Mario Esposito

Influence of ocean acidification on a natural winter-to-summer plankton succession: First insights from a long-term mesocosm study draw attention to periods of low nutrient concentrations

Every year, the oceans absorb about 30% of anthropogenic carbon dioxide (CO2) leading to a re-equilibration of the marine carbonate system and decreasing seawater pH. Today, there is increasing awareness that these changes–summarized by the term ocean acidification (OA)–could differentially affect the competitive ability of marine organisms, thereby provoking a restructuring of marine ecosystems and biogeochemical element cycles. In winter 2013, we deployed ten pelagic mesocosms in the Gullmar Fjord at the Swedish west coast in order to study the effect of OA on plankton ecology and biogeochemistry under close to natural conditions. Five of the ten mesocosms were left unperturbed and served as controls (~380 μatm pCO2), whereas the others were enriched with CO2-saturated water to simulate realistic end-of-the-century carbonate chemistry conditions (~760 μatm pCO2). We ran the experiment for 113 days which allowed us to study the influence of high CO2 on an entire winter-to-summer plankton succession and to investigate the potential of some plankton organisms for evolutionary adaptation to OA in their natural environment. This paper is the first in a PLOS collection and provides a detailed overview on the experimental design, important events, and the key complexities of such a “long-term mesocosm” approach. Furthermore, we analyzed whether simulated end-of-the-century carbonate chemistry conditions could lead to a significant restructuring of the plankton community in the course of the succession. At the level of detail analyzed in this overview paper we found that CO2-induced differences in plankton community composition were non-detectable during most of the succession except for a period where a phytoplankton bloom was fueled by remineralized nutrients. These results indicate: (1) Long-term studies with pelagic ecosystems are necessary to uncover OA-sensitive stages of succession. (2) Plankton communities fueled by regenerated nutrients may be more responsive to changing carbonate chemistry than those having access to high inorganic nutrient concentrations and may deserve particular attention in future studies.

Seasonal Changes in Fe along a Glaciated Greenlandic Fjord

Greenland’s ice sheet is the second largest on Earth, and is under threat from a warming Arctic climate. An increase in freshwater discharge from Greenland has the potential to strongly influence the composition of adjacent water masses with the largest impact on marine ecosystems likely to be found within the glaciated fjords. Here we demonstrate that physical and chemical estuarine processes within a large Greenlandic fjord are critical factors in determining the fate of meltwater derived nutrients and particles, especially for non-conservative elements such as Fe. Concentrations of Fe and macronutrients in surface waters along Godthabsfjord, a southwest Greenlandic fjord with freshwater input from 6 glaciers, changed markedly between the onset and peak of the meltwater season due to the development of a thin (<10 m), outflowing, low-salinity surface layer. Dissolved (<0.2 µm) Fe concentrations in meltwater entering Godthabsfjord (200 nM), in freshly melted glacial ice (mean 38 nM) and in surface waters close to a land terminating glacial system (80 nM) all indicated high Fe inputs into the fjord in summer. Total dissolvable (unfiltered at pH <2.0) Fe was similarly high with concentrations always in excess of 100 nM throughout the fjord and reaching up to 5.0 µM close to glacial outflows in summer. Yet, despite the large seasonal freshwater influx into the fjord, Fe concentrations near the fjord mouth in the out-flowing surface layer were similar in summer to those measured before the meltwater season. Furthermore, turbidity profiles indicated that sub-glacial particulate Fe inputs may not actually mix into the outflowing surface layer of this fjord. Emphasis has previously been placed on the possibility of increased Fe export from Greenland as meltwater fluxes increase. Here we suggest that in-fjord processes may be effective at removing Fe from surface waters before it can be exported to coastal seas.

Influence of Ocean Acidification and Deep Water Upwelling on Oligotrophic Plankton Communities in the Subtropical North Atlantic: Insights from an In situ Mesocosm Study

Oceanic uptake of anthropogenic carbon dioxide (CO2) causes pronounced shifts in marine carbonate chemistry and a decrease in seawater pH. Increasing evidence indicates that these changes—summarized by the term ocean acidification (OA)—can significantly affect marine food webs and biogeochemical cycles. However, current scientific knowledge is largely based on laboratory experiments with single species and artificial boundary conditions, whereas studies of natural plankton communities are still relatively rare. Moreover, the few existing community-level studies were mostly conducted in rather eutrophic environments, while less attention has been paid to oligotrophic systems such as the subtropical ocean gyres. Here we report from a recent in situ mesocosm experiment off the coast of Gran Canaria in the eastern subtropical North Atlantic, where we investigated the influence of OA on the ecology and biogeochemistry of plankton communities in oligotrophic waters under close-to-natural conditions. This paper is the first in this Research Topic of Frontiers in Marine Biogeochemistry and provides (1) a detailed overview of the experimental design and important events during our mesocosm campaign, and (2) first insights into the ecological responses of plankton communities to simulated OA over the course of the 62-day experiment. One particular scientific objective of our mesocosm experiment was to investigate how OA impacts might differ between oligotrophic conditions and phases of high biological productivity, which regularly occur in response to upwelling of nutrient-rich deep water in the study region. Therefore, we specifically developed a deep water collection system that allowed us to obtain ~85 m3 of seawater from ~650 m depth. Thereby, we replaced ~20% of each mesocosms volume with deep water and successfully simulated a deep water upwelling event that induced a pronounced plankton bloom. Our study revealed significant effects of OA on the entire food web, leading to a restructuring of plankton communities that emerged during the oligotrophic phase, and was further amplified during the bloom that developed in response to deep water addition. Such CO2-related shifts in plankton community composition could have consequences for ecosystem productivity, biomass transfer to higher trophic levels, and biogeochemical element cycling of oligotrophic ocean regions.

Ocean acidification effects on mesozooplankton community development: Results from a long-term mesocosm experiment

Ocean acidification may affect zooplankton directly by decreasing in pH, as well as indirectly via trophic pathways, where changes in carbon availability or pH effects on primary producers may cascade up the food web thereby altering ecosystem functioning and community composition. Here, we present results from a mesocosm experiment carried out during 113 days in the Gullmar Fjord, Skagerrak coast of Sweden, studying plankton responses to predicted end-of-century pCO2 levels. We did not observe any pCO2 effect on the diversity of the mesozooplankton community, but a positive pCO2 effect on the total mesozooplankton abundance. Furthermore, we observed species-specific sensitivities to pCO2 in the two major groups in this experiment, copepods and hydromedusae. Also stage-specific pCO2 sensitivities were detected in copepods, with copepodites being the most responsive stage. Focusing on the most abundant species, Pseudocalanus acuspes, we observed that copepodites were significantly more abundant in the high-pCO2 treatment during most of the experiment, probably fuelled by phytoplankton community responses to high-pCO2 conditions. Physiological and reproductive output was analysed on P. acuspes females through two additional laboratory experiments, showing no pCO2 effect on females’ condition nor on egg hatching. Overall, our results suggest that the Gullmar Fjord mesozooplankton community structure is not expected to change much under realistic end-of-century OA scenarios as used here. However, the positive pCO2 effect detected on mesozooplankton abundance could potentially affect biomass transfer to higher trophic levels in the future.

Multidecadal accumulation of anthropogenic and remineralized dissolved inorganic carbon along the Extended Ellett Line in the northeast Atlantic Ocean

Marine carbonate chemistry measurements have been carried out annually since 2009 during UK research cruises along the Extended Ellett Line (EEL), a hydrographic transect in the northeast Atlantic Ocean. The EEL intersects several water masses that are key to the global thermohaline circulation, and therefore the cruises sample a region in which it is critical to monitor secular physical and biogeochemical changes. We have combined results from these EEL cruises with existing quality-controlled observational data syntheses to produce a hydrographic time series for the EEL from 1981 to 2013. This reveals multidecadal increases in dissolved inorganic carbon (DIC) throughout the water column, with a near-surface maximum rate of 1.80 ± 0.45 μmol kg -1 yr -1 . Anthropogenic CO 2 accumulation was assessed, using simultaneous changes in apparent oxygen utilization (AOU) and total alkalinity (TA) as proxies for the biogeochemical processes that influence DIC. The stable carbon isotope composition of DIC (δ 13 C DIC ) was also determined and used as an independent test of our method. We calculated a volume-integrated anthropogenic CO 2 accumulation rate of 2.8 ± 0.4 mg C m -3 yr -1 along the EEL, which is about double the global mean. The anthropogenic CO 2 component accounts for only 31 ± 6% of the total DIC increase. The remainder is derived from increased organic matter remineralization, which we attribute to the lateral redistribution of water masses that accompanies subpolar gyre contraction. Output from a general circulation ecosystem model demonstrates that spatiotemporal heterogeneity in the observations has not significantly biased our multidecadal rate of change calculations and indicates that the EEL observations have been tracking distal changes in the surrounding North Atlantic and Nordic Seas.

High CO2 Under Nutrient Fertilization Increases Primary Production and Biomass in Subtropical Phytoplankton Communities: A Mesocosm Approach

The subtropical oceans are home to one of the largest ecosystems on Earth, contributing to nearly one third of global oceanic primary production. Ocean warming leads to enhanced stratification in the oligotrophic ocean but also intensification in cross-shore wind gradients and thus in eddy kinetic energy across eastern boundary regions of the subtropical gyres. Phytoplankton thriving in a future warmer oligotrophic subtropical ocean with enhanced CO2 levels could therefore be patchily fertilized by increased mesoscale and submesoscale variability inducing nutrient pumping into the surface ocean. Under this premise, we have tested the response of three size classes (0.2-2, 2-20, and > 20 μm) of subtropical phytoplankton communities in terms of primary production, chlorophyll and cell biomass, to increasing CO2 concentrations and nutrient fertilization during an in situ mesocosm experiment in oligotrophic waters offof the island of Gran Canaria. We found no significant CO2-related effect on primary production and biomass under oligotrophic conditions (phase I). In contrast, primary production, chlorophyll and biomass displayed a significant and pronounced increase under elevated CO2 conditions in all groups after nutrient fertilization, both during the bloom (phase II) and post-bloom (phase III) conditions. Although the relative increase of primary production in picophytoplankton (250%) was 2.5 higher than in microphytoplankton (100%) after nutrient fertilization, comparing the high and low CO2 treatments, microphytoplankton dominated in terms of biomass, contributing > 57% to the total. These results contrast with similar studies conducted in temperate and cold waters, where consistently small phytoplankton benefitted after nutrient additions at high CO2, pointing to different CO2-sensitivities across plankton communities and ecosystem types in the ocean.

Enhanced transfer of organic matter to higher trophic levels caused by ocean acidification and its implications for export production: A mass balance approach

Ongoing acidification of the ocean through uptake of anthropogenic CO2 is known to affect marine biota and ecosystems with largely unknown consequences for marine food webs. Changes in food web structure have the potential to alter trophic transfer, partitioning, and biogeochemical cycling of elements in the ocean. Here we investigated the impact of realistic end-of-the-century CO2 concentrations on the development and partitioning of the carbon, nitrogen, phosphorus, and silica pools in a coastal pelagic ecosystem (Gullmar Fjord, Sweden). We covered the entire winter-to-summer plankton succession (100 days) in two sets of five pelagic mesocosms, with one set being CO2 enriched (~760 μatm pCO2) and the other one left at ambient CO2 concentrations. Elemental mass balances were calculated and we highlight important challenges and uncertainties we have faced in the closed mesocosm system. Our key observations under high CO2 were: (1) A significantly amplified transfer of carbon, nitrogen, and phosphorus from primary producers to higher trophic levels, during times of regenerated primary production. (2) A prolonged retention of all three elements in the pelagic food web that significantly reduced nitrogen and phosphorus sedimentation by about 11 and 9%, respectively. (3) A positive trend in carbon fixation (relative to nitrogen) that appeared in the particulate matter pool as well as the downward particle flux. This excess carbon counteracted a potential reduction in carbon sedimentation that could have been expected from patterns of nitrogen and phosphorus fluxes. Our findings highlight the potential for ocean acidification to alter partitioning and cycling of carbon and nutrients in the surface ocean but also show that impacts are temporarily variable and likely depending upon the structure of the plankton food web.

Abundance of the iron containing biomolecule, heme b, during the progression of a spring phytoplankton bloom in a mesocosm experiment

Concentrations of heme b were determined in a mesocosm experiment situated in Gullmar Fjord off Sweden. The mesocosm experiment lasted for ca. one hundred days and was characterised by the growth of a primary nutrient replete and a secondary nutrient deplete phytoplankton bloom. Heme b varied between 40 ± 10 pmol L-1 in the prebloom period up to a maximum of 700 ± 400 pmol L-1 just prior to the time of the primary chlorophyll a maximum. Thereafter, heme b concentrations decreased again to an average of 120 ± 60 pmol L-1. When normalised to total particulate carbon, heme b was most abundant during the initiation of the nutrient replete spring bloom, when ratios reached 52 ± 24 μmol mol-1; ten times higher than values observed both pre and post the primary bloom. Concentrations of heme b correlated with those of chlorophyll a. Nevertheless, differences were observed in the relative concentrations of the two parameters, with heme b concentrations increasing relative to chlorophyll a during the growth of the primary bloom, decreasing over the period of the secondary bloom and increasing again through the latter period of the experiment. Heme b abundance was therefore influenced by nutrient concentrations and also likely by changing community composition. In half of the mesocosms, pCO2 was elevated and maintained at ca.1000 μatm, however we observed no significant differences between heme b in plus or ambient pCO2 mesocosms, either in absolute terms, or relative to total particulate carbon and chlorophyll a. The results obtained in this study contribute to our understanding of the distribution of this significant component of the biogenic iron pool, and provide an iron replete coastal water end member that aids the interpretation of the distributions of heme b in more iron deplete open ocean waters.