Diatoms constitute a major group of phytoplankton, accounting for ~20% of the world’s primary production. Biological dinitrogen (N2) fixation by diazotrophic cyanobacteria has great biogeochemical implications in nitrogen (N) cycling, being the major source of new N input to the oceans and thereby contributing significantly to carbon (C) export production. It has been shown that iron (Fe) can be the limiting nutrient for phytoplankton growth, in particular, in the HNLC (High Nutrient Low Chlorophyll) regions. Iron plays thus an essential role in governing the marine primary productivity and the efficiency of biological carbon pump. Oceanic systems are undergoing continuous modifications at varying rates and magnitudes as a result of changing climate. The objective of our research is to evaluate the effects of global climate change processes (changing dust deposition, ocean acidification and sea-surface warming) on phytoplankton growth, biological N2 fixation, biogeochemical cycles, and the controlling role of Fe within these impacts. Laboratory culture experiments using a marine diatom Chaetoceros socialis were conducted at two temperatures (13 ℃ and 18 ℃) and two carbon dioxide partial pressures (pCO2, 400 µatm and 800 µatm). The present study clearly highlights the effect of ocean acidification on enhancing the release of Fe upon dust deposition. Our results also confirm that being a potential source of Fe, mineral dust provides in addition a readily utilizable source of macronutrients such as phosphorus (P) and silicon (Si). However, elevated atmospheric CO2 concentrations and ocean acidification may also have an adverse impact on diatom growth, causing a decrease in cell size and possible further changes in phytoplankton composition. Meanwhile, increasing temperature and ocean warming may lead to the reduction of diatom production as well as cell size, inducing poleward shifts in the biogeographic distribution of diatoms. Numerous factors can affect the extent of N2 fixation. A better understanding of the major environmental and nutrient controls governing this process is highly required. Iron and/or phosphorus are thought to be limiting factors in most oceanic regions. Special attention has been given to studying the effects of mineral dust deposition which is believed to promote N2 fixation as it increases the availability of both Fe and P. Three laboratory bioassays (+Fe, +P, +Dust) via incubation experiments were performed on Trichodesmium IMS101, an important contributor to marine N2 fixation. Each addition of Fe, P or desert dust was found to stimulate the growth and the N2 fixation activity of Trichodesmium IMS101. Several adaptive nutrient utilization strategies were observed, such as a Fe luxury uptake mechanism, a P-sparing effect and colony formation. In addition, during a field study in the temperate Northeast Atlantic Ocean using natural phytoplankton assemblages, N2 fixation was remarkably stimulated through the addition of dissolved Fe under low temperature and depleted P conditions, highlighting the critical role of Fe. At the time of this study, no Trichodesmium filaments were found in the region of investigation. The diazotrophic community was dominated by the unicellular cyanobacteria symbiont (prymnesiophyte-UCYN-A1) and heterotrophic diazotrophs, therefore suggesting that Fe could be the ultimate factor limiting N2 fixation of these smaller diazotrophs as well. Recently, the effects of ongoing climate change (ocean warming and acidification) on N2 fixation drew much attention, but various studies led to controversial conclusions. Semi-continuous dilution growth experiments were conducted on Trichodesmium IMS101 under future high pCO2 and warming seawater conditions (800 µatm and 28 °C) and compared to the present-day situations (400 µatm and 24 °C). The results indicate that higher pCO2 and therefore ocean acidification may be beneficial for Trichodesmium growth and N2 fixation. However, the present study suggests that Fe or P limitation in oligotrophic seawaters may offset the stimulation induced on Trichodesmium IMS101 due to ocean acidification. In contrast, ocean warming may not play an important role in Trichodesmium growth and N2 fixation with a 4 °C increase from 24 °C to 28 °C. Nevertheless, ocean warming was previously predicted to cause a shift in the geographical distribution of Trichodesmium toward higher latitudes, extending its niche to subtropical regions and potentially reducing its range in tropical ocean basins.Overall, the biological responses of the marine diatom Chaetoceros socialis and the N2-fixing cyanobacteria Trichodesmium IMS101 to several key climate change processes were presented and discussed in this study. These processes included dust deposition, and ocean acidification and warming, which were shown to have a significant impact on oceanic phytoplankton growth, cell size and primary productivity, biological N2 fixation, phytoplankton distribution and community composition. They would thus affect the C, N, P, Si and Fe biogeochemical cycles in various ways. Iron, as one of the most crucial micronutrients for marine phytoplankton, has in particular strong links to climate change and biogeochemical feedback mechanisms.
Posts Tagged 'nitrogen fixation'
Marine iron biogeochemistry under a changing climate: impact on the phytoplankton and the diazotroph communities
Published 13 February 2018 Science ClosedTags: biogeochemistry, biological response, growth, laboratory, multiple factors, nitrogen fixation, phytoplankton, prokaryotes, temperature
Nutrient co-limited Trichodesmium as nitrogen source or sink in a future ocean
Published 4 December 2017 Science ClosedTags: biological response, laboratory, molecular biology, multiple factors, nitrogen fixation, nutrients, physiology, prokaryotes
Nitrogen-fixing (N2) cyanobacteria provide bioavailable nitrogen to vast ocean regions but are in turn limited by iron (Fe) and/or phosphorus (P), which may force them to employ alternative nitrogen acquisition strategies. The adaptive responses of nitrogen-fixers to global-change drivers under nutrient-limited conditions could profoundly alter the current ocean nitrogen and carbon cycles. Here, we show that the globally-important N2-fixer Trichodesmium fundamentally shifts nitrogen metabolism towards organic-nitrogen scavenging following long-term high-CO2 adaptation under iron and/or phosphorus (co)-limitation. Global shifts in transcripts and proteins under high CO2/Fe-limited and/or P-limited conditions include decreases in the N2-fixing nitrogenase enzyme, coupled with major increases in enzymes that oxidize trimethylamine (TMA). TMA is an abundant, biogeochemically-important organic nitrogen compound that supports rapid Trichodesmium growth while inhibiting N2 fixation. In a future high-CO2 ocean, this whole-cell energetic reallocation towards organic nitrogen scavenging and away from N2-fixation may reduce new-nitrogen inputs by Trichodesmium, while simultaneously depleting the scarce fixed-nitrogen supplies of nitrogen-limited open ocean ecosystems.
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Response to Comment on “The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium”
Published 16 October 2017 Science ClosedTags: biological response, growth, nitrogen fixation, physiology, phytoplankton
Hutchins et al. question the validity of our results showing that under fast growth conditions, the beneficial effect of high CO2 on Trichodesmium is overwhelmed by the deleterious effect of the concomitant decrease in ambient and cellular pH. The positive effect of acidification reported by Hutchins and co-workers is likely caused by culture conditions that support suboptimal growth rates.
Comment on “The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium”
Published 13 October 2017 Science ClosedTags: biological response, growth, laboratory, nitrogen fixation, physiology, phytoplankton
Hong et al. (Reports, 5 May 2017, p. 527) suggested that previous studies of the biogeochemically significant marine cyanobacterium Trichodesmium showing increased growth and nitrogen fixation at projected future high CO2 levels suffered from ammonia or copper toxicity. They reported that these rates instead decrease at high CO2 when contamination is alleviated. We present and discuss results of multiple published studies refuting this toxicity hypothesis.
The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium
Published 28 April 2017 Science ClosedTags: biological response, growth, laboratory, nitrogen fixation, physiology, phytoplankton
Acidification of seawater caused by anthropogenic carbon dioxide (CO2) is anticipated to influence the growth of dinitrogen (N2)–fixing phytoplankton, which contribute a large fraction of primary production in the tropical and subtropical ocean. We found that growth and N2-fixation of the ubiquitous cyanobacterium Trichodesmium decreased under acidified conditions, notwithstanding a beneficial effect of high CO2. Acidification resulted in low cytosolic pH and reduced N2-fixation rates despite elevated nitrogenase concentrations. Low cytosolic pH required increased proton pumping across the thylakoid membrane and elevated adenosine triphosphate production. These requirements were not satisfied under field or experimental iron-limiting conditions, which greatly amplified the negative effect of acidification.
Influence of ocean acidification and deep water upwelling on oligotrophic plankton communities in the subtropical North Atlantic: Insights from an in situ mesocosm study
Published 15 March 2017 Science ClosedTags: abundance, biogeochemistry, biological response, BRcommunity, chemistry, community composition, crustaceans, field, fish, mesocosms, methods, mollusks, multiple factors, nitrogen fixation, North Atlantic, nutrients, otherprocess, physiology, phytoplankton, primary production, prokaryotes, protists, virus, zooplankton
Oceanic uptake of anthropogenic carbon dioxide (CO2) causes pronounced shifts in marine carbonate chemistry and a decrease in seawater pH. Increasing evidence indicates that these changes – summarized by the term ocean acidification (OA) – can significantly affect marine food webs and biogeochemical cycles. However, current scientific knowledge is largely based on laboratory experiments with single species and artificial boundary conditions, whereas studies of natural plankton communities are still relatively rare. Moreover, the few existing community-level studies were mostly conducted in rather eutrophic environments, while less attention has been paid to oligotrophic systems such as the subtropical ocean gyres.
Here we report from a recent in situ mesocosm experiment off the coast of Gran Canaria in the eastern subtropical North Atlantic, where we investigated the influence of OA on the ecology and biogeochemistry of plankton communities in oligotrophic waters under close-to-natural conditions. This paper is the first in this Research Topic of Frontiers in Marine Biogeochemistry and provides (1) a detailed overview of the experimental design and important events during our mesocosm campaign, and (2) first insights into the ecological responses of plankton communities to simulated OA over the course of the 62-day experiment.
One particular scientific objective of our mesocosm experiment was to investigate how OA impacts might differ between oligotrophic conditions and phases of high biological productivity, which regularly occur in response to upwelling of nutrient-rich deep water in the study region. Therefore, we specifically developed a deep water collection system that allowed us to obtain ~85 m3 of seawater from ~650 m depth. Thereby, we replaced ~20% of each mesocosm’s volume with deep water, and thus successfully simulated a deep water upwelling event that induced a pronounced plankton bloom.
Our study revealed significant effects of OA on the entire food web, leading to a restructuring of plankton communities that emerged during the oligotrophic phase, and was further amplified during the bloom that developed in response to deep water addition. Such CO2-related shifts in plankton community composition could have consequences for ecosystem productivity, biomass transfer to higher trophic levels, and biogeochemical element cycling of oligotrophic ocean regions.
Special edition of Estuarine, Coastal and Shelf Science – “Ocean acidification in the Mediterranean Sea: pelagic mesocosm experiments”
Published 14 February 2017 Science ClosedTags: biogeochemistry, biological response, BRcommunity, community composition, field, Mediterranean, mesocosms, multiple factors, nitrogen fixation, nutrients, otherprocess, physiology, phytoplankton, primary production, prokaryotes, virus, zooplankton
The topic of ocean acidification has received extensive attention in a recently published special edition of the journal Estuarine, Coastal and Shelf Science. Volume 186, Part A presents a series of 12 research papers focusing on pelagic mesocosm experiments conducted in the Mediterranean Sea in 2012 and 2013. Plankton plays a key role in the global carbon cycle. It is therefore important to project the evolution of plankton community structure and function in a future high-CO2 world. Several results from experiments conducted at the community level have shown increased rates of community primary production and shifts in community composition as a function of increasing pCO2. However, the great majority of these – experiments have been performed under high natural or nutrient-enriched conditions and very few data are available in areas with naturally low levels of nutrient and chlorophyll i.e. oligotrophic areas such as the Mediterranean Sea, although they represent a large and expanding part of the ocean surface. In the frame of the European Mediterranean Sea Acidification in a changing climate project (MedSeA; http://medsea-project.eu), large-scale in situ mesocosms (9 x 50 m3, 12 m deep) have been used to quantify the potential effects of CO2 enrichment in two coastal areas of the Mediterranean Sea: the bay of Calvi (Corsica, France) in June/July 2012 and the bay of Villefranche (France) in February/March 2013. These two experiments gathered the expertise of more than 25 scientists from 7 institutes and 6 countries (France, Greece, Spain, UK, Italy, Belgium, US).
Molecular and physiological evidence of genetic assimilation to high CO2 in the marine nitrogen fixer Trichodesmium
Published 25 November 2016 Science ClosedTags: adaptation, biological response, BRcommunity, communitymodeling, growth, laboratory, modeling, molecular biology, nitrogen fixation, otherprocess, prokaryotes
Most investigations of biogeochemically important microbes have focused on plastic (short-term) phenotypic responses in the absence of genetic change, whereas few have investigated adaptive (long-term) responses. However, no studies to date have investigated the molecular progression underlying the transition from plasticity to adaptation under elevated CO2 for a marine nitrogen-fixer. To address this gap, we cultured the globally important cyanobacterium Trichodesmium at both low and high CO2 for 4.5 y, followed by reciprocal transplantation experiments to test for adaptation. Intriguingly, fitness actually increased in all high-CO2 adapted cell lines in the ancestral environment upon reciprocal transplantation. By leveraging coordinated phenotypic and transcriptomic profiles, we identified expression changes and pathway enrichments that rapidly responded to elevated CO2 and were maintained upon adaptation, providing strong evidence for genetic assimilation. These candidate genes and pathways included those involved in photosystems, transcriptional regulation, cell signaling, carbon/nitrogen storage, and energy metabolism. Conversely, significant changes in specific sigma factor expression were only observed upon adaptation. These data reveal genetic assimilation as a potentially adaptive response of Trichodesmium and importantly elucidate underlying metabolic pathways paralleling the fixation of the plastic phenotype upon adaptation, thereby contributing to the few available data demonstrating genetic assimilation in microbial photoautotrophs. These molecular insights are thus critical for identifying pathways under selection as drivers in plasticity and adaptation.
Mechanisms of increased Trichodesmium fitness under iron and phosphorus co-limitation in the present and future ocean
Published 13 July 2016 Science ClosedTags: biological response, growth, laboratory, molecular biology, morphology, multiple factors, nitrogen fixation, nutrients, physiology, prokaryotes
Nitrogen fixation by cyanobacteria supplies critical bioavailable nitrogen to marine ecosystems worldwide; however, field and lab data have demonstrated it to be limited by iron, phosphorus and/or CO2. To address unknown future interactions among these factors, we grew the nitrogen-fixing cyanobacterium Trichodesmium for 1 year under Fe/P co-limitation following 7 years of both low and high CO2 selection. Fe/P co-limited cell lines demonstrated a complex cellular response including increased growth rates, broad proteome restructuring and cell size reductions relative to steady-state growth limited by either Fe or P alone. Fe/P co-limitation increased abundance of a protein containing a conserved domain previously implicated in cell size regulation, suggesting a similar role in Trichodesmium. Increased CO2 further induced nutrient-limited proteome shifts in widespread core metabolisms. Our results thus suggest that N2-fixing microbes may be significantly impacted by interactions between elevated CO2 and nutrient limitation, with broad implications for global biogeochemical cycles in the future ocean.
No observed effect of ocean acidification on nitrogen biogeochemistry in a summer Baltic Sea plankton community (update)
Published 8 July 2016 Science ClosedTags: abundance, Baltic, biogeochemistry, biological response, BRcommunity, field, mesocosms, nitrogen fixation, otherprocess, prokaryotes
Nitrogen fixation by filamentous cyanobacteria supplies significant amounts of new nitrogen (N) to the Baltic Sea. This balances N loss processes such as denitrification and anammox, and forms an important N source supporting primary and secondary production in N-limited post-spring bloom plankton communities. Laboratory studies suggest that filamentous diazotrophic cyanobacteria growth and N2-fixation rates are sensitive to ocean acidification, with potential implications for new N supply to the Baltic Sea. In this study, our aim was to assess the effect of ocean acidification on diazotroph growth and activity as well as the contribution of diazotrophically fixed N to N supply in a natural plankton assemblage. We enclosed a natural plankton community in a summer season in the Baltic Sea near the entrance to the Gulf of Finland in six large-scale mesocosms (volume ∼ 55 m3) and manipulated fCO2 over a range relevant for projected ocean acidification by the end of this century (average treatment fCO2: 365–1231 µatm). The direct response of diazotroph growth and activity was followed in the mesocosms over a 47 day study period during N-limited growth in the summer plankton community. Diazotrophic filamentous cyanobacteria abundance throughout the study period and N2-fixation rates (determined only until day 21 due to subsequent use of contaminated commercial 15N-N2 gas stocks) remained low. Thus estimated new N inputs from diazotrophy were too low to relieve N limitation and stimulate a summer phytoplankton bloom. Instead, regeneration of organic N sources likely sustained growth in the plankton community. We could not detect significant CO2-related differences in neither inorganic nor organic N pool sizes, or particulate matter N : P stoichiometry. Additionally, no significant effect of elevated CO2 on diazotroph activity was observed. Therefore, ocean acidification had no observable impact on N cycling or biogeochemistry in this N-limited, post-spring bloom plankton assemblage in the Baltic Sea.
Implications of ocean acidification for marine microorganisms from the free-living to the host-associated
Published 11 April 2016 Science ClosedTags: abundance, biological response, BRcommunity, community composition, field, laboratory, nitrogen fixation, otherprocess, primary production, prokaryotes, review
Anthropogenic CO2 emissions are causing oceans to become more acidic, with consequences for all marine life including microorganisms. Studies reveal that from the microbes that occupy the open ocean to those intimately associated with their invertebrate hosts, changing ocean chemistry will alter the critical functions of these important organisms. Our current understanding indicates that bacterial communities associated with their host will shift as pH drops by another 0.2-0.4 units over the next 100 years. It is unclear what impacts this will have for host health, however increased vulnerability to disease seems likely for those associated with reef corals. Natural CO2 seeps have provided a unique setting for the study of microbial communities under OA in situ, where shifts in the bacterial communities associated with corals at the seep are correlated with a decline in abundance of the associated coral species. Changes to global biogeochemical cycles also appear likely as photosynthesis and nitrogen fixation by pelagic microbes becomes enhanced under low pH conditions. However, recent long-term studies have shown that pelagic microbes are also capable of evolutionary adaptation, with some physiological responses to a decline in pH restored after hundreds of generations at high pCO2 levels. The impacts of ocean acidification (OA) also will not work in isolation, thus synergistic interactions with other potential stressors, such as rising seawater temperatures, will likely exacerbate the microbial response to OA. This review discusses our existing understanding of the impacts of OA on both pelagic and host-associated marine microbial communities, whilst highlighting the importance of controlled laboratory studies and in situ experiments, to fill the current gaps in our knowledge.
Nitrate fertilisation does not enhance CO2 responses in two tropical seagrass species
Published 23 March 2016 Science ClosedTags: biological response, growth, laboratory, multiple factors, nitrogen fixation, nutrients, phanerogams, photosynthesis, physiology, primary production, respiration, South Pacific
Seagrasses are often considered “winners” of ocean acidification (OA); however, seagrass productivity responses to OA could be limited by nitrogen availability, since nitrogen-derived metabolites are required for carbon assimilation. We tested nitrogen uptake and assimilation, photosynthesis, growth, and carbon allocation responses of the tropical seagrasses Halodule uninervis and Thalassia hemprichii to OA scenarios (428, 734 and 1213 μatm pCO2) under two nutrients levels (0.3 and 1.9 μM NO3−). Net primary production (measured as oxygen production) and growth in H. uninervis increased with pCO2 enrichment, but were not affected by nitrate enrichment. However, nitrate enrichment reduced whole plant respiration in H. uninervis. Net primary production and growth did not show significant changes with pCO2 or nitrate by the end of the experiment (24 d) in T. hemprichii. However, nitrate incorporation in T. hemprichii was higher with nitrate enrichment. There was no evidence that nitrogen demand increased with pCO2 enrichment in either species. Contrary to our initial hypothesis, nutrient increases to levels approximating present day flood plumes only had small effects on metabolism. This study highlights that the paradigm of increased productivity of seagrasses under ocean acidification may not be valid for all species under all environmental conditions.
Ocean acidification impacts on nitrogen fixation in the coastal western Mediterranean Sea
Published 14 January 2016 Science ClosedTags: biogeochemistry, biological response, BRcommunity, chemistry, community composition, field, Mediterranean, mesocosms, nitrogen fixation, otherprocess, prokaryotes
The effects of ocean acidification on nitrogen (N2) fixation rates and on the community composition of N2-fixing microbes (diazotrophs) were examined in coastal waters of the North-Western Mediterranean Sea. Nine experimental mesocosm enclosures of ∼50 m3 each were deployed for 20 days during June-July 2012 in the Bay of Calvi, Corsica, France. Three control mesocosms were maintained under ambient conditions of carbonate chemistry. The remainder were manipulated with CO2 saturated seawater to attain target amendments of pCO2 of 550, 650, 750, 850, 1000 and 1250 μatm. Rates of N2 fixation were elevated up to 10 times relative to control rates (2.00 ± 1.21 nmol L-1d-1) when pCO2 concentrations were >1000 μatm and pHT (total scale) < 7.74. Diazotrophic phylotypes commonly found in oligotrophic marine waters, including the Mediterranean, were not present at the onset of the experiment and therefore, the diazotroph community composition was characterised by amplifying partial nifH genes from the mesocosms. The diazotroph community was comprised primarily of cluster III nifH sequences (which include possible anaerobes), and proteobacterial (α and γ) sequences, in addition to small numbers of filamentous (or pseudo-filamentous) cyanobacterial phylotypes. The implication from this study is that there is some potential for elevated N2 fixation rates in the coastal western Mediterranean before the end of this century as a result of increasing ocean acidification. Observations made of variability in the diazotroph community composition could not be correlated with changes in carbon chemistry, which highlights the complexity of the relationship between ocean acidification and these keystone organisms.
Impact of climate change and ocean acidification on the marine nitrogen cycle
Published 13 November 2015 Science ClosedTags: biogeochemistry, biological response, globalmodeling, modeling, multiple factors, nitrogen fixation, otherprocess, primary production, temperature
The marine nitrogen cycle is responsible for two climate feedbacks in the Earth System. Firstly, it modulates the fixed nitrogen pool available for phytoplankton growth and hence it modulates in part the strength of the biological pump, one of the mechanisms contributing to the oceanic uptake of anthropogenic CO2. Secondly, the nitrogen cycle produces a powerful greenhouse gas and ozone (O3) depletion agent called nitrous oxide (N2O). Future changes of the nitrogen cycle in response to global warming, ocean deoxygenation and ocean acidification are largely unknown. Processes such as N2-fixation, nitrification, denitrification and N2O production will experience changes under the simultaneous effect of these three stressors. Global ocean biogeochemical models allow us to study such interactions. Using NEMO-PISCES and the CMIP5 model ensemble we project changes in year 2100 under the business-as-usual high CO2 emissions scenario in global scale N2-fixation rates, nitrification rates, N2O production and N2O sea-to-air fluxes adding CO2 sensitive functions into the model parameterizations. Second order effects due to the combination of global warming in tandem with ocean acidification on the fixed nitrogen pool, primary productivity and N2O radiative forcing feedbacks are also evaluated in this thesis.
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No observed effect of ocean acidification on nitrogen biogeochemistry in a summer Baltic Sea plankton community
Published 30 October 2015 Science ClosedTags: abundance, Baltic, biogeochemistry, biological response, BRcommunity, field, mesocosms, nitrogen fixation, otherprocess, prokaryotes
Nitrogen fixation by filamentous cyanobacteria supplies significant amounts of new nitrogen (N) to the Baltic Sea. This balances N loss processes such as denitrification and anammox and forms an important N source supporting primary and secondary production in N-limited post-spring bloom plankton communities. Laboratory studies suggest that filamentous diazotrophic cyanobacteria growth and N2-fixation rates are sensitive to ocean acidification with potential implications for new N supply to the Baltic Sea. In this study, our aim was to assess the effect of ocean acidification on diazotroph growth and activity as well as the contribution of diazotrophically-fixed N to N supply in a natural plankton assemblage. We enclosed a natural plankton community in a summer season in the Baltic Sea near the entrance to the Gulf of Finland in six large-scale mesocosms (volume ~ 55 m3) and manipulated fCO2 over a range relevant for projected ocean acidification by the end of this century (average treatment fCO2: 365–1231 μatm). The direct response of diazotroph growth and activity was followed in the mesocosms over a 47 day study period during N-limited growth in the summer plankton community. Diazotrophic filamentous cyanobacteria abundance throughout the study period and N2-fixation rates (determined only until day 21 due to subsequent use of contaminated commercial 15N-N2 gas stocks) remained low. Thus estimated new N inputs from diazotrophy were too low to relieve N limitation and stimulate a summer phytoplankton bloom. Instead regeneration of organic N sources likely sustained growth in the plankton community. We could not detect significant CO2-related differences in inorganic or organic N pools sizes, or particulate matter N : P stoichiometry. Additionally, no significant effect of elevated CO2 on diazotroph activity was observed. Therefore, ocean acidification had no observable impact on N cycling or biogeochemistry in this N-limited, post-spring bloom plankton assemblage in the Baltic Sea.
Responses of elevated CO2 on photosynthesis and nitrogen metabolism in Ulva lactuca (Chlorophyta) at different temperature levels
Published 22 September 2015 Science ClosedTags: algae, biological response, growth, laboratory, multiple factors, nitrogen fixation, photosynthesis, physiology, respiration, temperature
Ulva lactuca was cultured in a laboratory at two CO2 levels (390 and 700 μl l−1) and at low (15°C) and high (25°C) temperature levels. Growth, biochemical composition, photosynthesis and nitrogen metabolism were examined to evaluate the impacts of elevated CO2 and temperature on this species. Elevated CO2 had little effect on the relative growth rate (RGR) in both low and high temperature levels. The levels of Chl a, carotenoids, soluble protein (SP) and soluble carbohydrates (SC) were also unaffected by CO2 enrichment. Elevated CO2 decreased the content of total fatty acids (FAs) but the higher temperature increased them. At 15°C, the light-saturated maximal photosynthetic rate (Pm), photosynthetic efficiency (α), and dark respiration (Rd) were barely affected by the increase in CO2 concentration. However, at 25°C the Pm, α and Rd were pronouncedly enhanced by elevated CO2. Additionally, high temperature increased the slopes of curves of electron transport rates (ETR) versus gross photosynthesis (Pg), indicating the loss of correlation between ETR and linear photosynthetic flow. The elevation of temperature significantly increased the nitrogen uptake rates (NUR) during the dark and light period, while elevated CO2 increased NUR only during the light period. Increased CO2 had no marked effect on nitrate reductase (NR) activity at 15°C, but high CO2 could remarkably improve the NR activity at 25°C. These results suggest that the responses to elevated CO2 in U. lactuca were temperature-dependent, and elevated CO2 alone was less effective than temperature levels.
Irreversibly increased nitrogen fixation in Trichodesmium experimentally adapted to elevated carbon dioxide
Published 3 September 2015 Science ClosedTags: adaptation, biological response, growth, laboratory, molecular biology, nitrogen fixation, otherprocess, prokaryotes
Nitrogen fixation rates of the globally distributed, biogeochemically important marine cyanobacterium Trichodesmium increase under high carbon dioxide (CO2) levels in short-term studies due to physiological plasticity. However, its long-term adaptive responses to ongoing anthropogenic CO2 increases are unknown. Here we show that experimental evolution under extended selection at projected future elevated CO2 levels results in irreversible, large increases in nitrogen fixation and growth rates, even after being moved back to lower present day CO2 levels for hundreds of generations. This represents an unprecedented microbial evolutionary response, as reproductive fitness increases acquired in the selection environment are maintained after returning to the ancestral environment. Constitutive rate increases are accompanied by irreversible shifts in diel nitrogen fixation patterns, and increased activity of a potentially regulatory DNA methyltransferase enzyme. High CO2-selected cell lines also exhibit increased phosphorus-limited growth rates, suggesting a potential advantage for this keystone organism in a more nutrient-limited, acidified future ocean.
Ocean acidification and marine microorganisms: responses and consequences
Published 5 August 2015 Science ClosedTags: abundance, biological response, BRcommunity, community composition, methods, nitrogen fixation, otherprocess, physiology, phytoplankton, primary production, prokaryotes, review
Ocean acidification (OA) is one of the global issues caused by rising atmospheric CO2. The rising pCO2 and resulting pH decrease has altered ocean carbonate chemistry. Microbes are key components of marine environments involved in nutrient cycles and carbon flow in marine ecosystems. However, these marine microbes and the microbial processes are sensitive to ocean pH shift. Thus, OA affects the microbial diversity, primary productivity and trace gases emission in oceans. Apart from that, it can also manipulate the microbial activities such as quorum sensing, extracellular enzyme activity and nitrogen cycling. Short-term laboratory experiments, mesocosm studies and changing marine diversity scenarios have illustrated undesirable effects of OA on marine microorganisms and ecosystems. However, from the microbial perspective, the current understanding on effect of OA is based mainly on limited experimental studies. It is challenging to predict response of marine microbes based on such experiments for this complex process. To study the response of marine microbes towards OA, multiple approaches should be implemented by using functional genomics, new generation microscopy, small-scale interaction among organisms and/or between organic matter and organisms. This review focuses on the response of marine microorganisms to OA and the experimental approaches to investigate the effect of changing ocean carbonate chemistry on microbial mediated processes.
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Inter- and intra-specific responses of coccolithophores to CO2-induced ocean acidification
Published 14 January 2015 Science ClosedTags: biogeochemistry, biological response, calcification, growth, laboratory, nitrogen fixation, North Pacific, photosynthesis, physiology, phytoplankton, primary production
Oceanic uptake of anthropogenic carbon dioxide (CO2) is altering the seawater chemistry of the world’s oceans with consequences for marine bioregions, especially calcareous organisms such as corals, foraminifera and coccolithophores. The coccolithophores, one of the most abundant and widespread groups of calcifying plankton, are responsible for a large proportion of modern oceanic carbonate production. However, culture experiments examining the response of coccolithophores to elevated CO2 partial pressure (pCO2) have mostly been based on investigations of a single strain and have yielded contradictory results from different experiments between and even within species. Here, four strains of the coccolithophores Emiliania huxleyi (E. huxleyi) and Gephyrocapsa oceanica (G. oceanica), which contained separately naked and calcifying strains, were investigated simultaneously for the first time in a bubbling batch culture at four CO2 grades ranging from approximately 380 to 2000 μatm. We synchronously determined multiple physiological parameters of four coccolithophore strains involving growth, photosynthesis, nitrogen uptake, elemental compositions and calcification efficiency in the process of cultivation. The results did not show a uniform response from different strains to elevated pCO2 up to 2000 μatm, and the naked strain E. huxleyi (N-E) was seriously suppressed, in sharp contrast to the positive response of the different levels of the other three algae. In addition, we fitted nitrogen uptake rate response curves relative to changing pCO2 for the four strains and applied kinetic constants from the response curves to further analyze the hypostatic difference among different strains, which reflected the same variational trend of the four stains above vs. increasing CO2. We determined that the responses of coccolithophores to ocean acidification are inter- and intra-specific, and this variation may cause changes to biodiversity and other ecosystem processes in the future ocean.
Ocean acidification rapidly reduces dinitrogen fixation associated with the hermatypic coral Seriatopora hystrix
Published 24 November 2014 Science ClosedTags: algae, biological response, laboratory, nitrogen fixation
Since productivity and growth of coral-associated dinoflagellate algae is nitrogen (N)-limited, dinitrogen (N2) fixation by coral-associated microbes is likely crucial for maintaining the coral-dinoflagellate symbiosis. It is thus essential to understand the effects future climate change will have on N2 fixation by the coral holobiont. This laboratory study is the first to investigate short-term effects of ocean acidification on N2 fixation activity associated with the tropical, hermatypic coral Seriatopora hystrix using the acetylene reduction assay in combination with calcification measurements. Findings reveal that simulated ocean acidification ( pCO2 1080 µatm) caused a rapid and significant decrease (53%) in N2 fixation rates associated with S. hystrix compared to the present day scenario ( pCO2 486 µatm). In addition, N2 fixation associated with the coral holobiont showed a positive exponential relationship with its calcification rates. This suggests that even small declines in calcification rates of hermatypic corals under high CO2 conditions may result in decreased N2 fixation activity, since these 2 processes may compete for energy in the coral holobiont. Ultimately, an intensified N limitation in combination with a decline in skeletal growth may trigger a negative feedback loop on coral productivity exacerbating the negative long-term effects of ocean acidification.
