Diatom-mediated food web functioning under ocean artificial upwelling.

Enhancing ocean productivity by artificial upwelling is evaluated as a nature-based solution for food security and climate change mitigation. Fish production is intended through diatom-based plankton food webs as these are assumed to be short and efficient. However, our findings from mesocosm experiments on artificial upwelling in the oligotrophic ocean disagree with this classical food web model. Here, diatoms did not reduce trophic length and instead impaired the transfer of primary production to crustacean grazers and small pelagic fish. The diatom-driven decrease in trophic efficiency was likely mediated by changes in nutritional value for the copepod grazers. Whilst diatoms benefitted the availability of essential fatty acids, they also caused unfavorable elemental compositions via high carbon-to-nitrogen ratios (i.e. low protein content) to which the grazers were unable to adapt. This nutritional imbalance for grazers was most pronounced in systems optimized for CO2 uptake through carbon-to-nitrogen ratios well beyond Redfield. A simultaneous enhancement of fisheries production and carbon sequestration via artificial upwelling may thus be difficult to achieve given their opposing stoichiometric constraints. Our study suggest that food quality can be more critical than quantity to maximize food web productivity during shorter-term fertilization of the oligotrophic ocean.

The appendicularian Oikopleura dioica can enhance carbon export in a high CO2 ocean.

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.

Impact of green and blue-green light on the growth, pigment concentration, and fatty acid unsaturation in the microalga Monoraphidium braunii.

The spectral composition of light is an important factor for the metabolism of photosynthetic organisms. Several blue light-regulated metabolic processes have already been identified in the industrially relevant microalga Monoraphidium braunii. However, little is known about the spectral impact on this species' growth, fatty acid (FA), and pigment composition. In this study, M. braunii was cultivated under different light spectra (white light: 400-700 nm, blue light: 400-550 nm, green light: 450-600 nm, and red light: 580-700 nm) at 25°C for 96 h. The growth was monitored daily. Additionally, the FA composition, and pigment concentration was analyzed after 96 h. The highest biomass production was observed upon white light and red light irradiation. However, green light also led to comparably high biomass production, fueling the scientific debate about the contribution of weakly absorbed light wavelengths to microalgal biomass production. All light spectra (white, blue, and green) that comprised blue-green light (450-550 nm) led to a higher degree of FA unsaturation and a greater concentration of all identified pigments than red light. These results further contribute to the growing understanding that blue-green light is an essential trigger for maximized pigment concentration and FA unsaturation in green microalgae.

Respiration rate scales inversely with sinking speed of settling marine aggregates.

Sinking marine aggregates have been studied for a long time to understand their role in carbon sequestration. Traditionally, sinking speed and respiration rates have been treated as independent variables, but two recent papers suggest that there is a connection albeit in contrasting directions. Here we collected recently formed (<2 days old) aggregates from sediment traps mounted underneath mesocosms during two different experiments. The mesocosms were moored off Gran Canaria, Spain (~ 27.9 N; 15.4 E) in a coastal, sub-tropical and oligotrophic ecosystem. We determined the respiration rates of organisms (mainly heterotrophic prokaryotes) attached to aggregates sinking at different velocities. The average respiration rate of fast sinking aggregates (>100 m d-1) was 0.12 d-1 ± 0.08 d-1 (SD). Slower sinking aggregates (<50 m d-1) had on average higher (p <0.001) and more variable respiration rates (average 0.31 d-1 ± 0.16 d-1, SD). There was evidence that slower sinking aggregates had higher porosity than fast sinking aggregates, and we hypothesize that higher porosity increase the settlement area for bacteria and the respiration rate. These findings provide insights into the efficiency of the biological carbon pump and help resolve the apparent discrepancy in the recent studies of the correlation between respiration and sinking speed.

Enhanced silica export in a future ocean triggers global diatom decline.

Diatoms account for up to 40% of marine primary production1,2 and require silicic acid to grow and build their opal shell3. On the physiological and ecological level, diatoms are thought to be resistant to, or even benefit from, ocean acidification4-6. Yet, global-scale responses and implications for biogeochemical cycles in the future ocean remain largely unknown. Here we conducted five in situ mesocosm experiments with natural plankton communities in different biomes and find that ocean acidification increases the elemental ratio of silicon (Si) to nitrogen (N) of sinking biogenic matter by 17 ± 6 per cent under [Formula: see text] conditions projected for the year 2100. This shift in Si:N seems to be caused by slower chemical dissolution of silica at decreasing seawater pH. We test this finding with global sediment trap data, which confirm a widespread influence of pH on Si:N in the oceanic water column. Earth system model simulations show that a future pH-driven decrease in silica dissolution of sinking material reduces the availability of silicic acid in the surface ocean, triggering a global decline of diatoms by 13-26 per cent due to ocean acidification by the year 2200. This outcome contrasts sharply with the conclusions of previous experimental studies, thereby illustrating how our current understanding of biological impacts of ocean change can be considerably altered at the global scale through unexpected feedback mechanisms in the Earth system.

Blue-green light is required for a maximized fatty acid unsaturation and pigment concentration in the microalga Acutodesmus obliquus.

Blue-green light is known to maximize the degree of fatty acid (FA) unsaturation in microalgae. However, knowledge on the particular waveband responsible for this stimulation of FA desaturation and its impact on the pigment composition in microalgae remains limited. In this study, Acutodesmus obliquus was cultivated for 96 h at 15°C with different light spectra (380-700 nm, 470-700 nm, 520-700 nm, 600-700 nm, and dark controls). Growth was monitored daily, and qualitative characterization of the microalgal FA composition was achieved via gas chromatography coupled with electron impact ionization mass spectrometry (GC-EI/MS). Additionally, a quantitative analysis of microalgal pigments was performed using high-performance liquid chromatography with diode array detection (HPLC-DAD). Spectra that included wavelengths between 470 and 520 nm led to a significantly higher percentage of the polyunsaturated fatty acids (PUFA) 18:3 and 16:4, compared to all other light conditions. However, no significant differences between the red light cultivations and the heterotrophic dark controls were observed for the FA 18:3 and 16:4. These results indicate, that exclusively the blue-green light waveband between 470 and 520 nm is responsible for a maximized FA unsaturation in A. obliquus. Furthermore, the growth and production of pigments were impaired if blue-green light (380-520 nm) was absent in the light spectrum. This knowledge can contribute to achieving a suitable microalgal pigment and FA composition for industrial purposes and must be considered in spectrally selective microalgae cultivation systems.

Differences in adaptation to light and temperature extremes of Chlorella sorokiniana strains isolated from a wastewater lagoon.

Presently, two Chlorella sorokiniana strains sampled during summer (CS-S) and winter (CS-W) from a maturation pond and isolated by dominance were studied on their behavior on temperature and light extremes in batch experiments. Although both strains showed no differences in their tolerance of temperatures up to 45 °C, the growth rates, pigment contents and fatty acid compositions in response to PAR at 700 and 1,500 µmol m-2sec-1 differed. CS-W was less affected by photoinhibition and maintained constantly high growth rates. High radiation resulted in both strains in an equivalent decrease of chlorophyll a and accessory pigments indicating that the latter did not function as a light filter. PUFAS (18:3 and 16:3) increased in CS-W at high radiation by > 60% and decreased in CS-S by 8 %. Results indicate that CS-W is highly favorable for mass cultivation particularly in outdoors, in which diurnal variations of solar radiation occur.

Ocean acidification induces distinct metabolic responses in subtropical zooplankton under oligotrophic conditions and after simulated upwelling.

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.

Exploring biogeochemical and ecological redundancy in phytoplankton communities in the global ocean.

Climate-change-induced alterations of oceanic conditions will lead to the ecological niches of some marine phytoplankton species disappearing, at least regionally. How will such losses affect the ecosystem and the coupled biogeochemical cycles? Here, we couch this question in terms of ecological redundancy (will other species be able to fill the ecological roles of the extinct species) and biogeochemical redundancy (can other species replace their biogeochemical roles). Prior laboratory and field studies point to a spectrum in the degree of redundancy. We use a global three-dimensional computer model with diverse planktonic communities to explore these questions further. The model includes 35 phytoplankton types that differ in size, biogeochemical function and trophic strategy. We run two series of experiments in which single phytoplankton types are either partially or fully eliminated. The niches of the targeted types were not completely reoccupied, often with a reduction in the transfer of matter from autotrophs to heterotrophs. Primary production was often decreased, but sometimes increased due to reduction in grazing pressure. Complex trophic interactions (such as a decrease in the stocks of a predator's grazer) led to unexpected reshuffling of the community structure. Alterations in resource utilization may cause impacts beyond the regions where the type went extinct. Our results suggest a lack of redundancy, especially in the 'knock on' effects on higher trophic levels. Redundancy appeared lowest for types on the edges of trait space (e.g. smallest) or with unique competitive strategies. Though highly idealized, our modelling findings suggest that the results from laboratory or field studies often do not adequately capture the ramifications of functional redundancy. The modelled, often counterintuitive, responses-via complex food web interactions and bottom-up versus top-down controls-indicate that changes in planktonic community will be key determinants of future ocean global change ecology and biogeochemistry.

Oxidative stress and antioxidant defence responses in two marine copepods in a high CO2 experiment.

We collected samples for oxidative stress and antioxidants in a high CO2 mesocosm experiment for two weeks, focussing on two common crustacean copepods Calanus finmarchicus and Temora longicornis. The samples were collected during a field experiment campaign studying responses of plankton communities to future ocean acidification (OA), off the Norwegian coast south of Bergen. The main results showed that there were species-specific differences between Temora and Calanus, especially in antioxidant defences (glutathione system) and oxidative stress (lipid peroxidation and reduced:oxidised glutathione ratio). Regular monitoring of chlorophyll a and jellyfish abundances taking place during the field campaign revealed that both chl a and predators may have affected the eco-physiological response. Antioxidant and oxidative stress levels are known to respond sensitively to both the food quality and quantity and the predator pressure, apart from environmental (i.e., abiotic) changes. Calanus was more robust towards OA, perhaps due to its high tolerance to a wide range of vertical physical-chemical conditions. Both top-down and bottom-up factors seem to play a role for the outcome of copepod responses to future ocean acidification.

Ocean acidification increases domoic acid contents during a spring to summer succession of coastal phytoplankton.

Enrichment of the oceans with CO2 may be beneficial for some marine phytoplankton, including harmful algae. Numerous laboratory experiments provided valuable insights into the effects of elevated pCO2 on the growth and physiology of harmful algal species, including the production of phycotoxins. Experiments close to natural conditions are the next step to improve predictions, as they consider the complex interplay between biotic and abiotic factors that can confound the direct effects of ocean acidification. We therefore investigated the effect of ocean acidification on the occurrence and abundance of phycotoxins in bulk plankton samples during a long-term mesocosm experiment in the Gullmar Fjord, Sweden, an area frequently experiencing harmful algal blooms. During the experimental period, a total of seven phycotoxin-producing harmful algal genera were identified in the fjord, and in accordance, six toxin classes were detected. However, within the mesocosms, only domoic acid and the corresponding producer Pseudo-nitzschia spp. was observed. Despite high variation within treatments, significantly higher particulate domoic acid contents were measured in the mesocosms with elevated pCO2. Higher particulate domoic acid contents were additionally associated with macronutrient limitation. The risks associated with potentially higher phycotoxin levels in the future ocean warrants attention and should be considered in prospective monitoring strategies for coastal marine waters.

In situ growth and bioerosion rates of Lophelia pertusa in a Norwegian fjord and open shelf cold-water coral habitat.

Coral reef resilience depends on the balance between carbonate precipitation, leading to reef growth, and carbonate degradation, for example, through bioerosion. Changes in environmental conditions are likely to affect the two processes differently, thereby shifting the balance between reef growth and degradation. In cold-water corals estimates of accretion-erosion processes in their natural habitat are scarce and solely live coral growth rates were studied with regard to future environmental changes in the laboratory so far, limiting our ability to assess the potential of cold-water coral reef ecosystems to cope with environmental changes. In the present study, growth rates of the two predominant colour morphotypes of live Lophelia pertusa as well as bioerosion rates of dead coral framework were assessed in different environmental settings in Norwegian cold-water coral reefs in a 1-year in situ experiment. Net growth (in weight gain and linear extension) of live L. pertusa was in the lower range of previous estimates and did not significantly differ between inshore (fjord) and offshore (open shelf) habitats. However, slightly higher net growth rates were obtained inshore. Bioerosion rates were significantly higher on-reef in the fjord compared to off-reef deployments in- and offshore. Besides, on-reef coral fragments yielded a broader range of individual growth and bioerosion rates, indicating higher turnover in live reef structures than off-reef with regard to accretion-bioerosion processes. Moreover, if the higher variation in growth rates represents a greater variance in (genetic) adaptations to natural environmental variability in the fjord, inshore reefs could possibly benefit under future ocean change compared to offshore reefs. Although not significantly different due to high variances between replicates, growth rates of orange branches were consistently higher at all sites, while mortality was statistically significantly lower, potentially indicating higher stress-resistance than the less pigmented white phenotype. Comparing the here measured rates of net accretion of live corals (regardless of colour morphotype) with net erosion of dead coral framework gives a first estimate of the dimensions of both processes in natural cold-water coral habitats, indicating that calcium carbonate loss through bioerosion amounts to one fifth to one sixth of the production rates by coral calcification (disregarding accretion processes of other organisms and proportion of live and dead coral framework in a reef). With regard to likely accelerating bioerosion and diminishing growth rates of corals under ocean acidification, the balance of reef accretion and degradation may be shifted towards higher biogenic dissolution in the future.

Shift towards larger diatoms in a natural phytoplankton assemblage under combined high-CO2 and warming conditions.

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 ~4.8 °C and ~535 ± 25 µatm pCO2), a "High-CO2" treatment (ambient temperature and initially 1020 ± 45 µatm pCO2) and a "Greenhouse" treatment (~8.5 °C and initially 990 ± 60 µatm pCO2). Nutrient replete conditions prevailed at the beginning of the experiment and light was provided at in situ levels upon reaching pCO2 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.

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.

Food web changes under ocean acidification promote herring larvae survival.

Ocean acidification-the decrease in seawater pH due to rising CO2 concentrations-has been shown to lower survival in early life stages of fish and, as a consequence, the recruitment of populations including commercially important species. To date, ocean-acidification studies with fish larvae have focused on the direct physiological impacts of elevated CO2, but largely ignored the potential effects of ocean acidification on food web interactions. In an in situ mesocosm study on Atlantic herring (Clupea harengus) larvae as top predators in a pelagic food web, we account for indirect CO2 effects on larval survival mediated by changes in food availability. The community was exposed to projected end-of-the-century CO2 conditions (~760 µatm pCO2) over a period of 113 days. In contrast with laboratory studies that reported a decrease in fish survival, the survival of the herring larvae in situ was significantly enhanced by 19 ± 2%. Analysis of the plankton community dynamics suggested that the herring larvae benefitted from a CO2-stimulated increase in primary production. Such indirect effects may counteract the possible direct negative effects of ocean acidification on the survival of fish early life stages. These findings emphasize the need to assess the food web effects of ocean acidification on fish larvae before we can predict even the sign of change in fish recruitment in a high-CO2 ocean.

Rapid evolution of highly variable competitive abilities in a key phytoplankton species.

Climate change challenges plankton communities, but evolutionary adaptation could mitigate the potential impacts. Here, we tested with the phytoplankton species Emiliania huxleyi whether adaptation to a stressor under laboratory conditions leads to equivalent fitness gains in a more natural environment. We found that fitness advantages that had evolved under laboratory conditions were masked by pleiotropic effects in natural plankton communities. Moreover, new genotypes with highly variable competitive abilities evolved on timescales significantly shorter than climate change.

Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change-A review.

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy-making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process-oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science-based policy formulation.

Growth performance and survival of larval Atlantic herring, under the combined effects of elevated temperatures and CO2.

In the coming decades, environmental change like warming and acidification will affect life in the ocean. While data on single stressor effects on fish are accumulating rapidly, we still know relatively little about interactive effects of multiple drivers. Of particular concern in this context are the early life stages of fish, for which direct effects of increased CO2 on growth and development have been observed. Whether these effects are further modified by elevated temperature was investigated here for the larvae of Atlantic herring (Clupea harengus), a commercially important fish species. Over a period of 32 days, larval survival, growth in size and weight, and instantaneous growth rate were assessed in a crossed experimental design of two temperatures (10°C and 12°C) with two CO2 levels (400 µatm and 900 µatm CO2) at food levels mimicking natural levels using natural prey. Elevated temperature alone led to increased swimming activity, as well as decreased survival and instantaneous growth rate (Gi). The comparatively high sensitivity to elevated temperature in this study may have been influenced by low food levels offered to the larvae. Larval size, Gi and swimming activity were not affected by CO2, indicating tolerance of this species to projected "end of the century" CO2 levels. A synergistic effect of elevated temperature and CO2 was found for larval weight, where no effect of elevated CO2 concentrations was detected in the 12°C treatment, but a negative CO2 effect was found in the 10°C treatment. Contrasting CO2 effects were found for survival between the two temperatures. Under ambient CO2 conditions survival was increased at 12°C compared to 10°C. In general, CO2 effects were minor and considered negligible compared to the effect of temperature under these mimicked natural food conditions. These findings emphasize the need to include biotic factors such as energy supply via prey availability in future studies on interactive effects of multiple stressors.

Simulated ocean acidification reveals winners and losers in coastal phytoplankton.

The oceans absorb ~25% of the annual anthropogenic CO2 emissions. This causes a shift in the marine carbonate chemistry termed ocean acidification (OA). OA is expected to influence metabolic processes in phytoplankton species but it is unclear how the combination of individual physiological changes alters the structure of entire phytoplankton communities. To investigate this, we deployed ten pelagic mesocosms (volume ~50 m3) for 113 days at the west coast of Sweden and simulated OA (pCO2 = 760 µatm) in five of them while the other five served as controls (380 µatm). We found: (1) Bulk chlorophyll a concentration and 10 out of 16 investigated phytoplankton groups were significantly and mostly positively affected by elevated CO2 concentrations. However, CO2 effects on abundance or biomass were generally subtle and present only during certain succession stages. (2) Some of the CO2-affected phytoplankton groups seemed to respond directly to altered carbonate chemistry (e.g. diatoms) while others (e.g. Synechococcus) were more likely to be indirectly affected through CO2 sensitive competitors or grazers. (3) Picoeukaryotic phytoplankton (0.2-2 µm) showed the clearest and relatively strong positive CO2 responses during several succession stages. We attribute this not only to a CO2 fertilization of their photosynthetic apparatus but also to an increased nutrient competitiveness under acidified (i.e. low pH) conditions. The stimulating influence of high CO2/low pH on picoeukaryote abundance observed in this experiment is strikingly consistent with results from previous studies, suggesting that picoeukaryotes are among the winners in a future ocean.