Posts Tagged 'phytoplankton'

Global record of “ghost” nannofossils reveals plankton resilience to high CO2 and warming


Fossil imprints from oceans of the past

Ghosts of the past

The marine geological records of some past global warming events contain relatively few nannoplankton fossils, the lack which some interpret as being evidence of the impact of ocean acidification and/or related environmental factors on biocalcification. Slater et al. present a global record of imprint, or “ghost,” nannofossils throughout several of those intervals during the Jurassic and Cretaceous periods (see the Perspective by Henderiks). This finding implies that a literal interpretation of the fossil record can be misleading, and demonstrates that nannoplankton were more resilient to past warming events than traditional fossil evidence would suggest. —HJS


Predictions of how marine calcifying organisms will respond to climate change rely heavily on the fossil record of nannoplankton. Declines in calcium carbonate (CaCO3) and nannofossil abundance through several past global warming events have been interpreted as biocalcification crises caused by ocean acidification and related factors. We present a global record of imprint—or “ghost”—nannofossils that contradicts this view, revealing exquisitely preserved nannoplankton throughout an inferred Jurassic biocalcification crisis. Imprints from two further Cretaceous warming events confirm that the fossil records of these intervals have been strongly distorted by CaCO3 dissolution. Although the rapidity of present-day climate change exceeds the temporal resolution of most fossil records, complicating direct comparison with past warming events, our findings demonstrate that nannoplankton were more resilient to past events than traditional fossil evidence suggests.

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A competitive advantage of middle-sized diatoms from increasing seawater CO2

Diatoms, one of the most important phytoplankton groups, fulfill their carbon demand from seawater mainly by obtaining passively diffused carbon dioxide (CO2) and/or actively consuming intracellular energy to acquire bicarbonate (HCO3). An anthropogenically induced increase in seawater CO2 reduces the HCO3 requirement of diatoms, potentially saving intracellular energy and benefitting their growth. This effect is commonly speculated to be most remarkable in larger diatoms that are subject to a stronger limitation of CO2 supply because of their smaller surface-to-volume ratios. However, we constructed a theoretical model for diatoms and revealed a unimodal relationship between the simulated growth rate response (GRR, the ratio of growth rates under elevated and ambient CO2) and cell size, with the GRR peaking at a cell diameter of ∼7 μm. The simulated GRR of the smallest diatoms was low because the CO2 supply was nearly sufficient at the ambient level, while the decline of GRR from a cell diameter of 7 μm was simulated because the contribution of seawater CO2 to the total carbon demand greatly decreased and diatoms became less sensitive to CO2 increase. A collection of historical data in CO2 enrichment experiments of diatoms also showed a roughly unimodal relationship between maximal GRR and cell size. Our model further revealed that the “optimal” cell size corresponding to peak GRR enlarged with the magnitude of CO2 increase but diminished with elevating cellular carbon demand, leading to projection of the smallest optimal cell size in the equatorial Pacific upwelling zone. Last, we need to emphasize that the size-dependent effects of increasing CO2 on diatoms are multifaceted, while our model only considers the inorganic carbon supply from seawater and optimal allocation of intracellular energy. Our study proposes a competitive advantage of middle-sized diatoms and can be useful in projecting changes in the diatom community in the future acidified high-CO2 ocean.

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Global climate change and the Baltic Sea ecosystem: direct and indirect effects on species, communities and ecosystem functioning

Climate change has multiple effects on Baltic Sea species, communities and ecosystem functioning through changes in physical and biogeochemical environmental characteristics of the sea. Associated indirect and secondary effects on species interactions, trophic dynamics and ecosystem function are expected to be significant. We review studies investigating species-, population- and ecosystem-level effects of abiotic factors that may change due to global climate change, such as temperature, salinity, oxygen, pH, nutrient levels, and the more indirect biogeochemical and food web processes, primarily based on peer-reviewed literature published since 2010.

For phytoplankton, clear symptoms of climate change, such as prolongation of the growing season, are evident and can be explained by the warming, but otherwise climate effects vary from species to species and area to area. Several modelling studies project a decrease of phytoplankton bloom in spring and an increase in cyanobacteria blooms in summer. The associated increase in N:P ratio may contribute to maintaining the “vicious circle of eutrophication”. However, uncertainties remain because some field studies claim that cyanobacteria have not increased and some experimental studies show that responses of cyanobacteria to temperature, salinity and pH vary from species to species. An increase of riverine dissolved organic matter (DOM) may also decrease primary production, but the relative importance of this process in different sea areas is not well known. Bacteria growth is favoured by increasing temperature and DOM, but complex effects in the microbial food web are probable. Warming of seawater in spring also speeds up zooplankton growth and shortens the time lag between phytoplankton and zooplankton peaks, which may lead to decreasing of phytoplankton in spring. In summer, a shift towards smaller-sized zooplankton and a decline of marine copepod species has been projected.

In deep benthic communities, continued eutrophication promotes high sedimentation and maintains good food conditions for zoobenthos. If nutrient abatement proceeds, improving oxygen conditions will first increase zoobenthos biomass, but the subsequent decrease of sedimenting matter will disrupt the pelagic–benthic coupling and lead to a decreased zoobenthos biomass. In the shallower photic systems, heatwaves may produce eutrophication-like effects, e.g. overgrowth of bladderwrack by epiphytes, due to a trophic cascade. If salinity also declines, marine species such as bladderwrack, eelgrass and blue mussel may decline. Freshwater vascular plants will be favoured but they cannot replace macroalgae on rocky substrates. Consequently invertebrates and fish benefiting from macroalgal belts may also suffer. Climate-induced changes in the environment also favour establishment of non-indigenous species, potentially affecting food web dynamics in the Baltic Sea.

As for fish, salinity decline and continuing of hypoxia is projected to keep cod stocks low, whereas the increasing temperature has been projected to favour sprat and certain coastal fish. Regime shifts and cascading effects have been observed in both pelagic and benthic systems as a result of several climatic and environmental effects acting synergistically.

Knowledge gaps include uncertainties in projecting the future salinity level, as well as stratification and potential rate of internal loading, under different climate forcings. This weakens our ability to project how pelagic productivity, fish populations and macroalgal communities may change in the future. The 3D ecosystem models, food web models and 2D species distribution models would benefit from integration, but progress is slowed down by scale problems and inability of models to consider the complex interactions between species. Experimental work should be better integrated into empirical and modelling studies of food web dynamics to get a more comprehensive view of the responses of the pelagic and benthic systems to climate change, from bacteria to fish. In addition, to better understand the effects of climate change on the biodiversity of the Baltic Sea, more emphasis should be placed on studies of shallow photic environments.

The fate of the Baltic Sea ecosystem will depend on various intertwined environmental factors and on development of the society. Climate change will probably delay the effects of nutrient abatement and tend to keep the ecosystem in its “novel” state. However, several modelling studies conclude that nutrient reductions will be a stronger driver for ecosystem functioning of the Baltic Sea than climate change. Such studies highlight the importance of studying the Baltic Sea as an interlinked socio-ecological system.

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The biological uptake of dissolved iron in the changing Daya Bay, South China Sea: effect of pH and DO


  • Fe bioavailability affected by pH and DO regulates phytoplanktonic Fe uptake.
  • Nano-phytoplankton is more sensitive to the variation of seawater pH and DO.
  • Phytoplankton community tend to be miniaturized in the changing DYB.
  • Fe requirement in DYB goes higher accompanied with the phytoplankton miniaturization.
  • DYB is not an Fe-rich environment derived from the relative low Fe:C ratio.


The oceanic acidification and coastal hypoxia have potential to enhance biological uptake of dissolved iron (Fe) by phytoplankton. In this study, the Fe uptake rate (FeUR) in Daya Bay was significantly negatively correlated with pH and dissolved oxygen (DO) (r = −0.81 and −0.73, respectively, p < 0.001). In addition, binary regression (FeUR = −1.45 × pH − 0.10 × DO + 13.64) also indicated that both pH and DO played key roles in FeUR variations. As pH and DO decreased, Fe uptake by phytoplankton was promoted, and the contribution of nano-phytoplankton to Fe uptake increased significantly, while that of pico-FeUR decreased. These will result in the phytoplankton community to be miniaturized and Fe requirement of phytoplankton goes higher, thereby leading changes of phytoplankton composition and coastal ecosystem. This study helps to understand how Fe could affect the coastal ecosystem under the increasing anthropogenic influences.

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No evidence of altered relationship between diet and consumer fatty acid composition in a natural plankton community under combined climate drivers

Fatty acids (FA), especially polyunsaturated fatty acids (PUFA), are key biomolecules involved in immune responses, reproduction, and membrane fluidity. PUFA in marine environments are synthesized exclusively by primary producers. Therefore the FA composition of these organisms at the base of the food web (i.e., phytoplankton) and their primary consumers (i.e., zooplankton) are important determinants of the health and productivity of entire ecosystems as they are transferred to higher trophic levels. However, environmental conditions such as seawater pH and temperature, which are already changing in response to climate change and predicted to continue to change in the future, can affect the FA composition of phytoplankton and zooplankton at both the organismal and community level. During a 20 day mesocosm experiment, we tested the effect of ocean acidification alone and in combination with ocean warming on 1) the fatty acid composition of a natural prey community for zooplankton (i.e. phytoplankton and microzooplankton), 2) the fatty acid composition of zooplankton, and 3) the relationship between prey and consumer fatty acid compositions in coastal waters. Significant effects of the climate stressors were not detected in the fatty acid composition of the prey or the relationship between diet and consumer fatty acids. A significant decrease in C18:4n-3 (stearidonic acid) was observed in the zooplankton but not their diet, but understanding the mechanism behind this decrease and its potential biological implications requires further investigation. These results highlight the importance of multi-stressor investigations on dynamics and variability contained within natural coastal plankton communities.

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Species specific responses to grazer cues and acidification in phytoplankton- winners and losers in a changing world

Phytoplankton induce defensive traits in response to chemical alarm signals from grazing zooplankton. However, these signals are potentially vulnerable to changes in pH and it is not yet known how predator recognition may be affected by ocean acidification. We exposed four species of diatoms and one toxic dinoflagellate to future pCO2 levels, projected by the turn of the century, in factorial combinations with predatory cues from copepods (copepodamides). We measured the change in growth, chain length, silica content, and toxin content. Effects of increased pCO2 were highly species specific. The induction of defensive traits was accompanied by a significant reduction in growth rate in three out of five species. The reduction averaged 39% and we interpret this as an allocation cost associated with defensive traits. Copepodamides induced significant chain length reduction in three of the four diatom species. Under elevated pCO2 Skeletonema marinoi reduced silica content by 30% and in Alexandrium minutum the toxin content was reduced by 30%. Using copepodamides to induce defensive traits in the absence of direct grazing provides a straightforward methodology to assess costs of defense in microplankton. We conclude that copepodamide signalling system is likely robust to ocean acidification. Moreover, the variable responses of different taxa to ocean acidification suggest that there will be winners and losers in a high pCO2 world, and that ocean acidification may have structuring effects on phytoplankton communities.

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Schizosphaerella size and abundance variations across the Toarcian Oceanic Anoxic Event in the Sogno Core (Lombardy Basin, Southern Alps)


  • Schizospharella spp. size and abundance variations during the Jenkyns event.
  • Abundance drop caused by the failure of S. punctulata > 7 μm.
  • Size decrease due to the relative increase in abundance of small specimens.
  • Drop in abundance and size consequence of ocean acidification and global warming.
  • Presence of diagenetic crust diagnostic to distinguish S. punctulata from S. astraea


Abundance and size variations of nannofossil Schizosphaerella punctulata were quantified in the uppermost Pliensbachian–Lower Toarcian succession recovered with the Sogno Core (Lombardy Basin, Northern Italy). High-resolution nannofossil biostratigraphy and C-isotopic chemostratigraphy identified the Jenkyns Event within the Toarcian oceanic anoxic event (T-OAE) interval. Absolute abundances and morphometric changes of “small S. punctulata” (< 7 μm), S. punctulata (7–10 μm; 10–14 μm; > 14 μm) and “encrusted S. punctulata” (specimens with a fringing crust) show large fluctuations across the negative δ13C Jenkyns Event. The Schizosphaerella crisis is further characterized by a decrease in average valve size in the early–middle Jenkyns Event. The abundance fall was caused by the failure of S. punctulata specimens >7 μm and “encrusted S. punctulata” that along with the increased relative abundance of small specimens, produced the reduction of average dimensions also documented in the Lusitanian and Paris Basins, although with a diachronous inception. The average valve size from the Lombardy Basin is ~2 μm smaller than in these other basins. Hyperthermal conditions associated with excess CO2 and ocean acidification possibly forced the drastic reduction of S. punctulata abundance/size. In the pelagic succession of the Sogno Core there is a strong positive correlation between the S. punctulata (> 7 μm) absolute abundance/size and the CaCO3 content, with a negligible contribution by small specimens (< 7 μm). Encrusted specimens testify selective neomorphic processes: the diagenetic crust seems diagnostic to separate S. punctulata from S. astraea.

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Whole community and functional gene changes of biofilms on marine plastic debris in response to ocean acidification

Plastics are accumulating in the world’s oceans, while ocean waters are becoming acidified by increased CO2. We compared metagenome of biofilms on tethered plastic bottles in subtidal waters off Japan naturally enriched in CO2, compared to normal ambient CO2 levels. Extending from an earlier amplicon study of bacteria, we used metagenomics to provide direct insights into changes in the full range of functional genes and the entire taxonomic tree of life in the context of the changing plastisphere. We found changes in the taxonomic community composition of all branches of life. This included a large increase in diatom relative abundance across the treatments but a decrease in diatom diversity. Network complexity among families decreased with acidification, showing overall simplification of biofilm integration. With acidification, there was decreased prevalence of genes associated with cell–cell interactions and antibiotic resistance, decreased detoxification genes, and increased stress tolerance genes. There were few nutrient cycling gene changes, suggesting that the role of plastisphere biofilms in nutrient processes within an acidified ocean may not change greatly. Our results suggest that as ocean CO2 increases, the plastisphere will undergo broad-ranging changes in both functional and taxonomic composition, especially the ecologically important diatom group, with possible wider implications for ocean ecology.

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Synergistic effects of ocean acidification and warming on coral host, Symbiodinium and nutrients exchange-based symbioses indicated by metatranscriptome

Global climate changes e.g. ocean acidification and warming caused by anthropogenic emission of CO2 are the greatest global threat to coral reef ecosystems. However, compared with the knowledge of Symbiodinium, little is known about the synergistic effects of combined ocean acidification and warming on the coral host and coral-Symbiodinium symbioses. In this study, metatranscriptomic analysis was performed to reveal the response of coral host and its symbiotic Symbiodinium to acidification (A), warming (H) and combined acidification and acidification (AH), using branching A. valida and massive G. fascicularis as models in a laboratory simulation system. RNA-Seq-based differently expressed genes (DEGs), together with coral’s morphological change, suggested the synergistic effects of AH on the coral host and coral-Symbiodinium symbioses, e.g. photosynthesis inhibition and negative effect on nutrients exchange between the host and its Symbiodinium. Particularly, AH showed a far greater impact on coral host than on Symbiodinium. These findings provide novel insights into the molecular mechanism of coral holobionts’ response to future extreme ocean acidification and warming, meanwhile highlight the molecular evidence for the different tolerance of branching and massive corals to environmental changes.

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Spatiotemporal distribution and environmental control factors of halocarbons in the Yangtze River Estuary and its adjacent marine area during autumn and spring


  • The source of air masses influenced volatile halocarbons (VHCs) levels in the air.
  • Spatiotemporal variations of VHCs in seawater and atmosphere were investigated.
  • Seasonal variations in VHCs concentrations were dependent on complex factors.
  • Ocean acidification and dust addition had an impact on the production of VHCs.


The oceanic production and release of volatile halocarbons (VHCs) to the atmosphere play a vital role in regulating the global climate. In this study, seasonal and spatial variations in VHCs, including trichlorofluoromethane (CFC-11), methyl iodide (CH3I), dibromomethane (CH2Br2), and bromoform (CHBr3), and environmental parameters affecting their concentrations were characterized in the atmosphere and seawater of the Yangtze River Estuary and its adjacent marine area during two cruises from October 17 to October 26, 2019 and from May 12 to May 25, 2020. Significant seasonal variations were observed in the atmosphere and seawater because of seasonal differences in the prevalent monsoon, water mass (Yangtze River Diluted Water), and biogenic production. VHCs concentrations were positively correlated with Chl-a concentrations in the surface water during autumn. The average sea-to-air fluxes of CH3I, CH2Br2, and CHBr3 in autumn were 19.7, 4.0, and 7.6 nmol m−2 d−1, respectively, while those in spring were 6.3, 6.4, and −3.6 nmol m−2 d−1. In the ship-based incubation experiments, ocean acidification and dust deposition had no significant effects on VHCs concentrations. The concentrations of CH2Br2 and CHBr3 were significantly positively correlated with phytoplankton biomass under lower pH condition (M3: pH 7.9). This result indicated that CH2Br2 and CHBr3 concentrations were mainly related to the biological release.

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Sensitivity of phytoplankton to climate change: direct and interactive effects of CO2 on primary production and community composition

Marine phytoplankton constitutes about half of the primary production on Earth. It forms the base of the marine food web and is a pivotal player in the marine biological carbon pump. The primary environmental drivers that control phytoplankton growth are temperature, nutrient availability, light, and the concentration of inorganic carbon species. Ongoing climate change modifies these drivers, leading to a warming, high-CO2 ocean with altered nutrient availabilities and light regimes. Changes in phytoplankton productivity and community composition resulting from these newly emerging environmental states in the ocean have important implications for the marine ecosystem and carbon cycling.

Biogeochemical ocean models are used to investigate how marine primary production may be affected by future climate change under different emission scenarios. Phytoplankton growth rates in models are typically determined by functions describing growth dependencies on temperature, light, and nutrients. However, a large body of laboratory studies on phytoplankton responses to environmental drivers reveals two points that are usually not considered in current biogeochemical models. Firstly, phytoplankton growth can be considerably modified by the state of the carbonate system. Changes in inorganic carbon species concentrations can be either growth-enhancing (CO2(aq) and bicarbonate are substrates for photosynthesis), or growth-dampening (increasing CO2(aq) levels lead to a shift in the carbonate equilibria and result in a pH decrease, a process which is called ocean acidification). Functions describing this growth dependence of phytoplankton on the carbonate system have not been implemented in large-scale ocean biogeochemical models so far. Secondly, growth responses towards one driver can be modified if the level of another driver is changing. Functions including these so-called interactive driver effects partly exist in models (e.g. the response to varying light levels may depend on the nutrient limitation term). However, the large number of laboratory studies on multiple driver effects has never been used to constrain driver interactions in large-scale ocean biogeochemical models. This holds especially true for the findings of growth responses to driver interactions that include ocean acidification, which make up the largest share of laboratory experiments.

This thesis aims to investigate sensitivities of marine phytoplankton to changing CO2(aq) levels as well as to interactive effects between CO2 and other environmental drivers. A comprehensive and reproducible literature search in combination with a statistical analysis (Publication I) reveals that increasing CO2(aq) levels robustly dampen the growth-increasing effects of warming and improving light conditions. In addition, the results show that the calcifying phytoplankton group of coccolithophores experiences the strongest negative effects by ocean acidification compared to other phytoplankton groups. A second study (Publication II) examines the effects of mechanistically described carbonate system dependencies on primary production and community composition in a model. To this end, carbonate system dependencies of phytoplankton growth and and coccolithophore calcification are implemented into the global biogeochemical ocean model REcoM. The study shows that responses to ocean acidification cascade on growth responses to other drivers, which partly balance or counteract the direct impact of the carbonate system on growth rates. In addition, warming is identified as the main driver of the observed recent increase of coccolithophore biomass in the North Atlantic. A final study (Publication III) investigates the interactive effects between CO2 and temperature as well as between CO2 and light on phytoplankton biomass and community composition in a high emission scenario. For the parametrization in REcoM, growth responses to interacting drivers as synthesized in Publication I are used. The decrease of global future phytoplankton biomass and net community production by the end of the century is similar in simulations with and without driver interactions (-6% and -8%, respectively). However, phytoplankton responses to future climate conditions are considerably modified on a regional scale and the share of individual phytoplankton groups in the community changes both globally and regionally when accounting for multiple driver effects. Globally, diatoms and coccolithophores are impacted more and small phytoplankton less severely by future oceanic conditions when accounting for driver interactions. Future projections of the Southern Ocean phytoplankton community are modified most dramatically with the new interactive growth formulation, as diatoms and coccolithophores become less and small phytoplankton more abundant, while it is the other way round in simulations without driver interactions.

The thesis highlights 1) that the carbonate system is a critical growth-modifying driver for phytoplankton in a high-CO2 ocean, which can furthermore modify growth responses to other drivers substantially, and 2) that driver interactions have considerable effects on climate-change induced alterations in the phytoplankton community as well as on regional biomass changes in a future ocean.

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Physiological control on carbon isotope fractionation in marine phytoplankton

One of the great challenges in biogeochemical research over the past half a century has been to quantify and understand the mechanisms underlying stable carbon isotope fractionation (εp) in phytoplankton in response to changing pCO2. Partly, this interest is grounded in the use of fossil photosynthetic organism remains as a proxy for past atmospheric CO2 concentrations. Phytoplankton organic carbon is depleted in 13C compared to its source because of kinetic fractionation by the enzyme RubisCO during photosynthetic carbon fixation, as well as through physiological pathways upstream of RubisCO. Moreover, other factors such as nutrient limitation, variations in light regime as well as phytoplankton culturing systems and inorganic carbon manipulation approaches may confound the influence of CO2 on εp. Here, based on experimental data compiled from the literature, we assess which underlying physiological processes cause the observed differences in εp for various phytoplankton groups in response to C-demand/C-supply and test potential confounding factors. Culturing approaches and methods of carbonate chemistry manipulation were found to best explain the differences in εp between studies, although daylength was an important predictor for εp in haptophytes. Extrapolating results from culturing experiments to natural environments and for proxy applications therefore requires caution, and it should be carefully considered whether culture methods and experimental conditions are representative of natural environments.

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Ocean acidification stimulation of phytoplankton growth depends on the extent of departure from the optimal growth temperature


  • Growth enhancement by HC was most prominent at temperatures close to optimum.
  • Cellular POC and PON were significantly reduced in HC than in LC near optimum.
  • High CO2-grown cells endured more stress at high and low growth temperatures.


Ocean acidification and warming are two major environmental stressors; however, the generality of how warming will alter growth responses of phytoplankton to ocean acidification is less known. Here, enhancement of growth by high CO2 (HC) in Phaeodactylum tricornutum and Thalassiosira weissflogii was most prominent at optimum temperature. The extent to which growth rates in HC cultures were raised compared to low CO2 (LC) cultures tended to decrease with increasing or decreasing temperature, compared to the optimum. Further mechanistic studies in P. tricornutum revealed that cellular carbon and nitrogen content, superoxide dismutase activity, and respiration were generally higher in HC than those in LC at high and low temperatures, whereas PSII photochemical parameters were generally lower in HC than in LC at high and low temperatures. These results indicate that HC-grown cells needed to invest more energy and materials to maintain intracellular homeostasis and repair damage induced by the unsuitable temperatures.

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Abrupt upwelling and CO2 outgassing episodes in the north-eastern Arabian Sea since mid-Holocene

Identifying the causes and consequences of natural variations in ocean acidification and atmospheric CO2 due to complex earth processes has been a major challenge for climate scientists in the past few decades. Recent developments in the boron isotope (δ11B) based seawater pH and pCO2 (or pCO2sw) proxy have been pivotal in understanding the various oceanic processes involved in air-sea CO2 exchange. Here we present the first foraminifera-based δ11B record from the north-eastern Arabian Sea (NEAS) covering the mid-late Holocene (~ 8–1 ka). Our record suggests that the region was overall a moderate to strong CO2 sink during the last 7.7 kyr. The region behaved as a significant CO2 source during two short intervals around 5.5–4 ka and 2.8–2.5 ka. The decreased pH and increased CO2 outgassing during those abrupt episodes are associated with the increased upwelling in the area. The upwelled waters may have increased the nutrient content of the surface water through either increased supply or weaker export production. This new dataset from the coastal NEAS suggests that, as a potential result of changes in the strength of the El-Nino Southern Oscillation, the region experienced short episodes of high CO2 outgassing and pre-industrial ocean acidification comparable to or even greater than that experienced during the last ~ 200 years.

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Parallel between the isotopic composition of coccolith calcite and carbon levels across Termination II: developing a new paleo-CO2 probe

Beyond the pCO2 records provided by ice core measurements, the quantification of atmospheric CO2 concentrations and changes thereof relies on proxy data, the development of which represents a foremost challenge in paleoceanography. In the paleoceanographic toolbox, the coccolithophores occupy a notable place, as the magnitude of the carbon isotopic fractionation between ambient CO2 and a type of organic compounds that these photosynthetic microalgae synthesize (the alkenones) represents a relatively robust proxy to reconstruct past atmospheric CO2 concentrations during the Cenozoic. The isotopic composition of coeval calcite biominerals found in the sediments and also produced by the coccolithophores (the coccoliths) have been found to record an ambient CO2 signal through culture and sediment analyses. These studies have, however, not yet formalized a transfer function that quantitatively ties the isotopic composition of coccolith calcite to the concentrations of aqueous CO2 and, ultimately, to atmospheric CO2 levels. Here, we make use of a microseparation protocol to compare the isotopic response of two size-restricted coccolith assemblages from the North Atlantic to changes in surface ocean CO2 during Termination II (ca. 130–140 ka). Performing paired measurements of the isotopic composition (δ13C and δ18O) of relatively large and small coccoliths provides an isotopic offset that can be designated as a “differential vital effect”. We find that the evolution of this offset follows that of aqueous CO2 concentrations computed from the ice core CO2 curve and an independent temperature signal. We interpret this biogeochemical feature to be the result of converging carbon fixation strategies between large and small cells as the degree of carbon limitation for cellular growth decreases across the deglaciation. We are therefore able to outline a first-order trend between the coccolith differential vital effects and aqueous CO2 in the range of Quaternary CO2 concentrations. Although this study would benefit from further constraints on the other controls at play on coccolith geochemistry (growth rate, air–sea gas exchange, etc.), this test of the drivers of coccolith Δδ13C and Δδ18O in natural conditions is a new step in the development of a coccolith paleo-CO2 probe.

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In contrast to diatoms, cryptophytes are susceptible to iron limitation, but not to ocean acidification

Previous field studies in the Southern Ocean (SO) indicated an increased occurrence and dominance of cryptophytes over diatoms due to climate change. To gain a better mechanistic understanding of how the two ecologically important SO phytoplankton groups cope with ocean acidification (OA) and iron (Fe) availability, we chose two common representatives of Antarctic waters, the cryptophyte Geminigera cryophila and the diatom Pseudo-nitzschia subcurvata. Both species were grown at 2°C under different pCO2 (400 vs. 900 μatm) and Fe (0.6 vs. 1.2 nM) conditions. For P. subcurvata, an additional high pCO2 level was applied (1400 μatm). At ambient pCO2 under low Fe supply, growth of G. cryophila almost stopped while it remained unaffected in P. subcurvata. Under high Fe conditions, OA was not beneficial for P. subcurvata, but stimulated growth and carbon production of G. cryophila. Under low Fe supply, P. subcurvata coped much better with OA than the cryptophyte, but invested more energy into photoacclimation. Our study reveals that Fe limitation was detrimental for the growth of G. cryophila and suppressed the positive OA effect. The diatom was efficient in coping with low Fe, but was stressed by OA while both factors together strongly impacted its growth. The distinct physiological response of both species to OA and Fe limitation explains their occurrence in the field. Based on our results, Fe availability is an important modulator of OA effects on SO phytoplankton, with different implications on the occurrence of cryptophytes and diatoms in the future.

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Surface ocean warming and acidification driven by rapid carbon release precedes Paleocene-Eocene Thermal Maximum

The Paleocene-Eocene Thermal Maximum (PETM) is recognized by a major negative carbon isotope (δ13C) excursion (CIE) signifying an injection of isotopically light carbon into exogenic reservoirs, the mass, source, and tempo of which continue to be debated. Evidence of a transient precursor carbon release(s) has been identified in a few localities, although it remains equivocal whether there is a global signal. Here, we present foraminiferal δ13C records from a marine continental margin section, which reveal a 1.0 to 1.5‰ negative pre-onset excursion (POE), and concomitant rise in sea surface temperature of at least 2°C and a decline in ocean pH. The recovery of both δ13C and pH before the CIE onset and apparent absence of a POE in deep-sea records suggests a rapid (< ocean mixing time scales) carbon release, followed by recovery driven by deep-sea mixing. Carbon released during the POE is therefore likely more similar to ongoing anthropogenic emissions in mass and rate than the main CIE.

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An integrated multiple driver mesocosm experiment reveals the effect of global change on planktonic food web structure

Global change puts coastal marine systems under pressure, affecting community structure and functioning. Here, we conducted a mesocosm experiment with an integrated multiple driver design to assess the impact of future global change scenarios on plankton, a key component of marine food webs. The experimental treatments were based on the RCP 6.0 and 8.5 scenarios developed by the IPCC, which were Extended (ERCP) to integrate the future predicted changing nutrient inputs into coastal waters. We show that simultaneous influence of warming, acidification, and increased N:P ratios alter plankton dynamics, favours smaller phytoplankton species, benefits microzooplankton, and impairs mesozooplankton. We observed that future environmental conditions may lead to the rise of Emiliania huxleyi and demise of Noctiluca scintillans, key species for coastal planktonic food webs. In this study, we identified a tipping point between ERCP 6.0 and ERCP 8.5 scenarios, beyond which alterations of food web structure and dynamics are substantial.

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Rising CO2 will increase toxicity of marine dinoflagellate Alexandrium minutum


  • High level CO2 significantly promoted growth of toxic Alexandrium minutum.
  • Total yields of paralytic shellfish toxins by A. minutum were enhanced by rising CO2.
  • Rising CO2 promoted transformation from GTX2&3 to more potent GTX1&4.
  • High level CO2 may depress the release of PSTs from inside to outside of the cells.
  • All the processes collectively increase the risk of A. minutum under CO2 enrichment.


Ocean acidification caused by increasing emission of carbon dioxide (CO2) is expected to have profound impacts on marine ecological processes, including the formation and evolution of harmful algal blooms (HABs). We designed a set of experiments in the laboratory to examine the effects of increasing CO2 on the growth and toxicity of a toxic dinoflagellate Alexandrium minutum producing paralytic shellfish toxins (PSTs). It was found that high levels of CO2 (800 and 1,200 ppm) significantly promoted the growth of A. minutum compared to the group (400 ppm) representing the current CO2 level. The total yields of PSTs by A. minutum, including both intracellular and extracellular toxins, were significantly enhanced, probably due to the induction of core enzyme activity and key amino acids synthesis for PST production. More interestingly, high level of CO2 promoted the transformation from gonyautoxin2&3 to gonyautoxin1&4 and depressed the release of PSTs from inside to outside of the cells. All these processes collectively led to an apparent increase of A. minutum toxicity. Our study demonstrated that rising CO2 would increase the risk of toxic A. minutum based on the comprehensive analyses of different processes including algal growth and toxin synthesis, transformation and release.

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Effects of elevated pCO2 on the photosynthetic performance of the sea ice diatoms Navicula directa and Navicula glaciei

Sea ice algal communities are generally dominated by pennate diatoms, which commonly occur at the ice-water interface and in brine channels. They also make a significant contribution to higher trophic levels associated with sea ice habitats. Here, the photosynthetic responses of two sea ice diatom species, Navicula directa and Navicula glaciei, to changes in pCO2 under controlled laboratory conditions were compared. pCO2 (390 ppm and 750 ppm) was manipulated to simulate a shift from present levels (1990) to predicted “IPCC year 2100 worst-case scenario” levels. To investigate these effects, a pulse-amplitude modulation (PAM) fluorometer was used to measure the photosynthetic performance. The ability of the sea ice algae to grow and photosynthesize within physio-chemical gradients in the sea ice suggests that both sea ice species are likely to be well adapted to cope with changes in pCO2 concentrations. Lower pH and higher pCO2 for 7 days resulted in increased biomass, especially for N. directa. However, a decline in photosynthetic capacity (rETRmax) was observed for both species (highest value 11.375 ± 0.163, control; and 8.322 ± 1.282, treatment). Navicula glaciei showed significant effects of elevated pCO2 (p < 0.05) on its photosynthetic response, while N. directa did not. Future changes in CO2 and pH may thus not significantly affect all diatoms but may lead to changes in the photosynthetic activities in some species.

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