Posts Tagged 'prokaryotes'

Differential responses of dominant and rare epiphytic bacteria from a submerged macrophyte to elevated CO2

Epiphytic bacteria develop complex interactions with their host macrophytes and play an important role in the ecological processes in freshwater habitats. However, how dominant and rare taxa respond to elevated atmospheric CO2 remains unclear. A manipulated experiment was carried out to explore the effects of elevated CO2 on the diversity or functional characteristics of leaf epiphytic dominant and rare bacteria from a submerged macrophyte. Three levels (high, medium, normal) of dissolved inorganic carbon (DIC) were applied to the overlying water. The physicochemical properties of the overlying water were measured. Elevated atmospheric CO2 significantly decreased the pH and dissolved oxygen (DO) of overlying water. Proteobacteria, Cyanobacteria, Bacteroidetes, Planctomycetes, and Actinobacteria are the dominant phyla of leaf epiphytic bacteria from Myriophyllum spicatum, occupying over 90% of the accumulated relative abundances. The aquatic DIC level and further pH significantly drove the epiphytic community composition differences among the three DIC levels. For dominant epiphytic bacteria, the functional potential of nutrient processes and mutualistic relationships were strongly affected by a high DIC level, while responses of rare epiphytic bacteria were more related to trace element processes, pathogens, and defense strategies under a high DIC level. Our results showed the responses of epiphytic bacteria to elevated CO2 varied across dominant and rare taxa.

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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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Transformations of diatom-derived dissolved organic matter by Bacillus pumilus under warming and acidification conditions

Heterotrophic bacteria are assumed to play an important role in processing of phytoplankton-derived dissolved organic matter (DOM). Although the algae-derived organic matter is commonly studied, the transformation and processing of DOM by epiphytic bacteria for phytoplankton have rarely been investigated, especially under warming and acidification. In this study, Bacillus pumilus is used to explore the ecologically important marine diatom Skeletonema dohrnii-derived DOM under different conditions (temperature, 27°C and 31°C; pCO2, 400 and 1,000 ppm), utilizing fluorescence excitation-emission matrix (EEM) combined with parallel factor analysis (EEM-PARAFAC). Fluorescence regional integration and the peak selecting method are used to generate B, T, N, A, M, and C peaks in the EEM fluorescence spectroscopy. The main known fluorophores including that protein-like components (peaks B and T), unknown components (peak N), and humic-like component (peaks A, M, and C). Our experimental results showed that under higher temperature and pressure of CO2 (pCO2) conditions, S. dohrnii-derived DOM fluorescence was dominated by a protein-like signal that slower waning throughout the experiment, becoming an increasingly humic-like substance, implying that processing by the epiphytic bacteria (B. pumilus) produced more complex molecules. In addition, spectroscopic indices (e.g., fluorescence index, biological index, freshness index β/α, and humification index) were changed in varying degrees. This study reveals and confirms the direct participation of heterotrophic bacteria in the transformation and generation of algae-derived DOM in the laboratory, underlining the influence of global warming and ocean acidification on this process.

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Microbial biofilms along a geochemical gradient at the shallow-water hydrothermal system of Vulcano Island, Mediterranean Sea

Shallow water hydrothermal vents represent highly dynamic environments where strong geochemical gradients can shape microbial communities. Recently, these systems are being widely used for investigating the effects of ocean acidification on biota as vent emissions can release high CO2 concentrations causing local pH reduction. However, other gas species, as well as trace elements and metals, are often released in association with CO2 and can potentially act as confounding factors. In this study, we evaluated the composition, diversity and inferred functional profiles of microbial biofilms in Levante Bay (Vulcano Island, Italy, Mediterranean Sea), a well-studied shallow-water hydrothermal vent system. We analyzed 16S rRNA transcripts from biofilms exposed to different intensity of hydrothermal activity, following a redox and pH gradient across the bay. We found that elevated CO2 concentrations causing low pH can affect the response of bacterial groups and taxa by either increasing or decreasing their relative abundance. H2S proved to be a highly selective factor shaping the composition and affecting the diversity of the community by selecting for sulfide-dependent, chemolithoautotrophic bacteria. The analysis of the 16S rRNA transcripts, along with the inferred functional profile of the communities, revealed a strong influence of H2S in the southern portion of the study area, and temporal succession affected the inferred abundance of genes for key metabolic pathways. Our results revealed that the composition of the microbial assemblages vary at very small spatial scales, mirroring the highly variable geochemical signature of vent emissions and cautioning for the use of these environments as models to investigate the effects of ocean acidification on microbial diversity.

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Modelling antifouling compounds of macroalgal holobionts in current and future pH conditions

Marine macroalgae are important ecosystem engineers in marine coastal habitats. Macroalgae can be negatively impacted through excessive colonization by harmful bacteria, fungi, microalgae, and macro-colonisers and thus employ a range of chemical compounds to minimize such colonization. Recent research suggests that environmental pH conditions potentially impact the functionality of such chemical compounds. Here we predict if and how naturally fluctuating pH conditions and future conditions caused by ocean acidification will affect macroalgal (antifouling) compounds and thereby potentially alter the chemical defence mediated by these compounds. We defined the relevant ecological pH range, analysed and scored the pH-sensitivity of compounds with antifouling functions based on their modelled chemical properties before assessing their distribution across the phylogenetic macroalgal groups, and the proportion of sensitive compounds for each investigated function. For some key compounds, we also predicted in detail how the associated ecological function may develop across the pH range. The majority of compounds were unaffected by pH, but compounds containing phenolic and amine groups were found to be particularly sensitive to pH. Future pH changes due to predicted average open ocean acidification pH were found to have little effect. Compounds from Rhodophyta were mainly pH-stable. However, key algal species amongst Phaeophyceae and Chlorophyta were found to rely on highly pH-sensitive compounds for their chemical defence against harmful bacteria, microalgae, fungi, and biofouling by macro-organisms. All quorum sensing disruptive compounds were found the be unaffected by pH, but the other ecological functions were all conveyed in part by pH-sensitive compounds. For some ecological keystone species, all of their compounds mediating defence functions were found to be pH-sensitive based on our calculations, which may not only affect the health and fitness of the host alga resulting in host breakdown but also alter the associated ecological interactions of the macroalgal holobiont with micro and macrocolonisers, eventually causing ecosystem restructuring and the functions (e.g. habitat provision) provided by macroalgal hosts. Our study investigates a question of fundamental importance because environments with fluctuating or changing pH are common and apply not only to coastal marine habitats and estuaries but also to freshwater environments or terrestrial systems that are subject to acid rain. Hence, whilst warranting experimental validation, this investigation with macroalgae as model organisms can serve as a basis for future investigations in other aquatic or even terrestrial systems.

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Can heat shock protein 70 (HSP70) serve as biomarkers in Antarctica for future ocean acidification, warming and salinity stress?

The Antarctic Peninsula is one of the fastest-warming places on Earth. Elevated sea water temperatures cause glacier and sea ice melting. When icebergs melt into the ocean, it “freshens” the saltwater around them, reducing its salinity. The oceans absorb excess anthropogenic carbon dioxide (CO2) causing decline in ocean pH, a process known as ocean acidification. Many marine organisms are specifically affected by ocean warming, freshening and acidification. Due to the sensitivity of Antarctica to global warming, using biomarkers is the best way for scientists to predict more accurately future climate change and provide useful information or ecological risk assessments. The 70-kilodalton (kDa) heat shock protein (HSP70) chaperones have been used as biomarkers of stress in temperate and tropical environments. The induction of the HSP70 genes (Hsp70) that alter intracellular proteins in living organisms is a signal triggered by environmental temperature changes. Induction of Hsp70 has been observed both in eukaryotes and in prokaryotes as response to environmental stressors including increased and decreased temperature, salinity, pH and the combined effects of changes in temperature, acidification and salinity stress. Generally, HSP70s play critical roles in numerous complex processes of metabolism; their synthesis can usually be increased or decreased during stressful conditions. However, there is a question as to whether HSP70s may serve as excellent biomarkers in the Antarctic considering the long residence time of Antarctic organisms in a cold polar environment which appears to have greatly modified the response of heat responding transcriptional systems. This review provides insight into the vital roles of HSP70 that make them ideal candidates as biomarkers for identifying resistance and resilience in response to abiotic stressors associated with climate change, which are the effects of ocean warming, freshening and acidification in Antarctic organisms.

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Proteome and microbiota analyses characterizing dynamic coral-algae-microbe tripartite interactions under simulated rapid ocean acidification


  • pH changes had a significant effect on the coral proteome- and 16S-profiling.
  • Maintenance of coral-algae-microbe interactions is a mechanism in coping with OA.
  • Proteome analysis identified some core biological pathways in OA early responses.
  • OA influences the microbial community, potentially compromising holobiont fitness.


Ocean acidification (OA) is a pressing issue currently and in the future for coral reefs. The importance of maintenance interactions among partners of the holobiont association in the stress response is well appreciated; however, the candidate molecular and microbial mechanisms that underlie holobiont stress resilience or susceptibility remain unclear. Here, to assess the effects of rapid pH change on coral holobionts at both the protein and microbe levels, combined proteomics and microbiota analyses of the scleractinian coral Galaxea fascicularis exposed to three relevant OA scenarios, including current (pHT = 8.15), preindustrial (pHT = 8.45) and future IPCC-2100 scenarios (pHT = 7.85), were conducted. The results demonstrated that pH changes had no significant effect on the physiological calcification rate of G. fascicularis in a 10-day experiment; however, significant differences were recorded in the proteome and 16S profiling. Proteome variance analysis identified some of the core biological pathways in coral holobionts, including coral host infection and immune defence, and maintaining metabolic compatibility involved in energy homeostasis, nutrient cycling, antibiotic activity and carbon budgets of coral-Symbiodiniaceae interactions were key mechanisms in the early OA stress response. Furthermore, microbiota changes indicate substantial microbial community and functional disturbances in response to OA stress, potentially compromising holobiont health and fitness. Our results may help to elucidate many complex mechanisms to describe scleractinian coral holobiont responses to OA and raise interesting questions for future studies.

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Carbonate chemistry in the microenvironment within cyanobacterial aggregates under present-day and future pCO2 levels

Photosynthesis and respiration cause distinct chemical microenvironments within cyanobacterial aggregates. Here, we used microsensors and a diffusion–reaction model to characterize gradients in carbonate chemistry and investigate how these are affected by ocean acidification in Baltic vs. Pacific aggregates (Nodularia and Dolichospermum vs. Trichodesmium). Microsensor measurements of O2 and pH were performed under in situ and expected future pCO2 levels on Nodularia and Dolichospermum aggregates collected in the Baltic Sea. Under in situ conditions, O2 and pH levels within the aggregates covered ranges of 80–175% air saturation and 7.7–9.4 in dark and light, respectively. Carbon uptake in the light was predicted to reduce HCO3 by 100–150 μmol L−1 and CO2 by 3–6 μmol L−1 in the aggregate center compared to outside, inducing strong CO2 depletion (down to 0.5 μmol L−1 CO2 remaining in the center) even when assuming that HCO3 covered 80–90% of carbon uptake. Under ocean acidification conditions, enhanced CO2 availability allowed for significantly lower activity of carbon concentrating mechanisms, including a reduction of the contribution of HCO3 to carbon uptake by up to a factor of 10. The magnification of proton gradients under elevated pCO2 that was predicted based on a lower buffer capacity was observed in measurements despite a concurrent decrease in photosynthetic activity. In summary, we provide a quantitative image of the inorganic carbon environment in cyanobacterial aggregates under present-day and expected future conditions, considering both the individual and combined effects of the chemical and biological processes that shape these environments.

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Response of microbial communities on culturing plates of post-settlement sea cucumbers to seawater acidification and warming

Seawater acidification and warming have been found to affect the early life of many marine organisms, but their effects on the microbial community in the environment related to the early development stage of aquaculture species have been rarely investigated. To understand how seawater acidification and warming impact the microbial community in aquaculture systems, we designed four microcosms to monitor and characterize the microbial composition on the corrugated plates in the Apostichopus japonicus culture tanks during its post-settlement stage. High-throughput 16S rRNA sequencing revealed that the bacterial community composition varied significantly in different periods of incubation. The bacterial diversity and community composition were obviously changed by seawater acidification and warming in the early period and then tended to revert to the level of the control group. Acidification significantly increased the relative abundance of dominant families Rhodobacteraceae and Flavobacteriaceae in the early period, suggesting that microbiota could increase the abundance of predominant taxa to adapt to increased CO2 concentration and reconstruct a stable community structure. No interaction effect of both factors was observed in the combined group. Results reveal that the microbial communities on the corrugated plates in A. japonicus culture tank were affected in the early period of incubation, and could then acclimatize to the increased CO2 and temperature. This study provides new insights into the variation and adaptation responses of the microbiota in aquaculture systems to seawater acidification and warming.

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Effects of ocean acidification on resident and active microbial communities of Stylophora pistillata

Ocean warming and ocean acidification (OA) are direct consequences of climate change and affect coral reefs worldwide. While the effect of ocean warming manifests itself in increased frequency and severity of coral bleaching, the effects of ocean acidification on corals are less clear. In particular, long-term effects of OA on the bacterial communities associated with corals are largely unknown. In this study, we investigated the effects of ocean acidification on the resident and active microbiome of long-term aquaria-maintained Stylophora pistillata colonies by assessing 16S rRNA gene diversity on the DNA (resident community) and RNA level (active community). Coral colony fragments of S. pistillata were kept in aquaria for 2 years at four different pCO2 levels ranging from current pH conditions to increased acidification scenarios (i.e., pH 7.2, 7.4, 7.8, and 8). We identified 154 bacterial families encompassing 2,047 taxa (OTUs) in the resident and 89 bacterial families including 1,659 OTUs in the active communities. Resident communities were dominated by members of Alteromonadaceae, Flavobacteriaceae, and Colwelliaceae, while active communities were dominated by families Cyclobacteriacea and Amoebophilaceae. Besides the overall differences between resident and active community composition, significant differences were seen between the control (pH 8) and the two lower pH treatments (7.2 and 7.4) in the active community, but only between pH 8 and 7.2 in the resident community. Our analyses revealed profound differences between the resident and active microbial communities, and we found that OA exerted stronger effects on the active community. Further, our results suggest that rDNA- and rRNA-based sequencing should be considered complementary tools to investigate the effects of environmental change on microbial assemblage structure and activity.

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Insignificant response of bacterioplankton community to elevated pCO2 during a short-term microcosm experiment in a subtropical eutrophic coastal ecosystem

Ocean acidification, as one of the major consequences of global climate change, markedly affects multiple ecosystem functions in disparate marine environments from coastal habitats to the deep ocean. Evaluation of the responses of marine microbial community to the increasing partial pressure of CO2 (pCO2) is crucial to explore the microbe-driven biogeochemical processes in the future ocean. In this study, a microcosm incubation of eutrophic coastal water from Xiamen Bay under elevated pCO2 (about 1,000 μatm) and control (ambient air, about 380–410 μatm) conditions was conducted to investigate the effect of ocean acidification on the natural bacterioplankton community. During the 5-day incubation period, the chlorophyll a concentration and bacterioplankton abundance were not significantly affected by increased pCO2. Hierarchical clustering and non-metric multidimensional scaling analysis based on Bray-Curtis similarity among the bacterioplankton community derived from the 16S rRNA genes revealed an inconspicuous impact of elevated pCO2 on the bacterial community. During the incubation period, Proteobacteria, Bacteroidetes, Actinobacteria, Cyanobacteria, and Epsilonbacteraeota were predominant in all microcosms. Despite the distinct temporal variation in the composition of the bacterioplankton community during the experimental period, statistical analyses showed that no significant difference was found on bacterioplankton taxa between elevated pCO2 and control, indicating that the bacterioplankton at the population-level were also insensitive to elevated pCO2. Our results therefore suggest that the bacterioplankton communities in the fluctuating and eutrophic coastal ecosystems appear to be adaptable to the short-term elevated pCO2.

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Composition and dominance of edible and inedible phytoplankton predict responses of Baltic Sea summer communities to elevated temperature and CO2

Previous studies with Baltic Sea phytoplankton combining elevated seawater temperature with CO2 revealed the importance of size trait-based analyses, in particular dividing the plankton into edible (>5 and <100 µm) and inedible (<5 and >100 µm) size classes for mesozoopankton grazers. While the edible phytoplankton responded predominantly negative to warming and the inedible group stayed unaffected or increased, independent from edibility most phytoplankton groups gained from CO2. Because the ratio between edible and inedible taxa changes profoundly over seasons, we investigated if community responses can be predicted according to the prevailing composition of edible and inedible groups. We experimentally explored the combined effects of elevated temperatures and CO2 concentrations on a late-summer Baltic Sea community. Total phytoplankton significantly increased in response to elevated CO2 in particular in combination with temperature, driven by a significant gain of the inedible <5 µm fraction and large filamentous cyanobacteria. Large flagellates disappeared. The edible group was low as usual in summer and decreased with both factors due to enhanced copepod grazing and overall decline of small flagellates. Our results emphasize that the responses of summer communities are complex, but can be predicted by the composition and dominance of size classes and groups.

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Highly variable and non-complex diazotroph communities in corals from ambient and high CO2 environments

The ecological success of corals depends on their association with microalgae and a diverse bacterial assemblage. Ocean acidification (OA), among other stressors, threatens to impair host-microbial metabolic interactions that underlie coral holobiont functioning. Volcanic CO2 seeps offer a unique opportunity to study the effects of OA in natural reef settings and provide insight into the long-term adaptations under a low pH environment. Here we compared nitrogen-fixing bacteria (diazotrophs) associated with four coral species (Pocillopora damicornisGalaxea fascicularisAcropora secale, and Porites rus) collected from CO2 seeps at Tutum Bay (Papua New Guinea) with those from a nearby ambient CO2 site using nifH amplicon sequencing to characterize the effects of seawater pH on bacterial communities and nitrogen cycling. Diazotroph communities were of generally low diversity across all coral species and for both sampling sites. Out of a total of 25 identified diazotroph taxa, 14 were associated with P. damicornis, of which 9 were shared across coral species. None of the diazotroph taxa, however, were consistently found across all coral species or across all samples within a species pointing to a high degree of diazotroph community variability. Rather, the majority of sampled colonies were dominated by one or two diazotroph taxa of high relative abundance. Pocillopora damicornis and Galaxea fascicularis that were sampled in both environments showed contrasting community assemblages between sites. In P. damicornis, Gammaproteobacteria and Cyanobacteria were prevalent under ambient pCO2, while a single member of the family Rhodobacteraceae was present at high relative abundance at the high pCO2 site. Conversely, in G. fascicularis diazotroph communities were indifferent between both sites. Diazotroph community changes in response to OA seem thus variable within as well as between host species, potentially arguing for haphazard diazotroph community assembly. This warrants further research into the underlying factors structuring diazotroph community assemblages and their functional role in the coral holobiont.

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Microbiomes of an oyster are shaped by metabolism and environment

Microbiomes can both influence and be influenced by metabolism, but this relationship remains unexplored for invertebrates. We examined the relationship between microbiome and metabolism in response to climate change using oysters as a model marine invertebrate. Oysters form economies and ecosystems across the globe, yet are vulnerable to climate change. Nine genetic lineages of the oyster Saccostrea glomerata were exposed to ambient and elevated temperature and PCO2 treatments. The metabolic rate (MR) and metabolic by-products of extracellular pH and CO2 were measured. The oyster-associated bacterial community in haemolymph was characterised using 16 s rRNA gene sequencing. We found a significant negative relationship between MR and bacterial richness. Bacterial community composition was also significantly influenced by MR, extracellular CO2 and extracellular pH. The effects of extracellular CO2 depended on genotype, and the effects of extracellular pH depended on CO2 and temperature treatments. Changes in MR aligned with a shift in the relative abundance of 152 Amplicon Sequencing Variants (ASVs), with 113 negatively correlated with MR. Some spirochaete ASVs showed positive relationships with MR. We have identified a clear relationship between host metabolism and the microbiome in oysters. Altering this relationship will likely have consequences for the 12 billion USD oyster economy.

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Impact of dust addition on the metabolism of Mediterranean plankton communities and carbon export under present and future conditions of pH and temperature (update)

Although atmospheric dust fluxes from arid as well as human-impacted areas represent a significant source of nutrients to surface waters of the Mediterranean Sea, studies focusing on the evolution of the metabolic balance of the plankton community following a dust deposition event are scarce, and none were conducted in the context of projected future levels of temperature and pH. Moreover, most of the experiments took place in coastal areas. In the framework of the PEACETIME project, three dust-addition perturbation experiments were conducted in 300 L tanks filled with surface seawater collected in the Tyrrhenian Sea (TYR), Ionian Sea (ION) and Algerian basin (FAST) on board the R/V Pourquoi Pas? in late spring 2017. For each experiment, six tanks were used to follow the evolution of chemical and biological stocks, biological activity and particle export. The impacts of a dust deposition event simulated at their surface were followed under present environmental conditions and under a realistic climate change scenario for 2100 (ca. +3 C and −0.3 pH units). The tested waters were all typical of stratified oligotrophic conditions encountered in the open Mediterranean Sea at this period of the year, with low rates of primary production and a metabolic balance towards net heterotrophy. The release of nutrients after dust seeding had very contrasting impacts on the metabolism of the communities, depending on the station investigated. At TYR, the release of new nutrients was followed by a negative impact on both particulate and dissolved 14C-based production rates, while heterotrophic bacterial production strongly increased, driving the community to an even more heterotrophic state. At ION and FAST, the efficiency of organic matter export due to mineral/organic aggregation processes was lower than at TYR and likely related to a lower quantity/age of dissolved organic matter present at the time of the seeding and a smaller production of DOM following dust addition. This was also reflected by lower initial concentrations in transparent exopolymer particles (TEPs) and a lower increase in TEP concentrations following the dust addition, as compared to TYR. At ION and FAST, both the autotrophic and heterotrophic community benefited from dust addition, with a stronger relative increase in autotrophic processes observed at FAST. Our study showed that the potential positive impact of dust deposition on primary production depends on the initial composition and metabolic state of the investigated community. This impact is constrained by the quantity of nutrients added in order to sustain both the fast response of heterotrophic prokaryotes and the delayed one of primary producers. Finally, under future environmental conditions, heterotrophic metabolism was overall more impacted than primary production, with the consequence that all integrated net community production rates decreased with no detectable impact on carbon export, therefore reducing the capacity of surface waters to sequester anthropogenic CO2.

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Diel metabolism of Yellow Sea green tide algae alters bacterial community composition under in situ seawater acidification of coastal areas


  • Metabolism of algae mat leads to a diel pH and CO2 fluctuation in affected seawater.
  • Bacterial communities in diffusive boundary layer of the algae had a diel change.
  • Flavobacteriaceae was shown increased at night but sharp decreased at daytime.
  • Harmful algal bloom might influence coastal ocean acidification.


Ocean acidification in coastal seawaters is a complex process, with coastal pH being affected by numerous factors including watershed and biological processes that also support metabolically diverse bacterial communities. The world’s largest macroalgal blooms have occurred consecutively in the Yellow Sea over the last 13 years. In particular, algal mats formed by Yellow Sea green tides (YSGT) significantly influence coastal environments. Herein, we hypothesized that 1) inorganic carbonate chemistry in coastal areas is altered by diel metabolism of these giant algal mats and that 2) bacterial community composition in diffusive boundary layers might be altered along diel cycles due to algal mat metabolism. In situ studies indicated that algal mat metabolism led to changes in diel pH and CO2 in affected seawaters. Such metabolic activities could intensify diel pH fluctuations in algal mat diffusive boundary layers, as noted by pH fluctuations of 0.22 ± 0.01 units, and pCO2 fluctuations of 214.62 ± 29.37 μatm per day. In contrast, pH fluctuations of 0.11 ± 0.02 units and pCO2 fluctuations of 79.02 ± 42.70 μatm were noted in unaffected areas. Furthermore, the bacterial community composition associated with diffusive algal boundary layers, including those of ambient bacteria and epiphytic bacteria, exhibited diel changes, while endophytic bacterial communities were relatively stable. Flavobacteriaceae were particularly highly abundant taxa in the ambient and epiphytic bacterial communities and exhibited increased abundances at night but sharp decreases in abundances during daytime. Flavobacteriaceae are heterotrophic taxa that could contribute to coastal area acidification at night due to the transformation of organic carbon to inorganic carbon. These results provide new insights to understand the variability in coastal ocean acidification via harmful algal blooms while providing a framework for evaluating the effects of YSGT on costal carbon cycling.

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Ocean acidification alters the diversity and structure of oyster associated microbial communities


Host-associated microbial communities are fundamental to host physiology, yet it is unclear how these communities will respond to environmental disturbances. Here, we disentangle the environment-linked and host-linked effects of ocean acidification on oyster-associated microbial communities. We exposed adult oysters (Crassostrea virginica) to CO2-induced ocean acidification (400 vs. 2800 ppm) for 80 d. We measured the oyster extrapallial fluid pH and sampled the gills for microbial analysis at six time points. We found that different subsets of microbes were linked to acidification (n = 34 amplicon sequence variants [ASVs]) and to host response (n = 20 ASVs) with little overlap (n = 8 ASVs), suggesting that some members of the oyster microbiome were more responsive to environmental conditions while others were more tightly linked to host condition. Our results provide insight into which members of the oyster microbiome may contribute to the health and resistance of their host, and which members are the most vulnerable to changing environmental conditions.

Scientific Significance Statement

Understanding microbial responses to environmental disturbances is critical. However, in host-associated microbial communities, it is unclear whether microbial response to disturbance is linked to the environment, or if it is mediated via host response. We used Eastern oysters as a model to demonstrate that both environment- and host-linked factors influence the composition and structure of gill microbial communities exposed to ocean acidification. Remarkably, members of the microbiome linked directly to elevated pCO2 were different from those linked to the host’s physiological response. Disentangling the microbial community’s response to environmental disturbance from its response to the host’s reaction to that disturbance is essential to understand and predict the effect of global change drivers on host-associated microbial communities.

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Effects of seawater scrubbing on a microplanktonic community during a summer-bloom in the Baltic Sea


  • Effects of seawater scrubbing on a microplanktonic community were assessed.
  • Biovolume increased with increasing concentrations of scrubber discharge water.
  • Group-specific impacts were recorded.
  • pH alone could not explain the observed results.
  • Other stressors in the scrubber water were responsible for the observed effect.


The International Maritime Organization (IMO) has gradually applied stricter regulations on the maximum sulphur content permitted in marine fuels and from January 1, 2020, the global fuel sulphur limit was reduced from 3.5% to 0.5%. An attractive option for shipowners is to install exhaust gas cleaning systems, also known as scrubbers, and continue to use high sulphur fuel oil. In the scrubber, the exhausts are led through a fine spray of water, in which sulphur oxides are easily dissolved. The process results in large volumes of acidic discharge water, but while regulations are focused on sulphur oxides removal and acidification, other pollutants e.g. polycyclic aromatic hydrocarbons, metals and nitrogen oxides can be transferred from the exhausts to the washwater and discharged to the marine environment. The aim of the current study was to investigate how different treatments of scrubber discharge water (1, 3 and 10%) affect a natural Baltic Sea summer microplanktonic community. To resolve potential contribution of acidification from the total effect of the scrubber discharge water, “pH controls” were included where the pH of natural sea water was reduced to match the scrubber treatments. Biological effects (e.g. microplankton species composition, biovolume and primary productivity) and chemical parameters (e.g. pH and alkalinity) were monitored and analysed during 14 days of exposure. Significant effects were observed in the 3% scrubber treatment, with more than 20% increase in total biovolume of microplankton compared to the control group, and an even greater effect in the 10% scrubber treatment. Group-specific impacts were recorded where diatoms, flagellates incertae sedis, chlorophytes and ciliates increased in biovolume with increasing concentrations of scrubber water while no effect was recorded for cyanobacteria. In contrast, these effects was not observed in the “pH controls”, a suggestion that other parameters/stressors in the scrubber water were responsible for the observed effects.

Continue reading ‘Effects of seawater scrubbing on a microplanktonic community during a summer-bloom in the Baltic Sea’

Climate change influences chlorophylls and bacteriochlorophylls metabolism in hypersaline microbial mat


  • Higher chlorophyll derivatives concentration on acidification treatment
  • Lower chlorophyll a concentration on acidification treatment
  • Production of bound carbohydrates EPS on acidification treatment
  • No impact of warming and acidification on mat photosynthetic efficiency


This study aimed to determine the effect of the climatic change on the phototrophic communities of hypersaline microbial mats. Ocean acidification and warming were simulated alone and together on microbial mats placed into mesocosms. As expected, the temperature in the warming treatments increased by 4 °C from the initial temperature. Surprisingly, no significance difference was observed between the water pH of the different treatments despite of a decrease of 0.4 unit pH in the water reserves of acidification treatments. The salinity increased on the warming treatments and the dissolved oxygen concentration increased and was higher on the acidification treatments. A total of 37 pigments were identified belonging to chlorophylls, carotenes and xanthophylls families. The higher abundance of unknown chlorophyll molecules called chlorophyll derivatives was observed in the acidification alone treatment with a decrease in chlorophyll a abundance. This change in pigmentary composition was accompanied by a higher production of bound extracellular carbohydrates but didn’t affect the photosynthetic efficiency of the microbial mats. A careful analysis of the absorption properties of these molecules indicated that these chlorophyll derivatives were likely bacteriochlorophyll c contained in the chlorosomes of green anoxygenic phototroph bacteria. Two hypotheses can be drawn from these results: 1/ the phototrophic communities of the microbial mats were modified under acidification treatment leading to a higher relative abundance of green anoxygenic bacteria, or 2/ the highest availability of CO2 in the environment has led to a shift in the metabolism of green anoxygenic bacteria being more competitive than other phototrophs.

Continue reading ‘Climate change influences chlorophylls and bacteriochlorophylls metabolism in hypersaline microbial mat’

Cyanobacteria net community production in the Baltic Sea as inferred from profiling pCO2 measurements

Organic matter production by cyanobacteria blooms is a major environmental concern for the Baltic Sea, as it promotes the spread of anoxic zones. Partial pressure of carbon dioxide (pCO2) measurements carried out on Ships of Opportunity (SOOP) since 2003 have proven to be a powerful tool to resolve the carbon dynamics of the blooms in space and time. However, SOOP measurements lack the possibility to directly constrain depth-integrated net community production (NCP) in moles of carbon per surface area due to their restriction to the sea surface. This study tackles the knowledge gap through (1) providing an NCP best guess for an individual cyanobacteria bloom based on repeated profiling measurements of pCO2 and (2) establishing an algorithm to accurately reconstruct depth-integrated NCP from surface pCO2 observations in combination with modelled temperature profiles.

Goal (1) was achieved by deploying state-of-the-art sensor technology from a small-scale sailing vessel. The low-cost and flexible platform enabled observations covering an entire bloom event that occurred in July–August 2018 in the Eastern Gotland Sea. For the biogeochemical interpretation, recorded pCO2 profiles were converted to C∗T, which is the dissolved inorganic carbon concentration normalised to alkalinity. We found that the investigated bloom event was dominated by Nodularia and had many biogeochemical characteristics in common with blooms in previous years. In particular, it lasted for about 3 weeks, caused a C∗T drawdown of 90 µmol kg−1, and was accompanied by a sea surface temperature increase of 10 C. The novel finding of this study is the vertical extension of the C∗T drawdown up to the compensation depth located at around 12 m. Integration of the C∗T drawdown across this depth and correction for vertical fluxes leads to an NCP best guess of ∼1.2 mol m−2 over the productive period.

Addressing goal (2), we combined modelled hydrographical profiles with surface pCO2 observations recorded by SOOP Finnmaid within the study area. Introducing the temperature penetration depth (TPD) as a new parameter to integrate SOOP observations across depth, we achieve an NCP reconstruction that agrees to the best guess within 10 %, which is considerably better than the reconstruction based on a classical mixed-layer depth constraint.

Applying the TPD approach to almost 2 decades of surface pCO2 observations available for the Baltic Sea bears the potential to provide new insights into the control and long-term trends of cyanobacteria NCP. This understanding is key for an effective design and monitoring of conservation measures aiming at a Good Environmental Status of the Baltic Sea.

Continue reading ‘Cyanobacteria net community production in the Baltic Sea as inferred from profiling pCO2 measurements’

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