Seawater carbonate chemistry, POC, PIC, TPC, SPM, N, TEP and growth rate during experiments with coccolithophores Emiliania huxleyi (AC472), Calcidiscus leptoporus (AC370) and Syracosphaera pulchra (AC418) during experiments, 2011

The response of three coccolithophores (Emiliania huxleyi, Calcidiscus leptoporus and Syracosphaera pulchra) to elevated partial pressure (pCO2) of carbon dioxide was investigated in batch cultures. For the first time, we also report on the response of the non calcifying (haploid) life stage of these three species. The growth rate, cell size, inorganic (PIC) and organic carbon (POC) of both life stages were measured at two different pCO2 (400and 760 ppm) and their organic and inorganic carbon production calculated. The two lifestages within the same species generally exhibited a similar response to elevated pCO2, theresponse of the haploid stage being often more pronounced than that of the diploid stage. Thegrowth rate was consistently higher at higher pCO2 but the response of other processes varied among species. The calcification rate of C. leptoporus and of S. pulchra did not change at elevated pCO2 while increased in E. huxleyi. The POC production as well as the cell size of both life stages of S. pulchra and of the haploid stage of E. huxleyi markedly decreased at elevated pCO2. It remained unaltered in the diploid stage of E. huxleyi and C. leptoporus and increased in the haploid stage of the latter. The PIC:POC ratio increased in E. huxleyi and was constant in C. leptoporus and S. pulchra. These results suggest that the non-calcifying stage, is more responsive than the calcifying stage and that the most versatile genera will proliferate in a more acidic ocean rather than all coccolithophores will decline.

Ocean acidification alleviates low-temperature effects on growth and photosynthesis of the red alga Neosiphonia harveyi (Rhodophyta)

This study aimed to examine interactive effects between ocean acidification and temperature on the photosynthetic and growth performance of Neosiphonia harveyi. N. harveyi was cultivated at 10 and 17.5 °C at present (~380 µatm), expected future (~800 µatm), and high (~1500 µatm) pCO2. Chlorophyll a fluorescence, net photosynthesis, and growth were measured. The state of the carbon-concentrating mechanism (CCM) was examined by pH-drift experiments (with algae cultivated at 10 °C only) using ethoxyzolamide, an inhibitor of external and internal carbonic anhydrases (exCA and intCA, respectively). Furthermore, the inhibitory effect of acetazolamide (an inhibitor of exCA) and Tris (an inhibitor of the acidification of the diffusive boundary layer) on net photosynthesis was measured at both temperatures. Temperature affected photosynthesis (in terms of photosynthetic efficiency, light saturation point, and net photosynthesis) and growth at present pCO2, but these effects decreased with increasing pCO2. The relevance of the CCM decreased at 10 °C. A pCO2 effect on the CCM could only be shown if intCA and exCA were inhibited. The experiments demonstrate for the first time interactions between ocean acidification and temperature on the performance of a non-calcifying macroalga and show that the effects of low temperature on photosynthesis can be alleviated by increasing pCO2. The findings indicate that the carbon acquisition mediated by exCA and acidification of the diffusive boundary layer decrease at low temperatures but are not affected by the cultivation level of pCO2, whereas the activity of intCA is affected by pCO2. Ecologically, the findings suggest that ocean acidification might affect the biogeographical distribution of N. harveyi.

Growth, respiration and photophysiology of coral massive Porites spp. in the experiment of Moorea

I tested the hypothesis that high pCO2 (76.6 Pa and 87.2 Pa vs. 42.9 Pa) has no effect on the metabolism of juvenile massive Porites spp. after 11 days at 28 °C and 545 µmol quanta/m**2/s. The response was assessed as aerobic dark respiration, skeletal weight (i.e., calcification), biomass, and chlorophyll fluorescence. Corals were collected from the shallow (3-4 m) back reef of Moorea, French Polynesia (17°28.614'S, 149°48.917'W), and experiments conducted during April and May 2011. An increase in pCO2 to 76.6 Pa had no effect on any dependent variable, but 87.2 Pa pCO2 reduced area-normalized (but not biomass-normalized) respiration 36 %, as well as maximum photochemical efficiency (Fv/Fm) of open RCIIs and effective photochemical efficiency of RCIIs in actinic light (Delta F/F'm ); neither biomass, calcification, nor the energy expenditure coincident with calcification (J/g) was effected. These results do not support the hypothesis that high pCO2 reduces coral calcification through increased metabolic costs and, instead, suggest that high pCO2 causes metabolic depression and photochemical impairment similar to that associated with bleaching. Evidence of a pCO2 threshold between 76.6 and 87.2 Pa for inhibitory effects on respiration and photochemistry deserves further attention as it might signal the presence of unpredictable effects of rising pCO2.

Seawater carbonate chemistry, stable carbon isotope fractionation and growth rate during experiments with marine phytoplankton community, 1999

Stable carbon isotope fractionation (%) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (pi), defined as the rate of carbon fixation during the photo period, when cultivated at 12:12 h. 16:8 h, or 186 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6% lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in pi. In contrast, E, in Phaeodactylum tn'cornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group 11) was largely insensitive to day length-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, pi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between E, and pi/[C02,,,] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [COz,,,], on isotopic fractionation. A decrease in [CO2,,,] from ca 26 to 3 pm01 kg-' caused a decrease in E, by less than 3%0 This indicates that variation in h in response to changes in day length has a similar or even greater effect on isotopic fractionation than [COz,,,] m some of the species tested. In both groups E, tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by l3C depletion in the 2 h before beginning of the following Light period. The daily amplitude of the algal isotopic composition (613C), however, was <1.5%0, which demonstrates that diurnal variation in Fl3C is relatively small.

Seawater carbonate chemistry, growth rate and processes during experiments with Coccolithus pelagicus and Calcidiscus leptoporus, 2006

Uptake of half of the fossil fuel CO2 into the ocean causes gradual seawater acidification. This has been shown to slow down calcification of major calcifying groups, such as corals, foraminifera, and coccolithophores. Here we show that two of the most productive marine calcifying species, the coccolithophores Coccolithus pelagicus and Calcidiscus leptoporus, do not follow the CO2-related calcification response previously found. In batch culture experiments, particulate inorganic carbon (PIC) of C. leptoporus changes with increasing CO2 concentration in a nonlinear relationship. A PIC optimum curve is obtained, with a maximum value at present-day surface ocean pCO2 levels (?360 ppm CO2). With particulate organic carbon (POC) remaining constant over the range of CO2 concentrations, the PIC/POC ratio also shows an optimum curve. In the C. pelagicus cultures, neither PIC nor POC changes significantly over the CO2 range tested, yielding a stable PIC/POC ratio. Since growth rate in both species did not change with pCO2, POC and PIC production show the same pattern as POC and PIC. The two investigated species respond differently to changes in the seawater carbonate chemistry, highlighting the need to consider species-specific effects when evaluating whole ecosystem responses. Changes of calcification rate (PIC production) were highly correlated to changes in coccolith morphology. Since our experimental results suggest altered coccolith morphology (at least in the case of C. leptoporus) in the geological past, coccoliths originating from sedimentary records of periods with different CO2 levels were analyzed. Analysis of sediment samples was performed on six cores obtained from locations well above the lysocline and covering a range of latitudes throughout the Atlantic Ocean. Scanning electron micrograph analysis of coccolith morphologies did not reveal any evidence for significant numbers of incomplete or malformed coccoliths of C. pelagicus and C. leptoporus in last glacial maximum and Holocene sediments. The discrepancy between experimental and geological results might be explained by adaptation to changing carbonate chemistry.

Seawater carbonate chemistry and carbon allocation, growth and morphology of the coccolithophore Emiliania huxleyi (calcifying strain CCMP 371) during experiments, 2011

Coccolithophores are unicellular phytoplankton that produce calcium carbonate coccoliths as an exoskeleton. Emiliania huxleyi, the most abundant coccolithophore in the world's ocean, plays a major role in the global carbon cycle by regulating the exchange of CO2 across the ocean-atmosphere interface through photosynthesis and calcium carbonate precipitation. As CO2 concentration is rising in the atmosphere, the ocean is acidifying and ammonium (NH4) concentration of future ocean water is expected to rise. The latter is attributed to increasing anthropogenic nitrogen (N) deposition, increasing rates of cyanobacterial N2 fixation due to warmer and more stratified oceans, and decreased rates of nitrification due to ocean acidification. Thus future global climate change will cause oceanic phytoplankton to experience changes in multiple environmental parameters including CO2, pH, temperature and nitrogen source. This study reports on the combined effect of elevated pCO2 and increased NH4 to nitrate (NO3) ratio (NH4/NO3) on E. huxleyi, maintained in continuous cultures for more than 200 generations under two pCO2 levels and two different N sources. Here we show that NH4 assimilation under N-replete conditions depresses calcification at both low and high pCO2, alters coccolith morphology, and increases primary production. We observed that N source and pCO2 synergistically drive growth rates, cell size and the ratio of inorganic to organic carbon. These responses to N source suggest that, compared to increasing CO2 alone, a greater disruption of the organic carbon pump could be expected in response to the combined effect of increased NH4/NO3 ratio and CO2 level in the future acidified ocean. Additional experiments conducted under lower nutrient conditions are needed prior to extrapolating our findings to the global oceans. Nonetheless, our results emphasize the need to assess combined effects of multiple environmental parameters on phytoplankton biology in order to develop accurate predictions of phytoplankton responses to ocean acidification.

Seawater carbonate chemistry, cell numbers and growth rate during experiments with dinoflagellates, 2007

The effects of dissolved inorganic carbon (DIC) on the growth of 3 red-tide dinoflagellates (Ceratium lineatum, Heterocapsa triquetra and Prorocentrum minimum) were studied at pH 8.0 and at higher pH levels, depending upon the pH tolerance of the individual species. The higher pH levels chosen for experiments were 8.55 for C. lineatum and 9.2 for the other 2 species. At pH 8.0, which approximates the pH found in the open sea, the maximum growth in all species was maintained until the total DIC concentration was reduced below ~0.4 and 0.2 mM for C. lineatum and the other 2 species, respectively. Growth compensation points (concentration of inorganic carbon needed for maintenance of cells) were reached at ~0.18 and 0.05 mM DIC for C. lineatum and the other 2 species, respectively. At higher pH levels, maximum growth rates were lower compared to growth at pH 8, even at very high DIC concentrations, indicating a direct pH effect on growth. Moreover, the concentration of bio-available inorganic carbon (CO2 + HCO3-) required for maintenance as well as the half-saturation constants were increased considerably at high pH compared to pH 8.0. Experiments with pH-drift were carried out at initial concentrations of 2.4 and 1.2 mM DIC to test whether pH or DIC was the main limiting factor at a natural range of DIC. Independent of the initial DIC concentrations, growth rates were similar in both incubations until pH had increased considerably. The results of this study demonstrated that growth of the 3 species was mainly limited by pH, while inorganic carbon limitation played a minor role only at very high pH levels and low initial DIC concentrations.

Seawater chemistry, nutrients, chlorophyll a, and growth rate of Phaeocystis globosa during experiments

Phaeocystis globosa (Prymnesiophyceae) is an ecologically dominating phytoplankton species in many areas around the world. It plays an important role in both the global sulfur and carbon cycles, by the production of dimethylsulfide (DMS) and the drawdown of inorganic carbon. Phaeocystis globosa has a polymorphic life cycle and is considered to be a harmful algal bloom (HAB) forming species. All these aspects make this an interesting species to study the effects of increasing carbon dioxide (CO2) concentrations, due to anthropogenic carbon emissions. Here, the combined effects of three different dissolved carbon dioxide concentrations (CO2(aq)) (low: 4 µmol/kg, intermediate: 6-10 µmol/kg and high CO2(aq): 21-24 µmol/kg) and two different light intensities (low light, suboptimal: 80 µmol photons/m**2/s and high light, light saturated: 240 µmol photons/m**2/s) are reported. The experiments demonstrated that the specific growth rate of P. globosa in the high light cultures decreased with increasing CO2(aq) from 1.4 to 1.1 /d in the low and high CO2 cultures, respectively. Concurrently, the photosynthetic efficiency (Fv/Fm) increased with increasing CO2(aq) from 0.56 to 0.66. The different light conditions affected photosynthetic efficiency and cellular chlorophyll a concentrations, both of which were lower in the high light cultures as compared to the low light cultures. These results suggest that in future inorganic carbon enriched oceans, P. globosa will become less competitive and feedback mechanisms to global change may decrease in strength.

Seawater carbonate chemistry, nutrients, particulate carbon and growth rate of Emiliania huxleyi (AC472), Calcidiscus leptoporus (AC370) and Syracosphaera pulchra (AC418) during experiments, 2011

The response of Emiliania huxleyi (Lohmann) W. W. Hay et H. Mohler, Calcidiscus leptoporus (G. Murray et V. H. Blackman) J. Schiller, andSyracosphaera pulchra Lohmann to elevated partial pressure of carbon dioxide (pCO2) was investigated in batch cultures. We reported on the response of both haploid and diploid life stages of these three species. Growth rate, cell size, particulate inorganic carbon (PIC), and particulate organic carbon (POC) of both life stages were measured at two different pCO2 (400 and 760 parts per million [ppm]), and their organic and inorganic carbon production were calculated. The two life stages within the same species generally exhibited a similar response to elevated pCO2, the response of the haploid stage being often more pronounced than that of the diploid stage. The growth rate was consistently higher at elevated pCO2, but the response of other processes varied among species. Calcification rate of C. leptoporusand of S. pulchra did not change at elevated pCO2, whereas it increased in E. huxleyi. POC production and cell size of both life stages of S. pulchra and of the haploid stage of E. huxleyi markedly decreased at elevated pCO2. It remained unaltered in the diploid stage of E. huxleyi and C. leptoporus and increased in the haploid stage of the latter. The PIC:POC ratio increased in E. huxleyi and was constant in C. leptoporus and S. pulchra. Elevated pCO2 has a significant effect on these three coccolithophore species, the haploid stage being more sensitive. This effect must be taken into account when predicting the fate of coccolithophores in the future ocean.

Abundance and Biomass from heterotrophic bacteria, Synechococcus, Prochlorococcus and Virus in the eastern Mediteranean Sea in October 2008 during SES_GR2

The HCMR_SES_LAGRANGIAN_GR2_ MICROBIAL PARAMETERS dataset is based on samples collected in the framework of the project SESAME, in the North Aegean Sea during October 2008. The objectives were to measure the standing stocks and calculate the production of the microbial compartment of the food web, describe the vertical distribution pattern and characterize its structure and function through the water column as influenced by the BSW. Heterotrophic bacteria, Synechococcus, Prochlorococcus and Virus abundance: Subsamples for virus, heterotrophic bacteria and cyanobacteria (Synechococcus spp. and Prochlorococcus spp.) counting were analyzed using a FACSCalibur (Becton Dickinson) flow cytometer equipped with a standard laser (488 nm) and filter set and using deionized water as sheath fluid. Fluorescent beads with a diameter of 0.97 µm (Polysciences) were added to each sample as an internal standard, and all parameters were normalized to the beads and expressed as relative units. SYBRGreen I stain (Molecular Probe) was used to stain viral and heterotrophic bacterial DNA. Viruses were counted according to (Brussaard 1984). In order to avoid bulk consentrations of viruses samples we dilluted to Tris-EDTA (pH=8,0) buffer to a final sollution of 1/5 to 1/100. Total abundance and nucleid content classes were calculated using the Paint-A-Gate software (Becton Dickinson). Heterotrophic Nanoflagellate abundance: Subsamples (30-150 ml) were concentrated on 25mm black polycarbonate filters of porosity 0.6?m and stained with DAPI for 10 min (Porter and Feig 1980). Under epifluorescence microscopy heterotrophic nanoflagellates (HNAN) were distinguished using UV and blue excitation and enumerated. Nanoflagellates were classified in size categories and the biovolume was calculated. Ciliate abundance: For ciliate identification and enumeration, 100-3000 ml samples were left for 24h-4d for sedimentation and then observed under an inverted microscope. Ciliates were counted, distinguished into size-classes and major taxonomic groups and identified down to genus or species level where possible (Pitta et al. 2005). Heterotrophic bacteria, Synechococcus, Prochlorococcus bacteria: Subsamples for virus, heterotrophic bacteria and cyanobacteria (Synechococcus spp. and Prochlorococcus spp.) counting were analyzed using a FACSCalibur (Becton Dickinson) flow cytometer equipped with a standard laser (488 nm) and filter set and using deionized water as sheath fluid. Fluorescent beads with a diameter of 0.97 µm (Polysciences) were added to each sample as an internal standard, and all parameters were normalized to the beads and expressed as relative units. SYBRGreen I stain (Molecular Probe) was used to stain viral and heterotrophic bacterial DNA. Viruses were counted according to (Brussaard 1984). In order to avoid bulk consentrations of viruses samples we dilluted to Tris-EDTA (pH=8,0) buffer to a final sollution of 1/5 to 1/100. Total abundance and nucleid content classes were calculated using the Paint-A-Gate software (Becton Dickinson). Abundance data were converted into C biomass using 250 fgC cell-1 (Kana & Glibert 1987) for Synechococcus, 50 fgC cell-1 (Campbell et al. 1994) for Prochlorococcus and 20fgC cell-1 (Lee & Fuhrman 1987) for heterotrophic bacteria. Heterotrophic Nanoflagellate biomass: Subsamples (30-150 ml) were concentrated on 25mm black polycarbonate filters of porosity 0.6µm and stained with DAPI for 10 min (Porter and Feig 1980). Under epifluorescence microscopy heterotrophic nanoflagellates (HNAN) were distinguished using UV and blue excitation and enumerated. Nanoflagellates were classified in size categories and the biovolume was calculated. Abundance data were converted into C biomass using 183 fgC µm**3 (Caron et al. 1995). Ciliate biomass: For ciliate identification and enumeration, 100-3000 ml samples were left for 24h-4d for sedimentation and then observed under an inverted microscope. Ciliates were counted, distinguished into size-classes and major taxonomic groups and identified down to genus or species level where possible (Pitta et al. 2005). Ciliate cell sizes were measured and converted into cell volumes using appropriate geometric formulae using image analysis. For biomass estimation, the conversion factor 190 fgC µm**3 was used (Putt and Stoecker 1989).

Integrated values of heterotrophic picoplankton and primary production (euphotic zone) in the Antarctic polar frontal region during POLARSTERN cruises ANT-XIII/2 and ANT-XVI/3

We assessed relationships between phytoplankton standing stock, measured as chlorophyll a (Chl a), primary production (PP), and heterotrophic picoplankton production (HPP), in the epipelagic zone (0-100 m) as well as in the mesopelagic zone (100-1,000 m) in the polar frontal zone of the Atlantic sector of the Southern Ocean in austral summer (late December to January) and fall (March to early May). Integrated epipelagic HPP was positively correlated to integrated PP in summer (data for fall are not available) but not to integrated Chl a. However, integrated mesopelagic HPP was positively correlated to Chl a in summer as well as fall. The mesopelagic fraction of HPP as a percentage of total HPP was also positively correlated to Chl a, whereas the epipelagic fraction of HPP was negatively correlated to it. These results indicate that with increasing phytoplankton standing stock, constituted mainly of highly silicified diatoms, the focus of its consumption by heterotrophic picoplankton shifts from epipelagic to mesopelagic waters. With a growth efficiency of 30%, our HPP data indicate that in both the epipelagic and mesopelagic zone heterotrophic picoplankton consume 20% of PP. Mesopelagic heterotrophic picoplankton consumed around 80% of the sinking flux, measured from depletion of 234Th, which is a lower fraction than that reported from the central and subarctic Pacific. Our analysis indicates that it is important to include mesopelagic HPP in comprehensive assessments of the microbial consumption of PP, phytoplankton biomass, and particulate organic matter in cold oceanic systems with high rates of export production.

Abundance and Biomass from heterotrophic bacteria, Synechococcus, Prochlorococcus and Virus in the Aegean Sea in April 2008 during SES_GR1

The HCMR_SES_LAGRANGIAN_GR1_ MICROBIAL PARAMETERS dataset is based on samples collected in the framework of the project SESAME, in the North Aegean Sea during April 2008. The objectives were to measure the standing stocks and calculate the production of the microbial compartment of the food web, describe the vertical distribution pattern and characterize its structure and function through the water column as influenced by the BSW. Heterotrophic bacteria, Synechococcus, Prochlorococcus and Virus abundance: Subsamples for virus, heterotrophic bacteria and cyanobacteria (Synechococcus spp. and Prochlorococcus spp.) counting were analyzed using a FACSCalibur (Becton Dickinson) flow cytometer equipped with a standard laser (488 nm) and filter set and using deionized water as sheath fluid. Fluorescent beads with a diameter of 0.97 µm (Polysciences) were added to each sample as an internal standard, and all parameters were normalized to the beads and expressed as relative units. SYBRGreen I stain (Molecular Probe) was used to stain viral and heterotrophic bacterial DNA. Viruses were counted according to (Brussaard 1984). In order to avoid bulk consentrations of viruses samples we dilluted to Tris-EDTA (pH=8,0) buffer to a final sollution of 1/5 to 1/100. Total abundance and nucleid content classes were calculated using the Paint-A-Gate software (Becton Dickinson). Heterotrophic Nanoflagellate abundance: Subsamples (30-150 ml) were concentrated on 25mm black polycarbonate filters of porosity 0.6µm and stained with DAPI for 10 min (Porter and Feig 1980). Under epifluorescence microscopy heterotrophic nanoflagellates (HNAN) were distinguished using UV and blue excitation and enumerated. Nanoflagellates were classified in size categories and the biovolume was calculated. Ciliate abundance: For ciliate identification and enumeration, 100-3000 ml samples were left for 24h-4d for sedimentation and then observed under an inverted microscope. Ciliates were counted, distinguished into size-classes and major taxonomic groups and identified down to genus or species level where possible (Pitta et al. 2005). Heterotrophic bacteria, Synechococcus, Prochlorococcus biomass: Subsamples for virus, heterotrophic bacteria and cyanobacteria (Synechococcus spp. and Prochlorococcus spp.) counting were analyzed using a FACSCalibur (Becton Dickinson) flow cytometer equipped with a standard laser (488 nm) and filter set and using deionized water as sheath fluid. Fluorescent beads with a diameter of 0.97 µm (Polysciences) were added to each sample as an internal standard, and all parameters were normalized to the beads and expressed as relative units. SYBRGreen I stain (Molecular Probe) was used to stain viral and heterotrophic bacterial DNA. Viruses were counted according to (Brussaard 1984). In order to avoid bulk consentrations of viruses samples we dilluted to Tris-EDTA (pH=8,0) buffer to a final sollution of 1/5 to 1/100. Total abundance and nucleid content classes were calculated using the Paint-A-Gate software (Becton Dickinson). Abundance data were converted into C biomass using 250 fgC cell-1 (Kana & Glibert 1987) for Synechococcus, 50 fgC cell-1 (Campbell et al. 1994) for Prochlorococcus and 20fgC cell-1 (Lee & Fuhrman 1987) for heterotrophic bacteria. Heterotrophic Nanoflagellate biomass: Subsamples (30-150 ml) were concentrated on 25mm black polycarbonate filters of porosity 0.6µm and stained with DAPI for 10 min (Porter and Feig 1980). Under epifluorescence microscopy heterotrophic nanoflagellates (HNAN) were distinguished using UV and blue excitation and enumerated. Nanoflagellates were classified in size categories and the biovolume was calculated. Abundance data were converted into C biomass using 183 fgC µm**3 (Caron et al. 1995). Ciliate biomass: For ciliate identification and enumeration, 100-3000 ml samples were left for 24h-4d for sedimentation and then observed under an inverted microscope. Ciliates were counted, distinguished into size-classes and major taxonomic groups and identified down to genus or species level where possible (Pitta et al. 2005). Ciliate cell sizes were measured and converted into cell volumes using appropriate geometric formulae using image analysis. For biomass estimation, the conversion factor 190 fgC µm**3 was used (Putt and Stoecker 1989).

The modulating effect of light intensity on the response of the coccolithophore Gephyrocapsa oceanica to ocean acidification

Global change leads to a multitude of simultaneous modifications in the marine realm among which shoaling of the upper mixed layer, leading to enhanced surface layer light intensities, as well as increased carbon dioxide (CO2) concentration are some of the most critical environmental alterations for phytoplankton. In this study, we investigated the responses of growth, photosynthetic carbon fixation and calcification of the coccolithophore Gephyrocapsa oceanica to elevated inline image (51 Pa, 105 Pa, and 152 Pa) (1 Pa ~ 10 µatm) at a variety of light intensities (50-800 µmol photons/m**2/s). By fitting the light response curve, our results showed that rising inline image reduced the maximum rates for growth, photosynthetic carbon fixation and calcification. Increasing light intensity enhanced the sensitivity of these rate responses to inline image, and shifted the inline image optima toward lower levels. Combining the results of this and a previous study (Sett et al. 2014) on the same strain indicates that both limiting low inline image and inhibiting high inline image levels (this study) induce similar responses, reducing growth, carbon fixation and calcification rates of G. oceanica. At limiting low light intensities the inline image optima for maximum growth, carbon fixation and calcification are shifted toward higher levels. Interacting effects of simultaneously occurring environmental changes, such as increasing light intensity and ocean acidification, need to be considered when trying to assess metabolic rates of marine phytoplankton under future ocean scenarios.