Posts Tagged 'nitrogen fixation'

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.

Continue reading ‘Long-term m5C methylome dynamics parallel phenotypic adaptation in the cyanobacterium Trichodesmium’

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.

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

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.

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

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.

Continue reading ‘Environmental and nutrient controls of marine nitrogen fixation’

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.

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

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.

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

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.

Continue reading ‘Marine iron biogeochemistry under a changing climate: impact on the phytoplankton and the diazotroph communities’

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.

Continue reading ‘Nutrient co-limited Trichodesmium as nitrogen source or sink in a future ocean’

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.

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

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”’


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