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Marine calcareous sediments provide a fundamental basis for palaeoceanographic studies aiming to reconstruct past oceanic conditions and understand key biogeochemical element cycles. Calcifying unicellular phytoplankton (coccolithophores) are a major contributor to both carbon and calcium cycling by photosynthesis and the production of calcite (coccoliths) in the euphotic zone, and the subsequent long-term deposition and burial into marine sediments. Here we present data from controlled laboratory experiments on four coccolithophore species and elucidate the relation between the divalent cation (Sr, Mg and Ca) partitioning in coccoliths and cellular physiology (growth, calcification and photosynthesis). Coccolithophores were cultured under different seawater temperature and carbonate chemistry conditions. The partition coefficient of strontium (DSr) was positively correlated with both carbon dioxide (pCO2) and temperature but displayed no coherent relation to particulate organic and inorganic carbon production rates. Furthermore, DSr correlated positively with cellular growth rates when driven by temperature but no correlation was present when changes in growth rates were pCO2-induced. Our results demonstrate the complex interaction between environmental forcing and physiological control on the strontium partitioning in coccolithophore calcite and challenge interpretations of the coccolith Sr / Ca ratio from high-pCO2 environments (e.g. Palaeocene–Eocene thermal maximum). The partition coefficient of magnesium (DMg) displayed species-specific differences and elevated values under nutrient limitation. No conclusive correlation between coccolith DMg and temperature was observed but pCO2 induced a rising trend in coccolith DMg. Interestingly, the best correlation was found between coccolith DMg and chlorophyll a production, suggesting that chlorophyll a and calcite associated Mg originate from the same intracellular pool. These and previous findings indicate that Mg is transported into the cell and to the site of calcification via different pathways than Ca and Sr. Consequently, the coccolith Mg / Ca ratio should be decoupled from the seawater Mg / Ca ratio. This study gives an extended insight into the driving factors influencing the coccolith Mg / Ca ratio and should be considered for future palaeoproxy calibrations.

Availability of phosphate for phytoplankton and bacteria and of glucose for bacteria at different pCO2 levels were studied in a mesocosm experiment (PeECE III). Using nutrient-depleted SW Norwegian fjord waters, three different levels of pCO2 (350 μatm: 1×CO2; 700 μatm: 2×CO2; 1050 μatm: 3×CO2) were set up, and nitrate and phosphate were added at the start of the experiment in order to induce a phytoplankton bloom. Despite similar responses of total particulate P concentration and phosphate turnover time at the three different pCO2 levels, the size distribution of particulate P and 33PO4 uptake suggested that phosphate transferred to the >10 μm fraction was greater in the 3×CO2 mesocosm during the first 6–10 days when phosphate concentration was high. During the period of phosphate depletion (after Day 12), specific phosphate affinity and specific alkaline phosphatase activity (APA) suggested a P-deficiency (i.e. suboptimal phosphate supply) rather than a P-limitation for the phytoplankton and bacterial community at the three different pCO2 levels. Specific phosphate affinity and specific APA tended to be higher in the 3×CO2 than in the 2×CO2 and 1×CO2 mesocosms during the phosphate depletion period, although no statistical differences were found. Glucose turnover time was correlated significantly and negatively with bacterial abundance and production but not with the bulk DOC concentration. This suggests that even though constituting a small fraction of the bulk DOC, glucose was an important component of labile DOC for bacteria. Specific glucose affinity of bacteria behaved similarly at the three different pCO2 levels with measured specific glucose affinities being consistently much lower than the theoretical maximum predicted from the diffusion-limited model. This suggests that bacterial growth was not severely limited by the glucose availability. Hence, it seems that the lower availability of inorganic nutrients after the phytoplankton bloom reduced the bacterial capacity to consume labile DOC in the upper mixed layer of the stratified mesocosms.

The coccolithophore Emiliania huxleyi is a marine phytoplankton species capable of forming small calcium carbonate scales (coccoliths) which cover the organic part of the cell. Calcification rates of E. huxleyi are known to be sensitive to changes in seawater carbonate chemistry. It has, however, not yet been clearly determined how these changes are reflected in size and weight of individual coccoliths and which specific parameter(s) of the carbonate system drive morphological modifications. Here, we compare data on coccolith size, weight, and malformation from a set of five experiments with a large diversity of carbonate chemistry conditions. This diversity allows distinguishing the influence of individual carbonate chemistry parameters such as carbon dioxide (CO2), bicarbonate (HCO3−), carbonate ion (CO32−), and protons (H+) on the measured parameters. Measurements of fine-scale morphological structures reveal an increase of coccolith malformation with decreasing pH suggesting that H+ is the major factor causing malformations. Coccolith distal shield area varies from about 5 to 11 μm2. Changes in size seem to be mainly induced by varying [HCO3−] and [H+] although influence of [CO32−] cannot be entirely ruled out. Changes in coccolith weight were proportional to changes in size. Increasing CaCO3 production rates are reflected in an increase in coccolith weight and an increase of the number of coccoliths formed per unit time. The combined investigation of morphological features and coccolith production rates presented in this study may help to interpret data derived from sediment cores, where coccolith morphology is used to reconstruct calcification rates in the water column.

Ocean acidification and associated shifts in carbonate chemistry speciation induced by increasing levels of atmospheric carbon dioxide (CO2) have the potential to impact marine biota in various ways. The process of biogenic calcification, for instance, is usually shown to be negatively affected. In coccolithophores, an important group of pelagic calcifiers, changes in cellular calcification rates in response to changing ocean carbonate chemistry appear to differ among species. By applying a wider CO2 range we show that a species previously reported insensitive to seawater acidification, Coccolithus braarudii, responds both in terms of calcification and photosynthesis, although at higher levels of CO2. Thus, observed differences between species seem to be related to individual sensitivities while the underlying mechanisms could be the same. On this basis we develop a conceptual model of coccolithophorid calcification and photosynthesis in response to CO2-induced changes in seawater carbonate chemistry speciation.

The effect of ocean acidification on the balance between gross community production (GCP) and community respiration (CR) (i.e., net community production, NCP) of plankton communities was investigated in summer 2010 in Kongsfjorden, west of Svalbard. Surface water, which was characterized by low concentrations of dissolved inorganic nutrients and chlorophyll a (a proxy of phytoplankton biomass), was enclosed in nine mesocosms and subjected to eight pCO2 levels (two replicated controls and seven enhanced pCO2 treatments) for one month. Nutrients were added to all mesocosms on day 13 of the experiment, and thereafter increase of chlorophyll a was provoked in all mesocosms. No clear trend in response to increasing pCO2 was found in the daily values of NCP, CR, and GCP. For further analysis, these parameters were cumulated for the following three periods: phase 1 – end of CO2 manipulation until nutrient addition (t4 to t13); phase 2 – nutrient addition until the second chlorophyll a minimum (t14 to t21); phase 3 – the second chlorophyll a minimum until the end of this study (t22 to t28). A significant response was detected as a decrease of NCP with increasing pCO2 during phase 3. CR was relatively stable throughout the experiment in all mesocosms. As a result, the cumulative GCP significantly decreased with increasing pCO2 during phase 3. After the nutrient addition, the ratios of cumulative NCP to cumulative consumption of NO3 and PO4 showed a significant decrease during phase 3 with increasing pCO2. The results suggest that elevated pCO2 influenced cumulative NCP and stoichiometric C and nutrient coupling of the plankton community in a high-latitude fjord only for a limited period. However provided that there were some differences or weak correlations between NCP data based on different methods in the same experiment, this conclusion should be taken with caution.

The influence of seawater carbon dioxide (CO2) concentration on the size distribution of suspended particles (2–60 μm) and on phytoplankton abundance was investigated during a mesocosm experiment at the large scale facility (LFS) in Bergen, Norway, in the frame of the Pelagic Ecosystem CO2 Enrichment study (PeECE II). In nine outdoor enclosures the partial pressure of CO2 in seawater was modified by an aeration system to simulate past (~190 μatm CO2), present (~370 μatm CO2) and future (~700 μatm CO2) CO2 conditions in triplicates. Due to the initial addition of inorganic nutrients, phytoplankton blooms developed in all mesocosms and were monitored over a period of 19 days. Seawater samples were collected daily for analysing the abundance of suspended particles and phytoplankton with the Coulter Counter and with Flow Cytometry, respectively. During the bloom period, the abundance of small particles (2 levels. At that time, a direct relationship between the total-surface-to-total-volume ratio of suspended particles and DIC concentration was determined for all mesocosms. Significant changes with respect to the CO2 treatment were also observed in the phytoplankton community structure. While some populations such as diatoms seemed to be insensitive to the CO2 treatment, others like Micromonas spp. increased with CO2, or showed maximum abundance at present day CO2 (i.e. Emiliania huxleyi). The strongest response to CO2 was observed in the abundance of small autotrophic nano-plankton that strongly increased during the bloom in the past CO2 mesocosms. Together, changes in particle size distribution and phytoplankton community indicate a complex interplay between the ability of the cells to physiologically respond to changes in CO2 and size selection. Size of cells is of general importance for a variety of processes in marine systems such as diffusion-limited uptake of substrates, resource allocation, predator-prey interaction, and gravitational settling. The observed changes in particle size distribution are therefore discussed with respect to biogeochemical cycling and ecosystem functioning.

The primary impacts of anthropogenic CO2 emissions on marine biogeochemical cycles predicted so far include ocean acidification, global warming induced shifts in biogeographical provinces, and a possible negative feedback on atmospheric CO2 levels by CO2‐fertilized biological production. Here we report a new potentially significant impact on the oxygen‐minimum zones of the tropical oceans. Using a model of global climate, ocean circulation, and biogeochemical cycling, we extrapolate mesocosm‐derived experimental findings of a pCO2‐sensitive increase in biotic carbon‐to‐nitrogen drawdown to the global ocean. For a simulation run from the onset of the industrial revolution until A.D. 2100 under a “business‐as‐usual” scenario for anthropogenic CO2 emissions, our model predicts a negative feedback on atmospheric CO2 levels, which amounts to 34 Gt C by the end of this century. While this represents a small alteration of the anthropogenic perturbation of the carbon cycle, the model results reveal a dramatic 50% increase in the suboxic water volume by the end of this century in response to the respiration of excess organic carbon formed at higher CO2 levels. This is a significant expansion of the marine “dead zones” with severe implications not only for all higher life forms but also for oxygen‐sensitive nutrient recycling and, hence, for oceanic nutrient inventories.

International audience; Availability of phosphate for phytoplankton and bacteria and of labile organic carbon for bacteria at different pCO2 levels were studied in a mesocosm experiment (PeECE III). Using nutrient-depleted SW Norwegian fjord waters, three different levels of pCO2 (350 ?atm: 1×CO2; 750 ?atm: 2×CO2; 1050 ?atm: 3×CO2) were set up, and nitrate and phosphate were added at the start of the experiment in order to induce a phytoplankton bloom. Despite similar responses of total particulate P concentration and phosphate turnover time at the three different pCO2 levels, the size distribution of particulate P and 33PO4 uptake suggested that phosphate transferred to the >10 ?m fraction was greater in the 3×CO2 mesocosm during the first 6?10 days when phosphate concentration was high. During the period of phosphate depletion (after Day 12), specific phosphate affinity and specific alkaline phosphatase activity (APA) suggested a P-deficiency (i.e. suboptimal phosphate supply) but not a P-limitation for the phytoplankton and bacterial community at the three different pCO2 levels. Although specific phosphate affinity and specific APA tended to be higher in 3×CO2 than in 2×CO2 and 1×CO2 mesocosms during the phosphate depletion period, no statistical differences were found. Responses of specific glucose affinity for bacteria were similar at the three different pCO2 levels. Measured specific glucose affinities were consistently much lower than the theoretical maximum predicted from the diffusion-limited model, suggesting that bacterial growth was not limited by the availability of labile dissolved organic carbon. These results suggest that availability of phosphate and glucose was similar at the three different pCO2 levels.