worldʼs oceans. Coccoliths accumulate in massive calcareous deposits on the sea floor that serve as a sedimentary buffer of ocean chemistry and are also a major long-term carbon storage that has a significant impact on the global carbon cycle and climate. Coccolithophores are also a major producer of dimethyl sulfide (DMS) that is the dominant precursor for cloud condensation nuclei in the marine atmosphere; DMS emissions decrease solar radiation due to increasing cloud cover. Coccolithophores play an important role in biogeochemical cycles in the ocean by influencing the oceanic uptake of atmospheric carbon dioxide, and producing calcium carbonate sediments and DMS. Coccolithophores often bloom on massive scales. Around the Japan archipelago, coccolithophore blooms have also been reported; a Gephyrocapsa oceanica bloom in Tokyo Bay and Sagami Bay (1995) and Hakata Bay (2004, 2007, 2008), and a Emiliania huxleyi bloom in Suruga Bay (2007). In mid-May 2020, a massive bloom of coccolithophores were observed in Sagami Bay, Japan, possibly for the first time in 25 years. The bloom was initially observed by the JAXA, GCOM-C ocean color remote sensing satellite “Shikisai” capturing a vast emerald green reflectance image in the bay. Researchers who received the report immediately conducted in situ observations, and found that coccolithophores caused the massive bloom. Coccolithophore is a common yet elusive group in the waters around Japan, and there has only been one report of a bloom observation in Sagami Bay. This study shows the characteristics of the coccolithophore bloom observed in Sagami Bay in 2020 and also reports on the marine environment during the bloom. Water temperature, salinity, nutrients, chlorophyll a, and cell density were collected from field samples. Satellite ocean color data was provided by “Shikisai”. Satellite images on May 13 and 17 in 2020 show that the bloom spread counterclockwise due to counterclockwise coastal currents in Sagami Bay. The species, Gephyrocapsa oceanica, was confirmed using light microscopy and a scanning electron microscope, and was the same species that occurred in 1995. Chlorophyll a concentration and cell density at the surface were 1.2 μg L-1 and 9.0×103 cells mL-1, respectively. NO2- and NO3-concentrations at the surface during the bloom were 0.10 μmol L-1 and 1.83 μmol L-1, respectively, which were relatively higher than previous years. The results suggest that one of the causes of coccolithophore bloom is the relatively high concentration of inorganic nitrogen. Previous studies of prokaryotic and eukaryotic community structure in the coastal areas of Sagami Bay using genetic analysis have reported occurrence of haptophytes and the occurrence of coccolithophores throughout the year. These results suggest that coccolithophores are frequently present in Sagami Bay and form massive blooms in a short time period if environmental conditions are favorable. Further investigation including culture experiments are needed to resolve the details and specific causes.

Beyond the pCO2 records provided by ice core measurements, the quantification of atmospheric CO2 concentrations and changes thereof relies on proxy data, the development of which represents a foremost challenge in paleoceanography. In the paleoceanographic toolbox, the coccolithophores occupy a notable place, as the magnitude of the carbon isotopic fractionation between ambient CO2 and a type of organic compounds that these photosynthetic microalgae synthesize (the alkenones) represents a relatively robust proxy to reconstruct past atmospheric CO2 concentrations during the Cenozoic. The isotopic composition of coeval calcite biominerals found in the sediments and also produced by the coccolithophores (the coccoliths) have been found to record an ambient CO2 signal through culture and sediment analyses. These studies have, however, not yet formalized a transfer function that quantitatively ties the isotopic composition of coccolith calcite to the concentrations of aqueous CO2 and, ultimately, to atmospheric CO2 levels. Here, we make use of a microseparation protocol to compare the isotopic response of two size-restricted coccolith assemblages from the North Atlantic to changes in surface ocean CO2 during Termination II (ca. 130–140 ka). Performing paired measurements of the isotopic composition (δ13C and δ18O) of relatively large and small coccoliths provides an isotopic offset that can be designated as a “differential vital effect”. We find that the evolution of this offset follows that of aqueous CO2 concentrations computed from the ice core CO2 curve and an independent temperature signal. We interpret this biogeochemical feature to be the result of converging carbon fixation strategies between large and small cells as the degree of carbon limitation for cellular growth decreases across the deglaciation. We are therefore able to outline a first-order trend between the coccolith differential vital effects and aqueous CO2 in the range of Quaternary CO2 concentrations. Although this study would benefit from further constraints on the other controls at play on coccolith geochemistry (growth rate, air–sea gas exchange, etc.), this test of the drivers of coccolith Δδ13C and Δδ18O in natural conditions is a new step in the development of a coccolith paleo-CO2 probe.

Carbonate dissolution and deposition in the deep ocean is a critical component of carbon cycle, but detailed pictures of carbonate dynamics below the Carbonate Compensation Depth (CCD) remains poorly understood. In this study, we present a ∼ 420 thousand years record of carbonate content below CCD in IODP Site U1433 at 4.4 km water depth in the South China Sea (SCS). Instead of carbonate-free sediment, we observed that the carbonate content in this deep site could reach as high as 40%. The dissolution-resistant biogenic carbonate, coccoliths, and stable isotope ratio of bulk carbonate were analyzed to trace the carbonate sources. The coccolith assemblage in IODP Site U1433 shows an unrealistic feature of high productivity, which indicates a lateral transport of coccoliths from the northern SCS to the central basin. Coccoliths' contribution to total carbonate was on average 30% and could be as high as ∼80%. Their contribution to carbonate increased at the expense of the dissolution of other biogenic carbonate, such as foraminifera, and decreased with the dilution of other terrestrial carbonate particles. According to the thickness of coccoliths, significant dissolution of coccoliths only happened when the carbonate content dropped below 3%. Marbles from Taiwan, with negative oxygen isotope ratios, and early diagenetic carbonates from carbonate platform nearby, with both negative carbon and oxygen isotope ratios, could be the other important sources besides the marine biogenic carbonate. With help of numerical simulations, we estimated that the lateral transport by turbidity currents could contribute up to 70–90% of the deep-sea sediment in the central basin of SCS. These new findings enrich our knowledge of the carbon fluxes from continental margins to the deep ocean as a key part of the carbon cycle., Marine Geology, 457, ISSN:0025-3227

The genus Palusphaera Lecal-Schlauder emend. R.E. Norris is a distinctive modern coccolithophore that accommodates monomorphic, monothecate coccospheres with one type of spine-bearing heterococcoliths. Having examined a set of scanning electron microscopy images from our collections, we were able to demonstrate that four distinct species of Palusphaera exist in the modern oceans, including the type species Palusphaera vandelii, an informally proposed form, Palusphaera sp. 1 (type robusta) that is herein taxonomically ratified as Palusphaera crosiae sp. nov., and a distinctive morphotype, which is formally described as Palusphaera bownii sp. nov. Biometric analyses and observations on the morphologies of numerous P. vandelii specimens revealed the existence of a further form, and its commonly distinguished coccolith morphotypes are therefore taxonomically separated from Palusphaera vandelii and established as a discrete species, Palusphaera probertii sp. nov.

Coccoliths are plates of biogenic calcium carbonate secreted by calcifying marine phytoplankton; annually these phytoplankton are responsible for exporting >1 billion tonnes (1015 g) of calcite to the deep ocean. Rapid and reliable methods for assessing the degree of calcification are technically challenging because the coccoliths are micron sized and contain picograms (pg) of calcite. Here we pioneer an opto‐eletrochemical acid titration of individual coccoliths which allows 3D reconstruction of each individual coccolith via in situ optical imaging enabling direct inference of the coccolith mass. Coccolith mass ranging from 2 to 400 pg are reported herein, evidencing both inter‐ and intra‐species variation over four different species. We foresee this scientific breakthrough, which is independent of knowledge regarding the species and calibration‐free, will allow continuous monitoring and reporting of the degree of coccolith calcification in the changing marine environment., Acid dissolution enables 3D image reconstruction of coccoliths, providing mass estimation in the range of 2 to 400 picograms.

Oceanic red beds (ORBs) are present in Upper Cretaceous and Danian deep-marine deposits in the Basque–Cantabrian Basin of northern Spain. The presence and regularity of the succession of marl–limestone couplets is exceptional based on the macroscopic, microscopic and geochemical evidence collected. Five types of marl–limestone couplets are identified based on the colour, and a high maximum sedimentation rate (3.6 cm ka–1 ) is noted. The oxidizing activity of deep, cold-water masses is indicated by the oxygen isotope signal in the lower–upper Maastrichtian and Danian sections and the presence of the boreal inoceramidSpyridoceramus tegulatus.In theory, the variation in colour from grey to greenish-yellow, purple and pink up to red tones correlates with the Fe2+/(Fe2++Fe3+) ratio. It is interpreted as the possible palaeoenvironmental transit of particles that sediment out slowly in oxic environments when they circulate through cooler, oxidizing water masses. The colour is considered to be a depositional feature, and hematite, detected by X-ray diffraction, is the main staining agent, without discarding the possible redistribution of previous oxyhydroxides passing to hematite as a final product. The cell filling of the foraminifer shells does not incorporate appreciable amounts of Fe and Mg during diagenesis. Bacterial activity is detected using scanning electron microscopy images, both in the coccolith debris and in the detrital micas, although there is uncertainty as to its importance in the staining process.

The localization of organic material within biominerals is central to developing biomineral formation mechanisms. Coccoliths, morphologically sophisticated calcite platelets of intracellularly calcifying coccolithophores, are not only eco-physiologically important, but also influence biogeochemical cycles through mass production. Despite their importance and over a century of research, the formation mechanism of coccoliths is still poorly understood. Crucial unsolved questions include the localization of organic material within coccoliths. In extracellular calcifiers the discovery of an organics-containing nano-structure within seemingly single crystals has led to the formulation of a two-step crystallization mechanism. Coccoliths are traditionally thought of as being formed by a different mechanism, but it is unclear whether coccolith crystals possess a nano-structure. Here we review the evidence for and against such a nano-structure. Current SXPD analyses suggest a nano-structure of some kind, while imaging methods (SEM, TEM, AFM) provide evidence against it. We suggest directions for future research which should help solve this puzzle. STATEMENT OF SIGNIFICANCE: Coccolithophores, unicellular calcifying algae, are important primary producers and contribute significantly to pelagic calcium carbonate export. Their calcite platelets, the coccoliths, are amongst the most sophisticated biomineral structures. Understanding the crystallization mechanism of coccolith crystals is not only central to coccolithophore cell biology but also lies at the heart of biomineralization research more generally. The crystallization mechanism of coccoliths has remained largely elusive, not least because it is still an open question whether the micron sized coccolith crystals are pure calcite, or contain organic material. Here we review the state of the art and suggest a way to solve this central problem.

Mesocosm experiments have been fundamental to investigate the effects of elevated CO2 and ocean acidification (OA) on planktic communities. However, few of these experiments have been conducted using naturally nutrient-limited waters and/or considering the combined effects of OA and ocean warming (OW). Coccolithophores are a group of calcifying phytoplankton that can reach high abundances in the Mediterranean Sea, and whose responses to OA are modulated by temperature and nutrients. We present the results of the first land-based mesocosm experiment testing the effects of combined OA and OW on an oligotrophic Eastern Mediterranean coccolithophore community. Coccolithophore cell abundance drastically decreased under OW and combined OA and OW (greenhouse, GH) conditions. Emiliania huxleyi calcite mass decreased consistently only in the GH treatment; moreover, anomalous calcifications (i.e. coccolith malformations) were particularly common in the perturbed treatments, especially under OA. Overall, these data suggest that the projected increase in sea surface temperatures, including marine heatwaves, will cause rapid changes in Eastern Mediterranean coccolithophore communities, and that these effects will be exacerbated by OA.

The Southern Ocean is experiencing rapid and profound changes in its physical and biogeochemical properties that may influence the distribution and composition of pelagic plankton communities. Coccolithophores are the most prolific carbonate-producing phytoplankton group playing an important role in Southern Ocean biogeochemical cycles. However, knowledge is scarce about the record of (sub-)fossil coccolith assemblages in the Southern Ocean, which are constituting invaluable indicators for paleoenvironmental reconstructions. This study investigates coccolith assemblages preserved in surface sediments of southernmost Chile and across the Drake Passage that were retrieved during R/V Polarstern Expedition PS97. We focused on the coccolith response to steep environmental gradients across the frontal system of the Antarctic Circumpolar Current and to hydrodynamic and post-depositional processes occurring in this region. We used statistical analyses to explore which environmental parameters influenced the coccolith assemblages by means of Cluster and Redundancy Analyses. We specifically assessed the morphological diversity of the dominant taxa, i.e. Emiliania huxleyi, emphasizing biogeographical variability of morphotypes, coccolith sizes and calcite carbonate mass estimations. High coccolith abundances and species diversity compared to studies in the same area and in other sectors of the Southern Ocean occur, with a high species richness especially south of the Polar Front. While the surface sediments offshore Chile and north of the Polar Front provide suitable material to reconstruct overlying surface ocean conditions, further factors such as temporary thriving coccolithophore communities in the surface waters or transport of settling coccoliths via surface and bottom currents and eddies are influencing the (sub-)fossil coccolith assemblages south of the Polar Front. Additionally, deeper samples in the southern part of the study area are particularly affected by selective carbonate dissolution. We identified five E. huxleyi morphotypes (A, A overcalcified, R, B/C and O), and estimated coccolith carbonate masses on the basis of scanning electron microscope images. E. huxleyi morphologies reflect diverging biogeographical distributions, trending towards smaller and lighter coccoliths to the south and emphasizing the importance of documenting those morphologies in relation to changing environmental conditions to assess their response to projected environmental change in the SO., Biogeosciences Discussions, ISSN:1810-6277, ISSN:1810-6285

Altres ajuts: Unidad de excelencia María de Maeztu CEX2019-000940-M; Universidade de Vigo's programme to attract excellent research talent (RR04092017) The eastern Mediterranean Sea lies under the influence of high- and low-latitude climatic systems. The northern part of the basin is affected by Atlantic depressions and continental and polar air masses that promote intermediate and deep-water formation. The southern part is influenced by subtropical conditions and monsoon activity. Monsoon intensification results in enhanced freshwater discharge from the Nile River and other (now dry) systems along the North African margin. This freshwater influx into the Mediterranean Sea reduces surface water buoyancy loss. Disentangling the influences of these diverse climatic forcings is hindered by inherent proxy data limitations and by interactions between the climatic forcings. Here we use a wealth of published and new paleoclimate records across Termination II to understand the impacts of the higher latitude and subtropical/monsoon climate influences on coccolithophore ecology and holococcolith preservation in Aegean Sea sediment core LC21. We then use these findingsto interpret coccolith assemblage variations at Ocean Drilling Program Site 967 (located nearby LC21, at the Eratosthenes Seamount) during multiple glacial-interglacial cycles across the Middle Pleistocene (marine isotopic stages 14-9). The LC21 analysis suggests that holococcolith preservation was enhanced during Heinrich Stadial 11 (∼133 ka) and cold spell C26 (∼119 ka). These two events have been previously linked to cold conditions in the North Atlantic and Atlantic Meridional Overturning Circulation weakening. We propose that associated atmospheric perturbations over the Mediterranean Sea promoted deep-water formation, and thus holococcolith preservation. Similarly, in the Middle Pleistocene (MIS 14-9) of Site 967, we observe temporal coincidence between ten episodes of enhanced holococcolith preservation and episodes of Atlantic Meridional Overturning Circulation slowdown. In Site 967, we also identified repeated fluctuations in placoliths and in Florisphaera profunda, which indicate nutricline depth variations. The development of a deep chlorophyll maximum is associated with the North Africa and wet phases, as recently observed using elemental proxy records at Site 967, during the deposition of sapropel layers. A further deep chlorophyll maximum development is identified during MISs 12 and 10, as a result of pycnocline and nutricline shoaling within the lower part of the photic zone due to glacial sea-level lowering and water mass transport reduction at both the Gibraltar and Sicily Straits. Finally, enhanced holococcolith preservation during cold/dry events is clearly correlated to weakened monsoon activity in both Africa and Asia.

Coccolithophores are a diverse group of calcifying microalgae that have left a prominent fossil record on Earth. Various coccolithophore relics, both organic and inorganic, serve as proxies for reconstruction of past oceanic conditions.Emiliania huxleyi is the most widely distributed representative of the coccolithophores in modern oceans, and is known to engage in dynamic interactions with bacteria. Algal-bacterial interactions influence various aspects of algal physiology and alter algal alkenone unsaturation (UK’37), a frequently used organic coccolithophore-derived paleotemperature proxy. Whether algal-bacterial interactions influence inorganic coccolithophore-derived paleo-proxies, is yet unknown.A commonly used inorganic proxy for past productivity and sea surface temperature is the Sr/Ca ratio of the coccolith calcite. Interestingly, during interactions between bacteria and a population of calcifying algae, bacteria were shown to physically attach only to non-calcified algal cells, suggesting an influence on algal calcification.In this study we explore the effects of algal-bacterial interactions on calcification and coccolith Sr/Ca ratios. We find that while bacteria attach only to non-calcified algal cells, coccolith cell coverage and overall calcite production in algal populations with and without bacteria, is similar. Furthermore, we find that Sr/Ca values are impacted only by water temperature and algal growth rate, regardless of bacterial influences on algal physiology. Our observations reinforce the robustness of coccolith Sr/Ca ratios as a paleo-proxy independent of microbial interactions, and highlight a fundamental difference between organic and inorganic paleo-proxies.Summary StatementThe current research investigates the effect of microbial interactions on coccolith Sr/Ca ratio and overall calcification in the coccolithophore Emiliania huxleyi. We co-cultured E. huxleyi with the marine bacterium Phaeobacter inhibens and compared coccolith Sr/Ca between different growth stages in a range of temperatures. Our results indicate that coccolith Sr/Ca depends on temperature and algal growth rate, and remains robust despite significant bacterial influences on algal physiology.

Pleurochrysis genus algae are widely distributed in ocean waters. Pleurochrysis sp. algae are popularly known for its coccolithophores. Calcium carbonate (CaCO3) shells are major components of the coccolithophore, and they are key absorbers of carbondioxide. In this study, we have reported the effects of potassium nitrate (KNO3) concentration on calcium accumulation and total lipid, carbohydrate and protein contents of Pleurochrysis dentata. Results obtained from complexometric titration and scanning electron microscopy analysis showed higher rates of CaCO3 accumulation on Pleurochrysis dentata cell surface. We have also observed that overall cell size of Pleurochrysis dentata reached maximum when it was cultured at 0.75 mmol L-1 of KNO3. During 10 days of Pleurochrysis dentata culture total lipids and carbohydrate contents decreased, with slightly increased protein content. Results obtained from Fourier-Transform Infrared Spectroscopy (FTIR) also reported an increase in protein and decrease in lipids and carbohydrate contents, respectively. Similarly, Pleurochrysis dentata cultured at 1 mmol L-1 concentration of KNO3 exhibited the lowest carbohydrate (21.08%) and highest protein (32.87%) contents. Interestingly, Pleurochrysis dentata cultured without KNO3 exhibited 33.61% of total lipid content which reduced to a total lipid content of 13.67% when cultured at 1 mmol L-1 concentration of KNO3. Thus, culture medium containing higher than 1 mmol L-1 of KNO3 could inhibit the cell size of Pleurochrysis dentata and CaCO3 accumulation in shells but it could promote its cell growth. For the first time we have reported a relatively complete coccolith structure devoid of its protoplast. In this study, we have also described about the special planar structure of Pleurochrysis dentata CaCO3 shells present on its inner tube of the R unit and parallel to the outer tube of the V unit which we named it as "doornail structure". We believe that this doornail structure provides structural stability and support to the developing coccoliths in Pleurochrysis dentata. Also, we have discussed about the "double-disc" structure of coccoliths which are closely arranged and interlocked with each other. The double-disc structure ensures fixation of each coccolith and objecting its free horizontal movement and helps in attaining a complementary coccolith structure.

Organic-rich layers (sapropels), preserved in eastern Mediterranean marine sediment records, represent pronounced perturbations to thermohaline circulation and environmental conditions in the basin, in response to enhanced African monsoon activity and subsequent massive freshwater discharge. During the most recent event, Sapropel S1 formed between 10.8 and 6.1 ka, when freshwater-driven stratification caused seafloor anoxia below ~1,800-m depth, as a result of both failure of deep water formation and enhanced productivity. Here we analyze coccolith assemblages from the open eastern Mediterranean that form a west-east transect across the basin and provide insights on past environmental changes. We focus on holococcoliths, which are specifically produced by coccolithophores as part of their life cycle during the haploid phase. Since holococcolith calcification is characterized by nanocrystals highly susceptible to dissolution, we are testing their potential preservation under different bottom environmental conditions, including the effect of postdepositional oxidation. A comparison with benthic foraminifera assemblages in a core recovered close to Lybia reveals that holococcolith preservation is enhanced during seafloor reventilation and benthic foraminiferal repopulation in the middle to upper part of the record, before the actual sapropel termination. There are two such events of improved deep-water oxygenation in the Aegean and Adriatic Seas at 8.2 and 7.4 ka. The latter episode marks the onset of the transition to restored circulation in the eastern Mediterranean Sea, due to resumption of deep-water formation in the southern Aegean Sea and the conclusion of enhanced biogenic productivity.

Coccolithophores produce the bulk of ocean biogenic calcium carbonate 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 voltage-gated 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.

Coccolithophores are among the most important calcifying pelagic organisms. To assess how coccolithophore species with different coccolith-carbonate mass and distinct ecological resilience to ocean warming will influence the “rain ratio” and the “biological carbon pump”, 1 yr of species-specific coccolith-carbonate export fluxes were quantified using sediment traps moored at four sites between NW Africa and the Caribbean (i.e., CB-20°N/21°W, at 1214 m; M1-12°N/23°W, at 1150 m; M2-14°N/37°W, at 1235 m; M4-12°N/49°W, at 1130 m). Highest coccolith-CaCO3 fluxes at the westernmost site M4, where the nutricline is deepest along the tropical North Atlantic, were dominated by deep-dwelling small-sized coccolithspecies Florisphaera profunda and Gladiolithus flabellatus. Total coccolith-CaCO3 fluxesof 371 mg m−2 yr−1 at M4 were followed by 165 mg m−2 yr−1 at the north-easternmost CB, 130 mg m−2 yr−1 at M1, and 114 mg m−2 yr −1 at M2 in between. Coccoliths accounted for nearly half of the total carbonate flux at M4 (45%), much higher compared to 23% at M2 and 15% at M1 and CB. At site M4, highest ratios of coccolith-CaCO3 to particulate organic carbon fluxes and weak correlations between the carbonate of deep-dwelling species and particulate organic carbon suggest that increasing productivity in the lower photic zone in response to ocean warming might enhance the rain ratio and reduce the coccolith-ballasting efficiency. The resulting weakened biological carbon pump could, however, be counterbalanced by increasing frequency of Saharan dust outbreaks across the tropical Atlantic, providing mineral ballast as well as nutrients to fuel fast-blooming and ballast-efficient coccolithophore species.

International audience; Heterococcoliths are calcite platelets produced inside diploid coccolithophore cells and extruded to form a covering on the cell surface called a coccosphere. The size of coccoliths is an important parameter sometimes used to identify species, and it is observed to be influenced in extant species by abiotic parameters (e.g., CO2, light). However, the variable distribution of coccolith sizes occurring within a single coccosphere questions the mechanisms controlling coccolith size. A relationship between cell/coccosphere size and mean coccolith size was previously identified, called the “coccolithophore size rules”. In this study, we query the mechanisms controlling the size of a coccolith during coccolithogenesis. A culture experiment on Gephyrocapsa huxleyi strain RCC1216 shows that coccolithogenesis occurs during the cell growth G1 interphase and newly produced coccoliths get bigger as the cell grows. These observations provide parameters for the development of two numerical models used to simulate the coccolith size distribution within a coccolithophore population. Neither model can accurately reproduce an empirical monoclonal coccolith size distribution, indicating that additional factors influence coccolith size. According to our results, coccolith size is only clearly related to cell size at the time of its formation. We confirm that application of the coccolithophore size rules model should be limited to inferring average cell dimensions from (fossil) coccolith biometry, and that comparisons are valid only in multipopulational studies. The coccolith size rule model – the constraining effect of coccolith production during G1 interphase cell growth on coccolith size – proposed here is applicable only for some placolith-forming species.

Coccolithophores are one of the major contributors to the pelagic production of calcium carbonate and their fossilized remains are a key component of the biogeochemical cycles of calcium (Ca), magnesium (Mg), and other divalent cations present in the intracellular precipitated calcitic structures (coccoliths). The geochemical signature of coccoliths (e.g., Sr/Ca and Mg/Ca ratios) is used as paleoproxy to reconstruct past environmental conditions and to understand the underlying physiological precipitation kinetics. Here, we present the elemental fractionation of Sr and Mg in calcite of the coccolithophore Gephyrocapsa oceanica from controlled laboratory experiments applying an extended temperature gradient (12 to 27°C). The physiological reaction norm of G. oceanica, in terms of growth rate, exhibited optimum behavior while the partition coefficient of Sr (DSr) was linearly correlated with temperature and DMg indicated no specific trend. Our results indicate: (1) a presumably secondary physiological control of DSr, and (2) the importance of calibrating coccolithophore-based proxies using experiments that include the full physiological reaction norms (i.e., a possible non-linear response) to environmental drivers (e.g., temperature, salinity, and pH, etc.). The presented results contribute to an improved understanding of the underlying physiological kinetics involved in regulating coccolith elemental fractionation and give additional implications for designing future laboratory experiments to calibrate and apply coccolithophore based paleoproxies on the fossil sediment record.

International audience; While numerous studies have highlighted the central role of Southern Ocean (SO) dynamics in modulating rapid increases in atmospheric CO2 concentrations during deglaciations, fewer studies have yet focused on the impact of the Biological Carbon Pump - and more specifically the Carbonate Counter Pump (CCP) - in contributing to increase the CO2 concentration in oceanic surface waters and thus, in the atmosphere. Here, we present micropaleontological (coccolith, planktonic foraminifera) and geochemical (CaCO3, CaXRF, δ13CN. pachyderma) constraints from sediment core MD04-2718 retrieved in the Polar Front Zone of the Indian Ocean covering the time interval spanning Marine Isotope Stage (MIS) 12 to MIS 10 (440,000-360,000 years). We compare our results with published records from the SO to reconstruct past changes in CCP and upwelling dynamics and understand their leverage on the ocean-atmosphere portioning of CO2. We demonstrate that the sharp increase in atmospheric pCO2 during Termination V was likely associated with enhanced deep-water ventilation in the SO, that promoted the release of previously sequestered CO2 to the ocean surface as the westerly wind belt and the frontal system migrated southwards. Enhanced CCP is observed later, during MIS 11, and is likely the consequence of higher sea surface temperature and higher nutrient availability due to the reinvigoration of SO upwelling leading to increased coccolith (and to a lesser degree, planktonic foraminifera) production and export. The low eccentricity signal recorded during MIS 11 might have additionally strengthened the CCP, exerting a specific control on Gephyrocapsa morphotypes. In addition to the strong global biological productivity and higher carbon storage on land, these synergistic mechanisms may have permitted to shape the distinctive 30 ka-long pCO2 plateau characteristic of MIS 11.

International audience; The development of calcification by the coccolithophores had a profound impact on ocean carbon cycling, but the evolutionary steps leading to the formation of these complex biomineralized structures are not clear. Heterococcoliths consisting of intricately shaped calcite crystals are formed intracellularly by the diploid life cycle phase. Holococcoliths consisting of simple rhombic crystals can be produced by the haploid life cycle stage but are thought to be formed extracellularly, representing an independent evolutionary origin of calcification. We use advanced microscopy techniques to determine the nature of coccolith formation and complex crystal formation in coccolithophore life cycle stages. We find that holococcoliths are formed in intracellular compartments in a similar manner to heterococcoliths. However, we show that silicon is not required for holococcolith formation and that the requirement for silicon in certain coccolithophore species relates specifically to the process of crystal morphogenesis in heterococcoliths. We therefore propose an evolutionary scheme in which the lower complexity holococcoliths represent an ancestral form of calcification in coccolithophores. The subsequent recruitment of a silicon-dependent mechanism for crystal morphogenesis in the diploid life cycle stage led to the emergence of the intricately shaped heterococcoliths, enabling the formation of the elaborate coccospheres that underpin the ecological success of coccolithophores.

Coccolithophores represent a major component of the marine phytoplankton and contribute to the bulk of biogenic calcite formation on Earth. These unicellular protists produce minute calcite scales (coccoliths) within the cell, which are secreted to the cell surface. Individual coccoliths and their arrangements on the cell surface display a wide range of morphological variations. This review explores some of the recent evidence that points to similarities and differences in the mechanisms of calcification, focussing on the transport mechanisms that bring substrates to, and remove products from the site of calcification, together with new findings on factors that regulate coccolith morphology. We argue that better knowledge of these mechanisms and their variations is needed to inform more generally how different species of coccolithophore are likely to respond to changes in ocean chemistry. STATEMENT OF SIGNIFICANCE: Coccolithophores, minute single celled phytoplankton are the major producers of biogenic carbonate on Earth. They also represent an important component of the ocean's biota and contribute significantly to global carbon fluxes. Coccolithophores produce intricate calcite scales (coccoliths) internally that they secrete onto their external surface. This review presents some recent key findings on the mechanisms underlying the production of coccoliths. It also considers the factors that regulate the rate of production as well as the variety of shapes of individual coccoliths and their arrangements at the cell surface. Understanding these processes is needed to allow better predictions of how coccolithophores may respond to changing ocean chemistry associated with climate change.

The Early Eocene Thermal Maximum 2 (ETM2) at ~53.7 Ma is one of multiple hyperthermal events that followed the Paleocene-Eocene Thermal Maximum (PETM, ~56 Ma). The negative carbon excursion and deep ocean carbonate dissolution which occurred during the event imply that a substantial amount (103 Gt) of carbon (C) was added to the ocean-atmosphere system, consequently increasing atmospheric CO2(pCO2). This makes the event relevant to the current scenario of anthropogenic CO2 additions and global change. Resulting changes in ocean stratification and pH, as well as changes in exogenic cycles which supply nutrients to the ocean, may have affected the productivity of marine phytoplankton, especially calcifying phytoplankton. Changes in productivity, in turn, may affect the rate of sequestration of excess CO2 in the deep ocean and sediments. In order to reconstruct the productivity response by calcareous nannoplankton to ETM2 in the South Atlantic (Site 1265) and North Pacific (Site 1209), we employ the coccolith Sr/Ca productivity proxy with analysis of well-preserved picked monogeneric populations by ion probe supplemented by analysis of various size fractions of nannofossil sediments by ICP-AES. The former technique of measuring Sr/Ca in selected nannofossil populations using the ion probe circumvents possible contamination with secondary calcite. Avoiding such contamination is important for an accurate interpretation of the nannoplankton productivity record, since diagenetic processes can bias the productivity signal, as we demonstrate for Sr/Ca measurements in the fine (

Coccolithophores are a group of phytoplankton widely distributed in the ocean, which secrete extracellular calcite plates termed coccoliths. Coccoliths have been increasingly employed as an archive for geochemical, ecological and paleoclimate studies in recent years. A robust application of coccolith-based geochemical proxies relies on understanding the carbon acquisition strategies and the pathways of carbon supply for calcification. Carbonic anhydrase (CA) plays important roles in the carbon concentrating mechanism s of aquatic algae and potentially also in calcification. However, it is difficult to independently assess the role of CA in carbon supply for photosynthesis versus calcification. To fill this gap, we explored a new method to detect the CA activity inside coccolithophore. To achieve this, coccolithophores were cultured with oxygen and carbon isotope labeled dissolved inorganic carbon (DIC). By exploiting the different behavior of oxygen and carbon isotopes with (sea)water, this double label method can elucidate the significance of CA activity in the calcification pathway. Application of this method to Emiliania huxleyi shows that CA is present in the calcification pathway, and that there is no significant difference in the CA activity between a high and low CO2 treatment. However, under low CO2 treatment E. huxleyi enhanced the bicarbonate pumping rate on both cell and chloroplast membranes. This novel method could be performed on other species of coccolithophores in the future and have a potential to extend our knowledge on coccolith oxygen isotope vital effects. (© 2020 Elsevier ), Geochimica et Cosmochimica Acta, 292, ISSN:0016-7037, ISSN:1872-9533

Culture experiments with coccolithophore algae—the dominant group of marine calcifying phytoplankton—imply a strong sensitivity in growth rate, degree of cellular calcification, and cell size to changes in the carbon chemistry of their growth environment. These results underpin recent studies that have explored how these physiological parameters have varied on geological time scales, in response to changing surface ocean habitats and the concentrations of carbon in the ocean-atmosphere system. Here, we add to this work with a study of reticulofenestrid coccolith size—the dominant coccolithophore family of the Cenozoic—over the Oligocene to Early Miocene time interval. We examine sediments from contrasting latitudes and regional environmental settings, comparing sites using coherent, updated age models to distinguish globally synchronous trends in cell size from regional trends. Our results confirm several changes in coccolith size—which is strongly correlated to cell size—that are globally reproducible within the ~1 Myr age uncertainty, including a reduction in mean size by >2 μm from 30.2 to 27 and 24.5 to 23 Ma, and then increase in mean size after 20 Ma. The main difference among regions is the presence/absence of coccoliths larger than 8 μm. We evaluate which scenarios of change in carbon dioxide, temperature, and nutrient availability could have exerted selective pressure on cell size for different size classes to produce the observed size trends at each studied site. These million-year scale adaptations of ancient coccolithophores contribute to the understanding of phytoplankton physiology in the transition to the modern “icehouse” world. ©2020. American Geophysical Union. All Rights Reserved. ISSN:2572-4525 ISSN:2572-4517

The effect of global climate cycles driven by Earth’s orbital variations on evolution is poorly understood because of difficulties achieving sufficiently-resolved records of past evolution. The fossil remains of coccolithophores, a key calcifying phytoplankton group, enable an exceptional assessment of the impact of cyclic orbital-scale climate change on evolution because of their abundance in marine sediments, and because coccolithophores demonstrate extreme morphological plasticity in response to the changing environment1,2. Recently, evolutionary genetic analyses linked broad changes in Pleistocene fossil coccolith morphology to species radiation events3. Here, using high-resolution coccolith data, we show that during the last 2.8 million years coccolithophore evolution was forced by Earth’s orbital eccentricity with rhythms of ~100,000 years and 405,000 years - a distinct spectral signature to that of coeval global climate cycles4. Simulations with an Earth System Model5 including the marine carbon cycle6 demonstrate that eccentricity directly impacts the diversity of ecological niches occurring over the annual cycle in the tropical ocean. Reduced seasonality favours species with mid-size coccoliths that bloom year-round, increasing coccolith carbonate export and burial. We posit that eccentricity pacing of phytoplankton evolution contributed to the strong 405,000-year pacing seen in records of the global carbon cycle. ; The effect of global climate cycles driven by Earth's orbital variations on evolution is poorly understood because of difficulties achieving 14 sufficiently-resolved records of past evolution. The fossil remains of coccolithophores, a key calcifying phytoplankton group, enable an 15 exceptional assessment of the impact of cyclic orbital-scale climate change on evolution because of their abundance in marine sediments, 16 and because coccolithophores demonstrate extreme morphological plasticity in response to the changing environment 1,2. Recently, 17 evolutionary genetic analyses linked broad changes in Pleistocene fossil coccolith morphology to species radiation events 3. Here, using 18 high-resolution coccolith data, we show that during the last 2.8 million years coccolithophore evolution was forced by Earth's orbital 19 eccentricity with rhythms of ~100,000 years and 405,000 years-a distinct spectral signature to that of coeval global climate cycles 4. 20 Simulations with an Earth System Model 5 including the marine carbon cycle 6 demonstrate that eccentricity directly impacts the diversity 21 of ecological niches occurring over the annual cycle in the tropical ocean. Reduced seasonality favours species with mid-size coccoliths 22 that bloom year-round, increasing coccolith carbonate export and burial. We posit that eccentricity pacing of phytoplankton evolution 23 contributed to the strong 405,000-year pacing seen in records of the global carbon cycle. 24 25 Coccolithophores are important producers of CaCO3 in the ocean and their fossil remains (calcite platelets called coccoliths) first appeared in 26 sediments during the late Triassic (~210 million years ago, Ma). Coccolithophores rose to dominance in the open ocean during the early 27 Cretaceous 7 and thereafter become a key biological modulator of the global carbon cycle, via both photosynthesis and calcification 8. In the 28 dominant Noëlaerhabdaceae family (including the cosmopolitan genera Emiliania, Gephyrocapsa and Reticulofenestra), species are defined by 29 the morphological characteristics of their coccoliths, with size being a key criterion 9. For Gephyrocapsa and Emiliania, phylogenies 30 reconstructed from gene sequence data indicate that morphology-based definitions correspond to biological species 3,10. Yet, within a given 31 Noëlaerhabdaceae population, interspecific and intraspecific changes in coccolith length and mass (the latter encoding degree of calcification as 32 well as size) occur in response to environmental parameters such as carbonate chemistry 1 and temperature 2. Existing studies of coccolithophore 33 evolution have focused on geological-timescale changes in species richness and turnover 11 , coccolith carbonate accumulation 7,12 , or calcification 34 potentially driven by carbon cycle changes 13. However, to date a lack of records that are both long-term and high-resolution has precluded an 35 understanding of the impact of orbital cycles (on timescales of tens to hundreds of thousands of years) on coccolithophore evolution and 36 carbonate production. 37 38