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.

Description: Phytoplankton experience strong and abrupt variations in light intensity. How cells cope with these changes influences their competitiveness in a highly dynamical environment. While a considerable amount of work has focused on photoacclimation, it is still unknown whether processes specific of phytoplankton groups (e.g. calcification and silicification) influence their response to changing light. Here we show that the diatom Phaeodactylum tricornutum and the coccolithophore Emiliania huxleyi respond to an abrupt increase in irradiance by increasing carbon fixation rates, decreasing light absorption through the decrease of light-harvesting pigments and increasing energy dissipation through the xanthophyll cycle. In addition, E. huxleyi rapidly increases calcium carbonate precipitation in response to elevated light intensity, thereby providing an additional sink for excess energy. Differences between the 2 species also emerge with regard to the magnitude and timing of their individual responses. While E. huxleyi show a pronounced decrease in chlorophyll a and fucoxanthin cellular contents following increased light intensity, P. tricornutum has a faster increase in diadinoxanthin quota, a slower decrease in Fv/Fm (ratio of variable to maximum fluorescence) and a stronger increase in organic carbon fixation rate during the first 10 min. Our findings provide further evidence of species-specific responses to abrupt changes in light intensity, which may partly depend on the phytoplankton functional groups, with coccolithophores having a supplementary path (calcification) for the rapid dissipation of excess energy produced after an abrupt increase in light intensity. These differences might influence competition between coexisting species and may therefore have consequences at the community level.

Description: A mesocosm experiment was conducted to investigate the impact of rising fCO2 on the build-up and decline of organic matter during coastal phytoplankton blooms. Five mesocosms (∼38 m³ each) were deployed in the Baltic Sea during spring (2009) and enriched with CO2 to yield a gradient of 355–862 µatm. Mesocosms were nutrient fertilized initially to induce phytoplankton bloom development. Changes in particulate and dissolved organic matter concentrations, including dissolved high-molecular weight (〉1 kDa) combined carbohydrates, dissolved free and combined amino acids as well as transparent exopolymer particles (TEP), were monitored over 21 days together with bacterial abundance, and hydrolytic extracellular enzyme activities. Overall, organic matter followed well-known bloom dynamics in all CO2 treatments alike. At high fCO2, higher ΔPOC:ΔPON during bloom rise, and higher TEP concentrations during bloom peak, suggested preferential accumulation of carbon-rich components. TEP concentration at bloom peak was significantly related to subsequent sedimentation of particulate organic matter. Bacterial abundance increased during the bloom and was highest at high fCO2. We conclude that increasing fCO2 supports production and exudation of carbon-rich components, enhancing particle aggregation and settling, but also providing substrate and attachment sites for bacteria. More labile organic carbon and higher bacterial abundance can increase rates of oxygen consumption and may intensify the already high risk of oxygen depletion in coastal seas in the future.

Description: It is currently under debate whether organisms that regulate their acid–base status under environmental hypercapnia demand additional energy. This could impair animal fitness, but might be compensated for via increased ingestion rates when food is available. No data are yet available for dominant Calanus spp. from boreal and Arctic waters. To fill this gap, we incubated Calanus glacialis at 390, 1120, and 3000 µatm for 16 d with Thalassiosira weissflogii (diatom) as food source on-board RV Polarstern in Fram Strait in 2012. Every 4 d copepods were subsampled from all CO2 treatments and clearance and ingestion rates were determined. During the SOPRAN mesocosm experiment in Bergen, Norway, 2011, we weekly collected Calanus finmarchicus from mesocosms initially adjusted to 390 and 3000 µatm CO2 and measured grazing at low and high pCO2. In addition, copepods were deep frozen for body mass analyses. Elevated pCO2 did not directly affect grazing activities and body mass, suggesting that the copepods did not have additional energy demands for coping with acidification, neither during long-term exposure nor after immediate changes in pCO2. Shifts in seawater pH thus do not seem to challenge these copepod species.

Description: Marine bacteria are the main consumers of freshly produced organic matter. Many enzymatic processes involved in the bacterial digestion of organic compounds were shown to be pH sensitive in previous studies. Due to the continuous rise in atmospheric CO2 concentration, seawater pH is presently decreasing at a rate unprecedented during the last 300 million years but the consequences for microbial physiology, organic matter cycling and marine biogeochemistry are still unresolved. We studied the effects of elevated seawater pCO2 on a natural plankton community during a large-scale mesocosm study in a Norwegian fjord. Nine Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS) were adjusted to different pCO2 levels ranging initially from ca. 280 to 3000 µatm and sampled every second day for 34 days. The first phytoplankton bloom developed around day 5. On day 14, inorganic nutrients were added to the enclosed, nutrient-poor waters to stimulate a second phytoplankton bloom, which occurred around day 20. Our results indicate that marine bacteria benefit directly and indirectly from decreasing seawater pH. During the first phytoplankton bloom, 5-10% more transparent exopolymer particles were formed in the high pCO2 mesocosms. Simultaneously, the efficiency of the protein-degrading enzyme leucine aminopeptidase increased with decreasing pH resulting in up to three times higher values in the highest pCO2/lowest pH mesocosm compared to the controls. In general, total and cell-specific aminopeptidase activities were elevated under low pH conditions. The combination of enhanced enzymatic hydrolysis of organic matter and increased availability of gel particles as substrate supported up to 28% higher bacterial abundance in the high pCO2 treatments. We conclude that ocean acidification has the potential to stimulate the bacterial community and facilitate the microbial recycling of freshly produced organic matter, thus strengthening the role of the microbial loop in the surface ocean.

Description: Rates of gross primary production (GPP), respiration (R), and net calcification (Gnet) in coral reef sediments are expected to change in response to global warming (and the consequent increase in sea surface temperature) and coastal eutrophication (and the subsequent increase in the concentration of organic matter (OM) being filtered by permeable coral reef carbonate sediments). To date, no studies have examined the combined effect of seawater warming and OM enrichment on coral reef carbonate sediment metabolism and dissolution. This study used 22-hour in situ benthic chamber incubations to examine the combined effect of temperature (T) and OM, in the form of coral mucus and phytodetritus, on GPP, R, and Gnet in the permeable coral reef carbonate sediments of Heron Island lagoon, Australia. Compared to control incubations, both warming (+2.4 ºC) and OM increased R and GPP. Under warmed conditions, R was enhanced to a greater extent than GPP, resulting in a shift to net heterotrophy and net dissolution. Under both phytodetritus and coral mucus treatments, GPP was enhanced to a greater extent than R, resulting in a net increase in GPP/R and Gnet. The combined effect of warming and OM enhanced R and GPP, but the net effect on GPP/R and Gnet was not significantly different from control incubations. These findings show that a shift to net heterotrophy and dissolution due to short-term increases in seawater warming may be countered by a net increase GPP/R and Gnet due to short-term increases in nutrient release from OM.

Description: Ocean acidification (OA) and organic matter enrichment (due to coastal eutrophication) could act in concert to shift coral reef carbonate sediments from a present state of net calcification to a future state of net dissolution, but no studies have examined the combined effect of these stressors on sediment metabolism and dissolution. This study used 22-hour incubations in flume aquaria with captive sediment communities to measure the combined effect of OA and organic matter (OM) enrichment, on coral reef sediment gross primary productivity (GPP), respiration (R), and net calcification (Gnet). Relative to control sediment communities, both OA ( 1000 µatm) and OM enrichment (+ 40 µmol C/L) significantly decreased rates of sediment Gnet by 98% and 15% mmol CaCO3/m**2/h, respectively , but the mechanism behind this decrease differed. The OA-mediated transition to net dissolution was geochemical, as rates of GPP and R remained unaffected and dissolution was solely enhanced by a decline in the aragonite saturation state (Omega arg) of the overlying water column. In contrast, the OM-mediated decline in Gnet was due to a decline in GPP/R, thereby biologically reducing overlying seawater Ωarg due to the increased respiratory addition of CO2. The decrease in Gnet in response to a combination of both stressors was additive (- 10% relative to OA alone) but this decrease did not significantly differ from the effect of OA alone. In this study OA was the primary driver of future carbonate sediment dissolution, but longer-term experiments with chronic organic matter enrichment are required.