ALBERT

Description: The flux of organic particles below the mixed layer is one major pathway of carbon from the surface into the deep ocean. The magnitude of this export flux depends on two major processes—remineralization rates and sinking velocities. Here, we present an efficient method to measure sinking velocities of particles in the size range from approximately 3–400 μm by means of video microscopy (FlowCAM®). The method allows rapid measurement and automated analysis of mixed samples and was tested with polystyrene beads, different phytoplankton species, and sediment trap material. Sinking velocities of polystyrene beads were close to theoretical values calculated from Stokes’ Law. Sinking velocities of the investigated phytoplankton species were in reasonable agreement with published literature values and sinking velocities of material collected in sediment trap increased with particle size. Temperature had a strong effect on sinking velocities due to its influence on seawater viscosity and density. An increase in 9 °C led to a measured increase in sinking velocities of ~40 %. According to this temperature effect, an average temperature increase in 2 °C as projected for the sea surface by the end of this century could increase sinking velocities by about 6 % which might have feedbacks on carbon export into the deep ocean.

Description: Increasing atmospheric CO2 concentrations are expected to impact pelagic ecosystem functioning in the near future by driving ocean acidification and increasing global average temperatures. We examined the effects of these two processes on the growth rate and carbon production of the calcifying coccolithophore Gephyrocapsa oceanica. 12 pCO2treatment levels (20-2500 atm) at three temperatures (15, 20 and 25°C) were established in artificial seawater by combined additions of Na2CO3 and HCl (total alkalinity constant in all treatments). Specific growth rate, particulate inorganic carbon (PIC) and particulate organic carbon (POC) production rates showed an optimum curve response to increasing pCO2 at all temperatures. Optima were found between pCO2 levels of 250-650 atm with a tendency to higher pCO2 levels at higher temperatures. Carbon production rates increased with temperature but sensitivity to increasing pCO2 remained similar at all temperatures. The PIC:POC ratio was temperature-independent and linearly decreased with increasing pCO2. The results obtained here contribute to a better understanding of the responses of coccolithophores to changing environmental conditions and may help to parameterize synergistic effects of ocean warming and acidification in marine biogeochemical models.

Description: Marine calcareous sediments provide a fundamental basis for paleoceanographic 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. The results demonstrate the complex interaction between environmental forcing and physiological control on the strontium partitioning in coccolithophore calcite. 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 results give an extended insight into the driving factors that lead to variations in the coccolith Mg / Ca ratio and can be used for Sr / Ca and Mg / Ca paleoproxy calibration.

Description: Diatoms can occur as single cells or as chain-forming aggregates. These two strategies affect buoyancy, predator evasion, light absorption and nutrient uptake. Adjacent cells in chains establish connections through various processes that determine strength and flexibility of the bonds, and at distinct cellular locations defining colony structure. Chain length has been found to vary with temperature and nutrient availability as well as being positively correlated with growth rate. However, the potential effect of enhanced carbon dioxide (CO2) concentrations and consequent changes in seawater carbonate chemistry on chain formation is virtually unknown. Here we report on experiments with semi-continuous cultures of the freshly isolated diatom Asterionellopsis glacialis grown under increasing CO2 levels ranging from 320 to 3400 mu atm. We show that the number of cells comprising a chain, and therefore chain length, increases with rising CO2 concentrations. We also demonstrate that while cell division rate changes with CO2 concentrations, carbon, nitrogen and phosphorus cellular quotas vary proportionally, evident by unchanged organic matter ratios. Finally, beyond the optimum CO2 concentration for growth, carbon allocation changes from cellular storage to increased exudation of dissolved organic carbon. The observed structural adjustment in colony size could enable growth at high CO2 levels, since longer, spiral-shaped chains are likely to create microclimates with higher pH during the light period. Moreover increased chain length of Asterionellopsis glacialis may influence buoyancy and, consequently, affect competitive fitness as well as sinking rates. This would potentially impact the delicate balance between the microbial loop and export of organic matter, with consequences for atmospheric carbon dioxide.

Description: Increasing atmospheric CO2 concentrations are expected to impact pelagic ecosystem functioning in the near future by driving ocean warming and acidification. While numerous studies have investigated impacts of rising temperature and seawater acidification on planktonic organisms separately, little is presently known on their combined effects. To test for possible synergistic effects we exposed two coccolithophore species, Emiliania huxleyi and Gephyrocapsa oceanica, to a CO2 gradient ranging from ,0.5–250 mmol kg21 (i.e. ,20–6000 matm pCO2) at three different temperatures (i.e. 10, 15, 20uC for E. huxleyi and 15, 20, 25uC for G. oceanica). Both species showed CO2-dependent optimum-curve responses for growth, photosynthesis and calcification rates at all temperatures. Increased temperature generally enhanced growth and production rates and modified sensitivities of metabolic processes to increasing CO2. CO2 optimum concentrations for growth, calcification, and organic carbon fixation rates were only marginally influenced from low to intermediate temperatures. However, there was a clear optimum shift towards higher CO2 concentrations from intermediate to high temperatures in both species. Our results demonstrate that the CO2 concentration where optimum growth, calcification and carbon fixation rates occur is modulated by temperature. Thus, the response of a coccolithophore strain to ocean acidification at a given temperature can be negative, neutral or positive depending on that strain’s temperature optimum. This emphasizes that the cellular responses of coccolithophores to ocean acidification can only be judged accurately when interpreted in the proper eco-physiological context of a given strain or species. Addressing the synergistic effects of changing carbonate chemistry and temperature is an essential step when assessing the success of coccolithophores in the future ocean.

Description: An indoor mesocosm experiment was carried out to investigate the combined effects of ocean acidification and warming on the species composition and biogeochemical element cycling during a winter/spring bloom with a natural phytoplankton assemblage from the Kiel fjord, Germany. The experimental setup consisted of a "Control" (ambient temperature of similar to 4.8 degrees C and similar to 535 +/- 25 mu atm pCO(2)), a "High-CO2" treatment (ambient temperature and initially 1020 +/- 45 mu atm pCO(2)) and a "Greenhouse" treatment (similar to 8.5 degrees C and initially 990 +/- 60 mu atm pCO(2)). Nutrient replete conditions prevailed at the beginning of the experiment and light was provided at in situ levels upon reaching pCO(2) target levels. A diatom-dominated bloom developed in all treatments with Skeletonema costatum as the dominant species but with an increased abundance and biomass contribution of larger diatom species in the Greenhouse treatment. Conditions in the Greenhouse treatment accelerated bloom development with faster utilization of inorganic nutrients and an earlier peak in phytoplankton biomass compared to the Control and High CO2 but no difference in maximum concentration of particulate organic matter (POM) between treatments. Loss of POM in the Greenhouse treatment, however, was twice as high as in the Control and High CO2 treatment at the end of the experiment, most likely due to an increased proportion of larger diatom species in that treatment. We hypothesize that the combination of warming and acidification can induce shifts in diatom species composition with potential feedbacks on biogeochemical element cycling.