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A comparative analysis of data, obtained during four indoor-mesocosm experiments with natural spring plankton communities from the Baltic Sea, was conducted to investigate whether biogeochemical cycling is affected by an increase in water temperature of up to 6 °C above present-day conditions. In all experiments, warming stimulated in particular heterotrophic bacterial processes and had an accelerating effect on the temporal development of phytoplankton blooms. This was also mirrored in the build-up and partitioning of organic matter between particulate and dissolved phases. Thus, warming increased both the magnitude and rate of dissolved organic carbon (DOC) build-up, whereas the accumulation of particulate organic carbon (POC) and phosphorus (POP) decreased with rising temperature. In concert, the observed temperature-mediated changes in biogeochemical components suggest strong shifts in the functioning of marine pelagic food webs and the ocean’s biological carbon pump, hence providing potential feedback mechanisms to Earth’s climate system.

To improve our mechanistic understanding and predictive capacities with respect to climate change effects on the spring phytoplankton bloom in temperate marine systems, we used a process-driven dynamical model to disentangle the impact of potentially relevant factors which are often correlated in the field. The model was based on comprehensive indoor mesocosm experiments run at four temperature and three light regimes. It was driven by time-series of water temperature and irradiance, considered edible and less edible phytoplankton separately, and accounted for density- dependent grazing losses. It successfully reproduced the observed dynamics of well edible phytoplankton in the different temperature and light treatments. Four major factors influenced spring phytoplankton dynamics: temperature, light (cloudiness), grazing, and the success of overwintering phyto- and zooplankton providing the starting biomasses for spring growth. Our study predicts that increasing cloudiness as anticipated for warmer winters for the Baltic Sea region will retard phytoplankton net growth and reduce peak heights. Light had a strong direct effect in contrast to temperature. However, edible phytoplankton was indirectly strongly temperature-sensitive via grazing which was already important in early spring at moderately high algal biomasses and counter-intuitively provoked lower and later algal peaks at higher temperatures. Initial phyto- and zooplankton composition and biomass also had a strong effect on spring algal dynamics indicating a memory effect via the broadly under-sampled overwintering plankton community. Unexpectedly, increased initial phytoplankton biomass did not necessarily lead to earlier or higher spring blooms since the effect was counteracted by subsequently enhanced grazing. Increasing temperature will likely exhibit complex indirect effects via changes in overwintering phytoplankton and grazer biomasses and current grazing pressure. Additionally, effects on the phytoplankton composition due to the species-specific susceptibility to grazing are expected. Hence, we need to consider not only direct but also indirect effects, e.g. biotic interactions, when addressing climate change impacts.

A CO2 enrichment experiment (PeECE III) was carried out in 9 mesocosms in which the seawater carbonate system was manipulated to achieve three different levels of pCO2. At the onset of the experimental period, nutrients were added to all mesocosms in order to initiate phytoplankton blooms. Primary production rates were measured by in-vitro incubations based on 14C-incorporation and oxygen production/consumption. Size fractionated particulate primary production was also determined by 14C incubation and is discussed in relation to phytoplankton composition. Primary production rates increased in response to nutrient addition and a net autotrophic phase with 14C-fixation rates up to 4 times higher than initial was observed midway through the 24 days experiment before net community production (NCP) returned to near-zero and 14C-fixation rates dropped below initial values. No clear heterotrophic phase was observed during the experiment. Based on the 14C-measurements we found higher cumulative primary production at higher pCO2 towards the end of the experiment. CO2 related differences were also found in size fractionated primary production. The most noticeable responses to CO2 treatments with respect to primary production rates occurred in the second half of the experiment when phytoplankton growth had become nutrient limited, and the phytoplankton community changed from diatom to flagellate dominance. This opens for two alternative hypotheses that the effects are either associated with mineral nutrient limited growth, and/or with a change in phytoplankton species composition. The lack of a clear net heterotrophic phase in the last part of the experiment supports the idea that a substantial part of production in the upper layer was not degraded locally, but either accumulated or exported vertically.

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 diazotrophic cyanobacterium Trichodesmium is a significant contributor to marine nitrogen and carbon cycles and has been incorporated in biogeochemical ocean circulation models. To date, parameterization of light as a controlling factor for nitrogen fixation has been based on field observations, where factors other than light also affect Trichodesmium physiology. Here we present data on light-dependent (15 to 1100 µmol quanta m–2 s–1) diazotrophic growth from controlled laboratory experiments and their implications for modeling approaches. We supply a simple empirical model to describe nitrogen fixation by Trichodesmium in batch cultures. Diazotrophic growth of axenic Trichodesmium IMS-101 was light saturated at 180 µmol quanta m–2 s–1 and did not vary significantly at higher photon irradiances up to 1100 µmol quanta m–2 s–1 (μcarbon based ≈ 0.26 d–1). Chlorophyll a (chl a) normalized N2 fixation rates were significantly affected by light intensity during mid-exponential growth (0.74 to 4.45 mol N fixed mol chl a–1 h–1) over the range of photon irradiances tested. In contrast, nitrogen fixation rates normalized to the cellular carbon content were relatively unaffected by light intensity (0.42 to 0.59, averaging 0.5 mmol N mol particulate organic carbon [POC]–1 h–1). Trichodesmium carbon biomass can be used to estimate the nitrogen input by this diazotroph into the ocean; the maximum input rate is 350 nmol N fixed l–1 h–1.

A microcosm experiment was conducted to investigate the interactive effects of rising sea-surface temperature and altered nutrient stoichiometry on the biogeochemical cycling of organic matter in a pelagic algal–bacterial assemblage. Natural seawater, containing a mixed bacterial community, was inoculated with an axenic culture of the bloom-forming diatom species Skeletonema costatum. A factorial combination of three temperatures, simulating weak to strong warming as projected for the end of the 21st century, and either nitrogen (N)-replete or -deficient growth conditions were applied. Depending on the type of nutrient limitation, the mixed algal–bacterial communities displayed pronounced differences in the accumulation and microbial utilization of organic matter in response to warming. Under N-deficient conditions, the build-up of organic matter occurred, irrespective of temperature, dominantly in the particulate pool, and only small amounts of dissolved material accumulated. The subsequent bacterial consumption of organic matter was low, as indicated by measurements of bacterial secondary production and extracellular enzyme activities, and remained also largely unaffected by an increase in temperature from 4°C up to 12°C. Contrastingly, warming resulted in a distinct temperature-dependent increase in the accumulation of dissolved organic carbon compounds under N-replete growth conditions. Moreover, rising temperature notably stimulated the bacterial activity, indicating an enhanced flow of organic matter through the microbial loop. These findings suggest that there will be strong shifts in the biogeochemical cycling of organic matter in the upper ocean in response to increased temperature and nutrient loading that will affect pelagic food-web structures and the biological sequestration of organic matter.

An experimental study was conducted to test the effects of projected sea surface warming (according to the IPPC scenarios) on the accumulation and composition of dissolved organic matter (DOM) during marine phytoplankton blooms in cold seas (

Effects of elevated temperature on the formation and subsequent degradation of diatom aggregates were studied in a laboratory experiment with a natural plankton community from the Kiel Fjord (Baltic Sea). Aggregates were derived from diatom blooms that developed in indoor mesocosms at 2.5 and 8.5 degrees C, corresponding to the 1993 to 2002 mean winter in situ temperature of the Western Baltic Sea and the projected sea surface temperature during winter in 2100, respectively. Formation and degradation of diatom aggregates at these 2 temperatures in the dark were promoted with roller tanks over a period of 11 d. Comparison of the 2 temperature settings revealed an enhanced aggregation potential of diatom cells at elevated temperature, which was likely induced by an increased concentration of transparent exopolymer particles (TEP). The enhanced aggregation potential led to a significantly higher proportion of particulate organic matter in aggregates at 8.5 degrees C. Moreover, the elevated temperature favoured the growth of bacteria, bacterial biomass production, and the activities of sugar- and protein-degrading extracellular enzymes in aggregates. Stimulating effects of rising temperature on growth and metabolism of the bacterial community resulted in an earlier onset of aggregate degradation and silica dissolution. Remineralization of carbon in aggregates at elevated temperature was partially compensated by the formation of carbon-rich TEP during dark incubation. Hence, our results suggest that increasing temperature will affect both formation and degradation of diatom aggregates. We conclude that the vertical export of organic matter through aggregates may change in the future, depending on the magnitude and vertical depth penetration of warming in the ocean.

The pelagic ocean harbors one of the largest ecosystems on Earth. It is responsible for approximately half of global primary production, sustains worldwide fisheries, and plays an important role in the global carbon cycle. Ocean warming caused by anthropogenic climate change is already starting to impact the marine biota, with possible consequences for ocean productivity and ecosystem services. Because temperature sensitivities of marine autotrophic and heterotrophic processes differ greatly, ocean warming is expected to cause major shifts in the flow of carbon and energy through the pelagic system. Attempts to integrate such biological responses into marine ecosystem and biogeochemical models suffer from a lack of empirical data. Here, we show, using an indoor-mesocosm approach, that rising temperature accelerates respiratory consumption of organic carbon relative to autotrophic production in a natural plankton community. Increasing temperature by 2–6 °C hence decreased the biological drawdown of dissolved inorganic carbon in the surface layer by up to 31%. Moreover, warming shifted the partitioning between particulate and dissolved organic carbon toward an enhanced accumulation of dissolved compounds. In line with these findings, the loss of organic carbon through sinking was significantly reduced at elevated temperatures. The observed changes in biogenic carbon flow have the potential to reduce the transfer of primary produced organic matter to higher trophic levels, weaken the ocean's biological carbon pump, and hence provide a positive feedback to rising atmospheric CO 2 .

The oceans have absorbed nearly half of the fossil-fuel carbon dioxide (CO2) emitted into the atmosphere since pre-industrial times1, causing a measurable reduction in seawater pH and carbonate saturation2. If CO2 emissions continue to rise at current rates, upper-ocean pH will decrease to levels lower than have existed for tens of millions of years and, critically, at a rate of change 100 times greater than at any time over this period3. Recent studies have shown effects of ocean acidification on a variety of marine life forms, in particular calcifying organisms4, 5, 6. Consequences at the community to ecosystem level, in contrast, are largely unknown. Here we show that dissolved inorganic carbon consumption of a natural plankton community maintained in mesocosm enclosures at initial CO2 partial pressures of 350, 700 and 1,050 μatm increases with rising CO2. The community consumed up to 39% more dissolved inorganic carbon at increased CO2 partial pressures compared to present levels, whereas nutrient uptake remained the same. The stoichiometry of carbon to nitrogen drawdown increased from 6.0 at low CO2 to 8.0 at high CO2, thus exceeding the Redfield carbon:nitrogen ratio of 6.6 in today’s ocean7. This excess carbon consumption was associated with higher loss of organic carbon from the upper layer of the stratified mesocosms. If applicable to the natural environment, the observed responses have implications for a variety of marine biological and biogeochemical processes, and underscore the importance of biologically driven feedbacks in the ocean to global change.

An indoor mesocosm system was set up to study the response of phytoplankton and zooplankton spring succession to winter and spring warming of sea surface temperatures. The experimental temperature regimes consisted of the decadal average of the Kiel Bight, Baltic Sea, and three elevated regimes with 2 degrees C, 4 degrees C, and 6 degrees C temperature difference from that at baseline. While the peak of the phytoplankton spring bloom was accelerated only weakly by increasing temperatures (1.4 days per degree Celsius), the subsequent biomass minimum of phytoplankton was accelerated more strongly (4.25 days per degree Celsius). Phytoplankton size structure showed a pronounced response to warming, with large phytoplankton being more dominant in the cooler mesocosms. The first seasonal ciliate peak was accelerated by 2.1 days per degree Celsius and the second one by 2.0 days per degree Celsius. The over-wintering copepod populations declined faster in the warmer mesocosm, and the appearance of nauplii was strongly accelerated by temperature (9.2 days per degree Celsius). The strong difference between the acceleration of the phytoplankton peak and the acceleration of the nauplii could be one of the "Achilles heels" of pelagic systems subject to climate change, because nauplii are the most starvation-sensitive life cycle stage of copepods and the most important food item of first-feeding fish larvae.

Increasing atmospheric carbon dioxide (CO2) concentrations due to anthropogenic fossil fuel combustion are currently changing the ocean's chemistry. Increasing oceanic [CO2] and consequently decreasing seawater pH have the potential to significantly impact marine life. Here we describe and analyze the build-up and decline of a natural phytoplankton bloom initiated during the 2005 mesocosm Pelagic Ecosystem CO2 Enrichment study (PeECE III). The draw-down of inorganic nutrients in the upper surface layer of the mesocosms was reflected by a concomitant increase of organic matter until day t11, the peak of the bloom. From then on, biomass standing stocks steadily decreased as more and more particulate organic matter was lost into the deeper layer of the mesocosms. We show that organic carbon export to the deeper layer was significantly enhanced at elevated CO2. This phenomenon might have impacted organic matter remineralization leading to decreased oxygen concentrations in the deeper layer of the high CO2 mesocosms as indicated by deep water ammonium concentrations. This would have important implications for our understanding of pelagic ecosystem functioning and future carbon cycling.