Description: Particle aggregation and the consequent formation of marine snow alter important properties of biogenic particles (size, sinking rate, degradability), thus playing a key role in controlling the vertical flux of organic matter to the deep ocean. However, there are still large uncertainties about rates and mechanisms of particle aggregation, as well as the role of plankton community structure in modifying biomass transfer from small particles to large fast-sinking aggregates.Here we present data from a high-resolution underwater camera system that we used to observe particle size distributions and formation of marine snow (aggregates 〉0.5 mm) over the course of a 9-week in situ mesocosm experiment in the Eastern Subtropical North Atlantic. After an oligotrophic phase of almost 4 weeks, addition of nutrient-rich deep water (650 m) initiated the development of a pronounced diatom bloom and the subsequent formation of large marine snow aggregates in all 8 mesocosms. We observed a substantial time lag between the peaks of chlorophyll a and marine snow biovolume of 9-12 days, which is much longer than previously reported and indicates a marked temporal decoupling of phytoplankton growth and marine snow formation during our study. Despite this time lag, our observations revealed substantial transfer of biomass from small particle sizes (single phytoplankton cells and chains) to marine snow aggregates of up to 2.5 mm diameter (ESD), with most of the biovolume being contained in the 0.5-1 mm size range. Notably, the abundance and community composition of mesozooplankton had a substantial influence on the temporal development of particle size spectra and formation of marine snow aggregates: While higher copepod abundances were related to reduced aggregate formation and biomass transfer towards larger particle sizes, the presence of appendicularia and doliolids enhanced formation of large marine snow.Furthermore, we combined in situ particle size distributions with measurements of particle sinking velocity to compute instantaneous (potential) vertical mass flux. However, somewhat surprisingly, we did not find a coherent relationship between our computed flux and measured vertical mass flux (collected by sediment traps in 15 m depth). Although the onset of measured vertical flux roughly coincided with the emergence of marine snow, we found substantial variability in mass flux among mesocosms that was not related to marine snow numbers, and was instead presumably driven by zooplankton-mediated alteration of sinking biomass and export of small particles (fecal pellets).Altogether, our findings highlight the role of zooplankton community composition and feeding interactions on particle size spectra and formation of marine snow aggregates, with important implications for our understanding of particle aggregation and vertical flux of organic matter in the ocean.

Description: Highlights: • Ocean acidification increases phytoplankton standing stock. • This increase is more pronounced in smaller-sized taxa. • Primary consumers reac differently depending on nutrient availability. • Bacteria and micro-heterotrophs benefited under limiting conditions. • In general, heterotrophs are negatively affected at nutrient replete periods. Abstract: In situ mesocosm experiments on the effect of ocean acidification (OA) are an important tool for investigating potential OA-induced changes in natural plankton communities. In this study we combined results from various in-situ mesocosm studies in two different ocean regions (Arctic and temperate waters) to reveal general patterns of plankton community shifts in response to OA and how these changes are modulated by inorganic nutrient availability. Overall, simulated OA caused an increase in phytoplankton standing stock, which was more pronounced in smaller-sized taxa. This effect on primary producers was channelled differently into heterotroph primary consumers depending on the inorganic nutrient availability. Under limiting conditions, bacteria and micro-heterotrophs benefited with inconsistent responses of larger heterotrophs. During nutrient replete periods, heterotrophs were in general negatively affected, although there was an increase of some mesozooplankton developmental stages (i.e. copepodites). We hypothesize that changes in phytoplankton size distribution and community composition could be responsible for these food web responses.

Description: Oceanic uptake and long-term storage of atmospheric carbon dioxide (CO2) are strongly driven by the marine “biological pump,” i.e., sinking of biotically fixed inorganic carbon and nutrients from the surface into the deep ocean (Sarmiento and Bender, 1994; Volk and Hoffert, 1985). Sinking velocity of marine particles depends on seawater viscosity, which is strongly controlled by temperature (Sharqawy et al., 2010). Consequently, marine particle flux is accelerated as ocean temperatures increase under global warming (Bach et al., 2012). Here we show that this previously overlooked “viscosity effect” could have profound impacts on marine biogeochemical cycling and carbon uptake over the next centuries to millennia. In our global warming simulation, the viscosity effect accelerates particle sinking by up to 25%, thereby effectively reducing the portion of organic matter that is respired in the surface ocean. Accordingly, the biological carbon pump's efficiency increases, enhancing the sequestration of atmospheric CO2 into the ocean. This effect becomes particularly important on longer time scales when warming reaches the ocean interior. At the end of our simulation (4000 A.D.), oceanic carbon uptake is 17% higher, atmospheric CO2 concentration is 180 ppm lower, and the increase in global average surface temperature is 8% weaker when considering the viscosity effect. Consequently, the viscosity effect could act as a long-term negative feedback mechanism in the global climate system.

Description: The ocean's potential to export carbon to depth partly depends on the fraction of primary production (PP) sinking out of the euphotic zone (i.e., the e-ratio). Measurements of PP and export flux are often performed simultaneously in the field, although there is a temporal delay between those parameters. Thus, resulting e-ratio estimates often incorrectly assume an instantaneous downward export of PP to export flux. Evaluating results from four mesocosm studies, we find that peaks in organic matter sedimentation lag chlorophyll a peaks by 2 to 15 days. We discuss the implications of these time lags (TLs) for current e-ratio estimates and evaluate potential controls of TL. Our analysis reveals a strong correlation between TL and the duration of chlorophyll a buildup, indicating a dependency of TL on plankton food web dynamics. This study is one step further toward time-corrected e-ratio estimates

Description: A global Earth System model is employed to investigate the role of direct temperature effects in the response of marine ecosystems to climate change. While model configurations with and without consideration of explicit temperature effects can reproduce observed current biogeochemical tracer distributions and estimated carbon export about equally well, carbon flow through the model ecosystem reveals strong temperature sensitivities. Depending on whether biological processes are assumed temperature sensitive or not, simulated marine net primary production (NPP) increases or decreases under projected climate change driven by a business-as-usual CO2 emission scenario for the 21st century. This suggests that indirect temperature effects such as changes in the supply of nutrients and light are not the only relevant factors to be considered when modeling the response of marine ecosystems to climate change. A better understanding of direct temperature effects on marine ecosystems is required before even the direction of change in NPP can be reliably predicted.

Description: Gravitational sinking of photosynthetically fixed particulate organic carbon (POC) constitutes a key component of the biological carbon pump. The fraction of POC leaving the surface ocean depends on POC sinking velocity (SV) and remineralization rate (Cremin), both of which depend on plankton community structure. However, the key drivers in plankton communities controlling SV and Cremin are poorly constrained. In fall 2014, we conducted a 6 weeks mesocosm experiment in the subtropical NE Atlantic Ocean to study the influence of plankton community structure on SV and Cremin. Oligotrophic conditions prevailed for the first 3 weeks, until nutrient‐rich deep water injected into all mesocosms stimulated diatom blooms. SV declined steadily over the course of the experiment due to decreasing CaCO3 ballast and – according to an optical proxy proposed herein – due to increasing aggregate porosity mostly during an aggregation event after the diatom bloom. Furthermore, SV was positively correlated with the contribution of picophytoplankton to the total phytoplankton biomass. Cremin was highest during a Synechococcus bloom under oligotrophic conditions and in some mesocosms during the diatom bloom after the deep‐water addition while it was particularly low during harmful algal blooms. The temporal changes were considerably larger in Cremin (max. 15‐fold) than in SV (max. 3‐fold). Accordingly, estimated POC transfer efficiency to 1000 m was mainly dependent on how the plankton community structure affected Cremin. Our approach revealed key players and interactions in the plankton food web influencing POC export efficiency thereby improving our mechanistic understanding of the biological carbon pump. Key points Sinking velocity was higher during oligotrophy than during blooms which is linked to ballast, porosity, and phytoplankton size structure Remineralization was highly variable but tended to be higher during Synechococcus or diatom blooms and lower during harmful algal blooms Plankton community structure had a considerably larger influence on particle remineralization rate than on sinking velocity

Description: Ocean acidification (OA) is one of the most critical anthropogenic threats to marine ecosystems. While significant ecological responses of plankton communities to OA have been revealed mainly by small-scale laboratory approaches, the interactive effect of OA-related changes on zooplankton metabolism and their biogeochemical implications in the natural environment still remains less well understood. Here, we explore the responses of zooplankton respiration and ammonium excretion, two key processes in the nutrient cycling, to high pCO2 levels in a 9-week in situ mesocosm experiment conducted during the autumn oligotrophic season in the subtropical northeast Atlantic. By simulating an upwelling event halfway through the study, we further evaluated the combined effects of OA and nutrient availability on the physiology of micro-and mesozooplankton. OA conditions generally resulted in a reduction in the biomass-specific metabolic and enzymatic rates, particularly in the mesozooplankton community. The situation reversed after the nutrient-rich deep-water addition, which initially promoted a diatom bloom and increased heterotrophic activities in all mesocosms. Under high pCO2 conditions (〉800 μatm), however, the nutrient fertilization triggered the proliferation of the harmful alga Vicicitus globosus, with important consequences for the metabolic performance of the two zooplankton size classes. Here, the zooplankton contribution to the remineralization of organic matter and nitrogen regeneration dropped by 30% and 24%, respectively, during the oligotrophic period, and by 40% and 70% during simulated upwelling. Overall, our results indicate a potential reduction in the biogeochemical role of zooplankton under future ocean conditions, with more evident effects on the large mesozooplankton and during high productivity events