Posts Tagged 'nitrogen fixation'

Impact of increasing carbon dioxide on dinitrogen and carbon fixation rates under oligotrophic conditions and simulated upwelling

Dinitrogen (N2) fixation is a major source of bioavailable nitrogen to oligotrophic ocean communities. Yet, we have limited understanding how ongoing climate change could alter N2 fixation. Most of our understanding is based on short-term laboratory experiments conducted on individual N2-fixing species whereas community-level approaches are rare. In this longer-term in situ mesocosm study, we aimed to improve our understanding on the role of rising atmospheric carbon dioxide (CO2) and simulated deep water upwelling on N2 and carbon (C) fixation rates in a natural oligotrophic plankton community. We deployed nine mesocosms in the subtropical North Atlantic Ocean and enriched seven of these with CO2 to yield a range of treatments (partial pressure of CO2pCO2 = 352–1025 μatm). We measured rates of N2 and C fixation in both light and dark incubations over the 55-day study period. High pCO2 negatively impacted light and dark N2 fixation rates in the oligotrophic phase before simulated upwelling, while the effect reversed in the light N2 fixation rates in the bloom decay phase after added nutrients were consumed. Dust deposition and simulated upwelling of nutrient-rich deep water increased N2 fixation rates and nifH gene abundances of selected clades including the unicellular diazotrophic cyanobacterium clade UCYN-B. Elevated pCO2 increased C fixation rates in the decay phase. We conclude that elevated pCO2 and pulses of upwelling have pronounced effects on diazotrophy and primary producers, and upwelling and dust deposition modify the pCO2 effect in natural assemblages.

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Everything is everywhere: physiological responses of the Mediterranean sea and Eastern Pacific ocean Epiphyte Cobetia Sp. to varying nutrient concentration

Bacteria are essential in the maintenance and sustainment of marine environments (e.g., benthic systems), playing a key role in marine food webs and nutrient cycling. These microorganisms can live associated as epiphytic or endophytic populations with superior organisms with valuable ecological functions, e.g., seagrasses. Here, we isolated, identified, sequenced, and exposed two strains of the same species (i.e., identified as Cobetia sp.) from two different marine environments to different nutrient regimes using batch cultures: (1) Cobetia sp. UIB 001 from the endemic Mediterranean seagrass Posidonia oceanica and (2) Cobetia sp. 4B UA from the endemic Humboldt Current System (HCS) seagrass Heterozostera chilensis. From our physiological studies, both strains behaved as bacteria capable to cope with different nutrient and pH regimes, i.e., N, P, and Fe combined with different pH levels, both in long-term (12 days (d)) and short-term studies (4 d/96 h (h)). We showed that the isolated strains were sensitive to the N source (inorganic and organic) at low and high concentrations and low pH levels. Low availability of phosphorus (P) and Fe had a negative independent effect on growth, especially in the long-term studies. The strain UIB 001 showed a better adaptation to low nutrient concentrations, being a potential N2-fixer, reaching higher growth rates (μ) than the HCS strain. P-acquisition mechanisms were deeply investigated at the enzymatic (i.e., alkaline phosphatase activity, APA) and structural level (e.g., alkaline phosphatase D, PhoD). Finally, these results were complemented with the study of biochemical markers, i.e., reactive oxygen species (ROS). In short, we present how ecological niches (i.e., MS and HCS) might determine, select, and modify the genomic and phenotypic features of the same bacterial species (i.e., Cobetia spp.) found in different marine environments, pointing to a direct correlation between adaptability and oligotrophy of seawater.

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Elevated pCO2 reinforces preference among intertidal algae in both a specialist and generalist herbivore

Highlights

  • Elevated pCO2 influences growth and chemical composition of some intertidal algae.
  • Herbivore preference is reinforced by resilience of preferred alga to pCO2 exposure.
  • Preference is also influenced by changes in lesser-preferred algal species.
  • Specialist and generalist feeding may be indirectly affected by ocean acidification.

Abstract

Ocean acidification (OA) can induce changes in marine organisms and species interactions. We examined OA effects on intertidal macroalgal growth, palatability, and consumption by a specialist crab (Pugettia producta) and a generalist snail (Tegula funebralis) herbivore. Moderate increases in pCO2 increased algal growth in most species, but effects of pCO2 on C:N and phenolic content varied by species. Elevated pCO2 had no effect on algal acceptability to herbivores, but did affect their preference ranks. Under elevated pCO2, electivity for a preferred kelp (Egregia menziesii) and preference rankings among algal species strengthened for both P. producta and T. funebralis, attributable to resilience of E. menziesii in elevated pCO2 and to changes in palatability among less-preferred species. Preferred algae may therefore grow more under moderate pCO2 increases in the future, but their appeal to herbivores may be strengthened by associated shifts in nutritional quality and defensive compounds in other species.

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Increased light availability modulates carbon and nitrogen accumulation in the macroalga Gracilariopsis lemaneiformis (Rhodophyta) in response to ocean acidification

Highlights

  • The effects of light and elevated pCO2 on Gracilariopsis were examined.
  • Ocean acidification enhanced algal biomass, photosynthesis and total C/N ratios.
  • Increasing light and elevated pCO2 lowered nutritional quality of G. lemaneiformis.

Abstract

The economically important red macroalga Gracilariopsis lemaneiformis has demonstrated positive ecological functions in nutrient bioextraction efficiency and high harvestable biomass, as well as being a food and agar source owing to its richness in proteins and polysaccharides. Carbon dioxide (CO2)-induced ocean acidification has resulted in mixed nutrient compound accumulations in this marine autotroph. G. lemaneiformis also experiences light variations resulting from self-shading and varied cultivation depths. Therefore, a factorial coupling experiment was conducted to examine how growth, photosynthesis performance, soluble cell components and metabolic enzyme-driven activities respond to light availability changes and CO2 enrichment. The ocean acidification enhanced the growth characteristics, total carbon/nitrogen ratios and metabolic nutrient accumulation processes in G. lemaneiformis regardless of the light level. Photosynthetic performances, including relative electron transport rate and maximum photochemical quantum yield, were increased by high pCO2 concentrations, resulting in soluble carbohydrate accumulation. The carbon and nitrogen accumulations might result from variations in carbonic anhydrase and nitrate reductase activities under high pCO2 conditions. The soluble protein and free amino acids contents declined in response to CO2 elevation, and this effect was more pronounced as the light intensity increased. Thus, future climate changes may cause greater algal biomass accumulations, but they may negatively affect the cell composition and nutritional quality of G. lemaneiformis.

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Effects of ocean acidification on carbon and nitrogen fixation in the hermatypic coral Galaxea fascicularis

The supply of metabolites from symbionts to scleractinian corals is crucial to coral health. Members of the Symbiodiniaceae can enhance coral calcification by providing photosynthetically fixed carbon (PFC) and energy, whereas dinitrogen (N2)-fixing bacteria can provide additional nutrients such as diazotrophically-derived nitrogen (DDN) that sustain coral productivity especially when alternative external nitrogen sources are scarce. How these mutualistic associations benefit corals in the future acidifying ocean is not well understood. In this study, we investigated the possible effects of ocean acidification (OA; pHs 7.7 and 7.4 vs. 8.1) on calcification in the hermatypic coral Galaxea fascicularis with respect to PFC and DDN assimilation. Our measurements based on isotopic tracing showed no significant differences in the assimilation of PFC among different pH treatments, but the assimilation of DDN decreased significantly after 28 days of stress at pH 7.4. The decreased DDN assimilation suggests a nitrogenous nutrient deficiency in the coral holotiont, potentially leading to reduced coral calcification and resilience to bleaching and other stressful events. This contrasting impact of OA on carbon and N flux demonstrates the flexibility of G. fascicularis in coping with OA, apparently by sustaining a largely undamaged photosystem at the expense of N2 fixation machinery, which competes with coral calcification for energy from photosynthesis. These findings shed new light on the critically important but more vulnerable N cycling in hospite, and on the trade-off between coral hosts and symbionts in response to future climate change.

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Coastal ocean acidification and nitrogen loading facilitate invasions of the non-indigenous red macroalga, Dasysiphonia japonica

Coastal ecosystems are prone to multiple anthropogenic and natural stressors including eutrophication, acidification, and invasive species. While the growth of some macroalgae can be promoted by excessive nutrient loading and/or elevated pCO2, responses differ among species and ecosystems. Native to the western Pacific Ocean, the filamentous, turf-forming rhodophyte, Dasysiphonia japonica, appeared in estuaries of the northeastern Atlantic Ocean during the 1980s and the northwestern Atlantic Ocean during the late 2000s. Here, we report on the southernmost expansion of the D. japonica in North America and the effects of elevated nutrients and elevated pCO2 on the growth of D. japonica over an annual cycle in Long Island, New York, USA. Growth limitation of the macroalga varied seasonally. During winter and spring, when water temperatures were < 15 °C, growth was significantly enhanced by elevated pCO2 (p < 0.05). During summer and fall, when the water temperature was 15–24 °C, growth was significantly higher under elevated nutrient treatments (p < 0.05). When temperatures reached 28 °C, the macroalga grew poorly and was unaffected by nutrients or pCO2. The δ13C content of regional populations of D. japonica was −30‰, indicating the macroalga is an obligate CO2-user. This result, coupled with significantly increased growth under elevated pCO2 when temperatures were < 15 °C, indicates this macroalga is carbon-limited during colder months, when in situ pCO2 was significantly lower in Long Island estuaries compared to warmer months when estuaries are enriched in metabolically derived CO2. The δ15N content of this macroalga (9‰) indicated it utilized wastewater-derived N and its N limitation during warmer months coincided with lower concentrations of dissolved inorganic N in the water column. Given the stimulatory effect of nutrients on this macroalga and that eutrophication can promote seasonally elevated pCO2, this study suggests that eutrophic estuaries subject to peak annual temperatures < 28 °C may be particularly vulnerable to future invasions of D. japonica as ocean acidification intensifies. Conversely, nutrient reductions would serve as a management approach that would make coastal regions more resilient to invasions by this macroalga.

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Ocean acidification decreases grazing pressure but alters morphological structure in a dominant coastal seaweed

Ocean acidification driven by anthropogenic climate change is causing a global decrease in pH, which is projected to be 0.4 units lower in coastal shallow waters by the year 2100. Previous studies have shown that seaweeds grown under such conditions may alter their growth and photosynthetic capacity. It is not clear how such alterations might impact interactions between seaweed and herbivores, e.g. through changes in feeding rates, nutritional value, or defense levels. Changes in seaweeds are particularly important for coastal food webs, as they are key primary producers and often habitat-forming species. We cultured the habitat-forming brown seaweed Fucus vesiculosus for 30 days in projected future pCO2 (1100 μatm) with genetically identical controls in ambient pCO2 (400 μatm). Thereafter the macroalgae were exposed to grazing by Littorina littorea, acclimated to the relevant pCO2-treatment. We found increased growth (measured as surface area increase), decreased tissue strength in a tensile strength test, and decreased chemical defense (phlorotannins) levels in seaweeds exposed to high pCO2-levels. The herbivores exposed to elevated pCO2-levels showed improved condition index, decreased consumption, but no significant change in feeding preference. Fucoid seaweeds such as Fvesiculosus play important ecological roles in coastal habitats and are often foundation species, with a key role for ecosystem structure and function. The change in surface area and associated decrease in breaking force, as demonstrated by our results, indicate that Fvesiculosus grown under elevated levels of pCO2 may acquire an altered morphology and reduced tissue strength. This, together with increased wave energy in coastal ecosystems due to climate change, could have detrimental effects by reducing both habitat and food availability for herbivores.

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Regulation mechanism of ocean acidification on key physiological processes of microalgae and the effect of environmental factors: A review

The inputs of carbon dioxide from anthropogenic activities to ocean through the sea-air interface exchange disturbs the balance of seawater carbonate system, resulting in ocean acidification (OA). OA affects the physical and chemical properties of both seawater and marine pollutants, which significantly regulates the physiological processes of planktonic algae living on the surface of ocean. As the main primary producers, the physiological function and processes of marine algae play an important role in marine ecosystem. We reviewed the underlying mechanisms of OA on the three key physiological processes of photosynthetic carbon fixation, calcification and nitrogen fixation of marine microalgae. OA could alter environmental factors (e.g., solar radiation, temperature, nutrient elements) and typical marine contaminants (e.g., organic contaminants, heavy metals, microplastics). We further summarized the effects of these factors on the regulation of physiological processes of microalgae. Finally, current research status and prospects for future research were addressed. This review provided important information for better understanding the potential impacts of OA on marine ecosystems.

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Long-term m5C methylome dynamics parallel phenotypic adaptation in the cyanobacterium Trichodesmium

A major challenge in modern biology is understanding how the effects of short-term biological responses influence long-term evolutionary adaptation, defined as a genetically determined increase in fitness to novel environments. This is particularly important in globally important microbes experiencing rapid global change, due to their influence on food webs, biogeochemical cycles, and climate. Epigenetic modifications like methylation have been demonstrated to influence short-term plastic responses, which ultimately impact long-term adaptive responses to environmental change. However, there remains a paucity of empirical research examining long-term methylation dynamics during environmental adaptation in non-model, ecologically important microbes. Here, we show the first empirical evidence in a marine prokaryote for long-term m5C methylome modifications correlated with phenotypic adaptation to CO2, using a 7-year evolution experiment (1000+ generations) with the biogeochemically-important marine cyanobacterium Trichodesmium. We identify m5C methylated sites that rapidly changed in response to high (750 µatm) CO2 exposure and were maintained for at least 4.5 years of CO2 selection. After 7 years of CO2 selection, however, m5C methylation levels that initially responded to high-CO2 returned to ancestral, ambient CO2 levels. Concurrently, high-CO2 adapted growth and N2 fixation rates remained significantly higher than those of ambient CO2 adapted cell lines irrespective of CO2 concentration, a trend consistent with genetic assimilation theory. These data demonstrate the maintenance of CO2-responsive m5C methylation for 4.5 years alongside phenotypic adaptation before returning to ancestral methylation levels. These observations in a globally distributed marine prokaryote provide critical evolutionary insights into biogeochemically important traits under global change.

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Projected expansion of Trichodesmium’s geographical distribution and increase in growth potential in response to climate change

Estimates of marine N2 fixation range from 52 to 73 Tg N/year, of which we calculate up to 84% is from Trichodesmium based on previous measurements of nifH gene abundance and our new model of Trichodesmium growth. Here, we assess the likely effects of four major climate change‐related abiotic factors on the spatiotemporal distribution and growth potential of Trichodesmium for the last glacial maximum (LGM), the present (2006–2015) and the end of this century (2100) by mapping our model of Trichodesmium growth onto inferred global surface ocean fields of pCO2, temperature, light and Fe. We conclude that growth rate was severely limited by low pCO2 at the LGM, that current pCO2 levels do not significantly limit Trichodesmium growth and thus, the potential for enhanced growth from future increases in CO2 is small. We also found that the area of the ocean where sea surface temperatures (SST) are within Trichodesmium‘s thermal niche increased by 32% from the LGM to present, but further increases in SST due to continued global warming will reduce this area by 9%. However, the range reduction at the equator is likely to be offset by enhanced growth associated with expansion of regions with optimal or near optimal Fe and light availability. Between now and 2100, the ocean area of optimal SST and irradiance is projected to increase by 7%, and the ocean area of optimal SST, irradiance and iron is projected to increase by 173%. Given the major contribution of this keystone species to annual N2 fixation and thus pelagic ecology, biogeochemistry and CO2 sequestration, the projected increase in the geographical range for optimal growth could provide a negative feedback to increasing atmospheric CO2 concentrations.

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A rise in ROS and EPS production: new insights into the Trichodesmium erythraeum response to ocean acidification

The diazotrophic cyanobacterium Trichodesmium is thought to be a major contributor to the new N in the parts of the oligotrophic, subtropical and tropical oceans. In this study physiological and biochemical methods and transcriptome sequencing were used to investigate the influences of ocean acidification (OA) on Trichodesmium erythraeum (T. erythraeum). We presented evidence that OA caused by CO2 slowed the growth rate and physiological activity of T. erythraeum. OA led to reduced development of proportion of the vegetative cells into diazocytes which included up‐regulated genes of nitrogen fixation. Reactive oxygen species (ROS) accumulation was increased due to the disruption of photosynthetic electron transport and decrease in antioxidant enzyme activities under acidified conditions. This study showed that OA increased the amounts of (exopolysaccharides) EPS in T. erythraeum, and the key genes of ribose‐5‐phosphate (R5P) and glycosyltransferases (Tery_3818) were up‐regulated. These results provide new insight into how ROS and EPS of T. erythraeum increase in an acidified future ocean to cope with OA‐imposed stress.

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Environmental and nutrient controls of marine nitrogen fixation

Highlights

• Fe, P and dust additions could stimulate N2 fixation of Trichodesmium.
• Novel nutrient acquisition strategies have been discovered for Trichodesmium.
• Fe could be the ultimate limiting factor for N2 fixation.
• Ocean acidification may be beneficial for N2 fixation of Trichodesmium.
• Ocean warming may not play an important role in N2 fixation of Trichodesmium.

Abstract

Biological dinitrogen (N2) fixation by diazotrophic cyanobacteria has great biogeochemical implications in nitrogen (N) cycling in the ocean as this process represents the major source of new N input to the oceans, thereby controlling the marine primary productivity. Numerous factors can affect the extent of N2 fixation. A better understanding of the major environmental and nutrient factors governing this process is highly required. Iron (Fe) and/or phosphorus (P) are thought to be limiting factors in most oceanic regions. Special attention has been given in the present study to evaluate the effects of mineral dust deposition which is believed to stimulate N2 fixation as it increases the availability of both Fe and P. Through three laboratory bioassays (+Fe, +P, +Dust) via incubation experiments performed on Trichodesmium IMS101, we found that each addition of Fe, P or desert dust could stimulate the growth and N2 fixation 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 using natural phytoplankton assemblages from the temperate Northeast Atlantic Ocean the critical role of dissolved Fe (DFe) was again highlighted by the remarkably enhanced N2 fixation rate observed after the addition of DFe under low temperature and P-depleted conditions. At the time of our study no Trichodesmium filaments were found in the studied region, the diazotrophic community was dominated by unicellular cyanobacteria symbiont (prymnesiophyte-UCYN-A1) and heterotrophic diazotrophs, therefore demonstrating that DFe could as well be the ultimate factor limiting N2 fixation of these smaller diazotrophs. 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 present-day and future high pCO2 (400 and 800 µatm, respectively) and warming seawater (24 and 28 °C) conditions. The results indicate that higher pCO2 and therefore ocean acidification may be beneficial for Trichodesmium growth and N2 fixation. However, our observations suggest that Fe or P limitation in oligotrophic seawaters may offset the stimulation induced on Trichodesmium IMS101 resulting from 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 is predicted to cause a shift in the geographical distribution of Trichodesmium species toward higher latitudes, extending its niche to subtropical ocean regions and potentially reducing its coverage in tropical ocean basins.

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The dissolution behavior of biogenic calcites in seawater and a possible role for magnesium and organic carbon

We present the dissolution kinetics of mixed planktic foraminifera, the benthic foraminifera Amphistegina, the coccolithophore Emiliania huxleyi, and the soft coral Rhythismia fulvum in seawater. Dissolution rates were measured across a large range of saturation states (Ω = 0.99–0.2) by dissolving 13C-labeled calcites in natural seawater undersaturated with respect to calcite. 13C-label was incorporated into biogenic calcite by culturing marine calcifiers in 13C-labeled natural seawater. Net dissolution rates were calculated as the slope of seawater δ13C versus time in a closed seawater-calcite system. All calcites show distinct, nonlinear, dependencies on seawater saturation state when normalized by mass or by specific surface area. For example, coccolith calcite dissolves at a similar rate to inorganic calcite near equilibrium when normalized by surface area, but dissolves much more slowly far from equilibrium. Mass loss from foraminiferal tests is correlated with a decrease in Mg/Ca of the solid, indicating that Mg-rich phases are preferentially leached out at even mild undersaturations. Dissolution also appears to strongly affect test B/Ca. Finally, we provide an interpretation of surface area-normalized biogenic calcite dissolution rates as a function of their Mg and organic carbon content. Near-equilibrium dissolution rates of all calcites measured here show a strong, nonlinear dependence on Mg content. Far-from-equilibrium dissolution rates decrease strongly as a function of organic carbon content. These results help to build a framework for understanding the underlying mechanisms of rate differences between biogenic calcites, and bear important implications for the dissolution of high-Mg calcites in view of ocean acidification.

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Effects of seawater acidification and alkalization on the farmed seaweed, Pyropia haitanensis (Bangiales, Rhodophyta), grown under different irradiance conditions

Highlights

• Either seawater acidification and alkalization or reduced light inhibited nitrogen metabolism of Pyropia haitanensis
• Reduced irradiance alleviate negative effects of seawater alkalization on the algal growth and photosynthesis
• Lowered irradiance aggravated adverse impacts of seawater acidification on the growth and photosynthesis of P. haitanensis

Abstract

The thalli of Pyropia haitanensis were cultured under different pH levels (7.8, 8.2, and 9.0) and under decreased (60 μmol photons mm−2 s−1) and ambient (300 μmol photons m−2 s−1) levels of light irradiance conditions, aiming to examine the influence of different pH and decreased light irradiance on this farmed seaweed species in Southern China. Either the decreased (7.8) or increased (9.0) pH values in seawater inhibited nitrogen uptake rates and nitrate reductase activity of P. haitanensis. The capacity of nitrogen uptake and maximum inorganic carbon (Ci)-saturated photosynthetic rate (Vmax) were reduced in P. haitanensis grown at decreased irradiance compared with the algae grown at ambient irradiance. Decreased pH had no significant effect on the algal growth and photosynthesis under ambient light conditions, but it significantly inhibited growth and photosynthesis under decreased light conditions. Increased seawater pH resulted in decreased relative growth rate (RGR), maximal quantum yield of photosystem II ((Fv/Fm), and non-photochemical quenching (NPQ) of P. haitanensis when the algae were grown under ambient light conditions. However, a slight decrease was observed with decreasing growth irradiances. Collectively, our results indicated that either the changed pH (acidification and alkalization) or reduced irradiance displayed a disadvantageous influence on nitrogen metabolism of P. haitanensis. We suggested that, during P. haitanensis mariculture, the decreased light irradiance resulting from increasing algal mats density alleviates the negative effects of seawater alkalization, but it aggravates the adverse effects of seawater acidification on the growth and photosynthesis of the algae.

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Marine iron biogeochemistry under a changing climate: impact on the phytoplankton and the diazotroph communities

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.

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Nutrient co-limited Trichodesmium as nitrogen source or sink in a future ocean

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”

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.

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Comment on “The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium”

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.

Continue reading ‘Comment on “The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium”’

The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium

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.

Continue reading ‘The complex effects of ocean acidification on the prominent N2-fixing cyanobacterium Trichodesmium’

Influence of ocean acidification and deep water upwelling on oligotrophic plankton communities in the subtropical North Atlantic: Insights from an in situ mesocosm study

Oceanic uptake of anthropogenic carbon dioxide (CO2) causes pronounced shifts in marine carbonate chemistry and a decrease in seawater pH. Increasing evidence indicates that these changes – summarized by the term ocean acidification (OA) – can significantly affect marine food webs and biogeochemical cycles. However, current scientific knowledge is largely based on laboratory experiments with single species and artificial boundary conditions, whereas studies of natural plankton communities are still relatively rare. Moreover, the few existing community-level studies were mostly conducted in rather eutrophic environments, while less attention has been paid to oligotrophic systems such as the subtropical ocean gyres.

Here we report from a recent in situ mesocosm experiment off the coast of Gran Canaria in the eastern subtropical North Atlantic, where we investigated the influence of OA on the ecology and biogeochemistry of plankton communities in oligotrophic waters under close-to-natural conditions. This paper is the first in this Research Topic of Frontiers in Marine Biogeochemistry and provides (1) a detailed overview of the experimental design and important events during our mesocosm campaign, and (2) first insights into the ecological responses of plankton communities to simulated OA over the course of the 62-day experiment.

One particular scientific objective of our mesocosm experiment was to investigate how OA impacts might differ between oligotrophic conditions and phases of high biological productivity, which regularly occur in response to upwelling of nutrient-rich deep water in the study region. Therefore, we specifically developed a deep water collection system that allowed us to obtain ~85 m3 of seawater from ~650 m depth. Thereby, we replaced ~20% of each 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.

Continue reading ‘Influence of ocean acidification and deep water upwelling on oligotrophic plankton communities in the subtropical North Atlantic: Insights from an in situ mesocosm study’


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