Description: Diazotrophic dinitrogen (N2) fixation contributes ~76% to "new" nitrogen inputs to the sunlit open ocean, but environmental factors determining N2 fixation rates are not well constrained. Excess phosphate (phosphate-nitrate/16 〉 0) and iron availability control N2 fixation rates in the eastern tropical North Atlantic (ETNA), but it remains an open question how excess phosphate is generated within or supplied to the phosphate-depleted sunlit layer. Our observations in the ETNA region (8°N-15°N, 19°W-23°W) suggest that Prochlorococcus and Synechococcus, the two ubiquitous non-diazotrophic cyanobacteria with cellular N:P ratios higher than the Redfield ratio, create an environment of excess phosphate, which cannot be explained by diapycnal mixing, atmospheric, and riverine inputs. Thus, our results unveil a new biogeochemical niche construction mechanism by non-diazotrophic cyanobacteria for their diazotrophic phylum group members (N2 fixers). Our observations may help to understand the prevalence of diazotrophy in low-phosphate, oligotrophic regions.

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: Coccolithophores are important calcifying phytoplankton predicted to be impacted by changes in ocean carbonate chemistry caused by the absorption of anthropogenic CO2. However, it is difficult to disentangle the effects of the simultaneously changing carbonate system parameters (CO2, bicarbonate, carbonate and protons) on the physiological responses to elevated CO2. Here, we adopted a multifactorial approach at constant pH or CO2 whilst varying dissolved inorganic carbon (DIC) to determine physiological and transcriptional responses to individual carbonate system parameters. We show that Emiliania huxleyi is sensitive to low CO2 (growth and photosynthesis) and low bicarbonate (calcification) as well as low pH beyond a limited tolerance range, but is much less sensitive to elevated CO2 and bicarbonate. Multiple up-regulated genes at low DIC bear the hallmarks of a carbon-concentrating mechanism (CCM) that is responsive to CO2 and bicarbonate but not to pH. Emiliania huxleyi appears to have evolved mechanisms to respond to limiting rather than elevated CO2. Calcification does not function as a CCM, but is inhibited at low DIC to allow the redistribution of DIC from calcification to photosynthesis. The presented data provides a significant step in understanding how E. huxleyi will respond to changing carbonate chemistry at a cellular level.

Description: About 50 Gt of carbon is fixed photosynthetically by surface ocean phytoplankton communities every year. Part of this organic matter is reprocessed within the plankton community to form aggregates which eventually sink and export carbon into the deep ocean. The fraction of organic matter leaving the surface ocean is partly dependent on aggregate sinking velocity which accelerates with increasing aggregate size and density, where the latter is controlled by ballast load and aggregate porosity. In May 2011, we moored nine 25 m deep mesocosms in a Norwegian fjord to assess on a daily basis how plankton community structure affects material properties and sinking velocities of aggregates (Ø 80–400 µm) collected in the mesocosms' sediment traps. We noted that sinking velocity was not necessarily accelerated by opal ballast during diatom blooms, which could be due to relatively high porosity of these rather fresh aggregates. Furthermore, estimated aggregate porosity (Pestimated) decreased as the picoautotroph (0.2–2 µm) fraction of the phytoplankton biomass increased. Thus, picoautotroph-dominated communities may be indicative for food webs promoting a high degree of aggregate repackaging with potential for accelerated sinking. Blooms of the coccolithophore Emiliania huxleyi revealed that cell concentrations of ~1500 cells/mL accelerate sinking by about 35–40%, which we estimate (by one-dimensional modeling) to elevate organic matter transfer efficiency through the mesopelagic from 14 to 24%. Our results indicate that sinking velocities are influenced by the complex interplay between the availability of ballast minerals and aggregate packaging; both of which are controlled by plankton community structure.

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: 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: Dinitrogen (N2) fixation is a major source of bioavailable nitrogen to oligotrophic ocean communities. Yet, we have limited understanding how ongoing climate change could alter N2 fixation. Most of our understanding is based on short-term laboratory experiments conducted on individual N2-fixing species whereas community-level approaches are rare. In this longer-term in situ mesocosm study, we aimed to improve our understanding on the role of rising atmospheric carbon dioxide (CO2) and simulated deep water upwelling on N2 and carbon (C) fixation rates in a natural oligotrophic plankton community. We deployed nine mesocosms in the subtropical North Atlantic Ocean and enriched seven of these with CO2 to yield a range of treatments (partial pressure of CO2, pCO2 = 352–1025 μatm). We measured rates of N2 and C fixation in both light and dark incubations over the 55-day study period. High pCO2 negatively impacted light and dark N2 fixation rates in the oligotrophic phase before simulated upwelling, while the effect reversed in the light N2 fixation rates in the bloom decay phase after added nutrients were consumed. Dust deposition and simulated upwelling of nutrient-rich deep water increased N2 fixation rates and nifH gene abundances of selected clades including the unicellular diazotrophic cyanobacterium clade UCYN-B. Elevated pCO2 increased C fixation rates in the decay phase. We conclude that elevated pCO2 and pulses of upwelling have pronounced effects on diazotrophy and primary producers, and upwelling and dust deposition modify the pCO2 effect in natural assemblages.

Description: Gelatinous zooplankton are increasingly recognized to play a key role in the ocean's biological carbon pump. Appendicularians, a class of pelagic tunicates, are among the most abundant gelatinous plankton in the ocean, but it is an open question how their contribution to carbon export might change in the future. Here, we conducted an experiment with large volume in situ mesocosms (~55–60 m3 and 21 m depth) to investigate how ocean acidification (OA) extreme events affect food web structure and carbon export in a natural plankton community, particularly focusing on the keystone species Oikopleura dioica, a globally abundant appendicularian. We found a profound influence of O. dioica on vertical carbon fluxes, particularly during a short but intense bloom period in the high CO2 treatment, during which carbon export was 42%–64% higher than under ambient conditions. This elevated flux was mostly driven by an almost twofold increase in O. dioica biomass under high CO2. This rapid population increase was linked to enhanced fecundity (+20%) that likely resulted from physiological benefits of low pH conditions. The resulting competitive advantage of O. dioica resulted in enhanced grazing on phytoplankton and transfer of this consumed biomass into sinking particles. Using a simple carbon flux model for O. dioica, we estimate that high CO2 doubled the carbon flux of discarded mucous houses and fecal pellets, accounting for up to 39% of total carbon export from the ecosystem during the bloom. Considering the wide geographic distribution of O. dioica, our findings suggest that appendicularians may become an increasingly important vector of carbon export with ongoing OA.

Description: About 50 Gt of carbon is fixed photosynthetically by surface ocean phytoplankton communities every year. Part of this organic matter is reprocessed within the plankton community to form aggregates which eventually sink and export carbon into the deep ocean. The fraction of organic matter leaving the surface ocean is partly dependent on aggregate sinking velocity which accelerates with increasing aggregate size and density, where the latter is controlled by ballast load and aggregate porosity. In May 2011, we moored nine 25 m deep mesocosms in a Norwegian fjord to assess on a daily basis how plankton community structure affects material properties and sinking velocities of aggregates (Ø 80–400 µm) collected in the mesocosms' sediment traps. We noted that sinking velocity was not necessarily accelerated by opal ballast during diatom blooms, which could be due to relatively high porosity of these rather fresh aggregates. Furthermore, estimated aggregate porosity (Pestimated) decreased as the picoautotroph (0.2–2 µm) fraction of the phytoplankton biomass increased. Thus, picoautotroph-dominated communities may be indicative for food webs promoting a high degree of aggregate repackaging with potential for accelerated sinking. Blooms of the coccolithophore Emiliania huxleyi revealed that cell concentrations of ~1500 cells/mL accelerate sinking by about 35–40%, which we estimate (by one-dimensional modeling) to elevate organic matter transfer efficiency through the mesopelagic from 14 to 24%. Our results indicate that sinking velocities are influenced by the complex interplay between the availability of ballast minerals and aggregate packaging; both of which are controlled by plankton community structure.