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One of the greatest threats facing the planet is the continued increase in excess greenhouse gasses, with CO 2 being the primary driver due to its rapid increase in only a century. Excess CO 2 is exacerbating known climate tipping points that will have cascading local and global effects including loss of biodiversity, global warming, and climate migration. However, global reduction of CO 2 emissions is not enough. Carbon dioxide removal (CDR) will also be needed to avoid the catastrophic effects of global warming. Although the drawdown and storage of CO 2 occur naturally via the coupling of the silicate and carbonate cycles, they operate over geological timescales (thousands of years). Here, we suggest that microbes can be used to accelerate this process, perhaps by orders of magnitude, while simultaneously producing potentially valuable by-products. This could provide both a sustainable pathway for global drawdown of CO 2 and an environmentally benign biosynthesis of materials. We discuss several different approaches, all of which involve enhancing the rate of silicate weathering. We use the silicate mineral olivine as a case study because of its favorable weathering properties, global abundance, and growing interest in CDR applications. Extensive research is needed to determine both the upper limit of the rate of silicate dissolution and its potential to economically scale to draw down significant amounts (Mt/Gt) of CO 2 Other industrial processes have successfully cultivated microbial consortia to provide valuable services at scale (e.g., wastewater treatment, anaerobic digestion, fermentation), and we argue that similar economies of scale could be achieved from this research., (Copyright © 2024 Cold Spring Harbor Laboratory Press; all rights reserved.)

Many marine organisms are exposed to decreasing O 2 levels due to warming-induced expansion of hypoxic zones and ocean deoxygenation (DeO 2 ). Nevertheless, effects of DeO 2 on phytoplankton have been neglected due to technical bottlenecks on examining O 2 effects on O 2 -producing organisms. Here we show that lowered O 2 levels increased primary productivity of a coastal phytoplankton assemblage, and enhanced photosynthesis and growth in the coastal diatom Thalassiosira weissflogii. Mechanistically, reduced O 2 suppressed mitochondrial respiration and photorespiration of T. weissflogii, but increased the efficiency of their CO 2 concentrating mechanisms (CCMs), effective quantum yield and improved light use efficiency, which was apparent under both ambient and elevated CO 2 concentrations leading to ocean acidification (OA). While the elevated CO 2 treatment partially counteracted the effect of low O 2 in terms of CCMs activity, reduced levels of O 2 still strongly enhanced phytoplankton primary productivity. This implies that decreased availability of O 2 with progressive DeO 2 could boost re-oxygenation by diatom-dominated phytoplankton communities, especially in hypoxic areas, with potentially profound consequences for marine ecosystem services in coastal and pelagic oceans., (© 2022. The Author(s).)

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 nonmodel, 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 (1,000+ 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., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

Coccolithophores are important oceanic primary producers not only in terms of photosynthesis but also because they produce calcite plates called coccoliths. Ongoing ocean acidification associated with changing seawater carbonate chemistry may impair calcification and other metabolic functions in coccolithophores. While short-term ocean acidification effects on calcification and other properties have been examined in a variety of coccolithophore species, long-term adaptive responses have scarcely been documented, other than for the single species Emiliania huxleyi. Here, we investigated the effects of ocean acidification on another ecologically important coccolithophore species, Gephyrocapsa oceanica, following 1,000 generations of growth under elevated CO 2 conditions (1,000 μatm). High CO 2 -selected populations exhibited reduced growth rates and enhanced particulate organic carbon (POC) and nitrogen (PON) production, relative to populations selected under ambient CO 2 (400 μatm). Particulate inorganic carbon (PIC) and PIC/POC ratios decreased progressively throughout the selection period in high CO 2 -selected cell lines. All of these trait changes persisted when high CO 2 -grown populations were moved back to ambient CO 2 conditions for about 10 generations. The results suggest that the calcification of some coccolithophores may be more heavily impaired by ocean acidification than previously predicted based on short-term studies, with potentially large implications for the ocean's carbon cycle under accelerating anthropogenic influences., (© 2018 The Authors. Global Change Biology Published by John Wiley & Sons Ltd.)

Nitrogen-fixing (N 2 ) 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 N 2 fixer Trichodesmium fundamentally shifts nitrogen metabolism toward organic-nitrogen scavenging following long-term high-CO 2 adaptation under iron and/or phosphorus (co)limitation. Global shifts in transcripts and proteins under high-CO 2 /Fe-limited and/or P-limited conditions include decreases in the N 2 -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 N 2 fixation. In a future high-CO 2 ocean, this whole-cell energetic reallocation toward organic nitrogen scavenging and away from N 2 fixation may reduce new-nitrogen inputs by Trichodesmium while simultaneously depleting the scarce fixed-nitrogen supplies of nitrogen-limited open-ocean ecosystems. IMPORTANCE Trichodesmium is among the most biogeochemically significant microorganisms in the ocean, since it supplies up to 50% of the new nitrogen supporting open-ocean food webs. We used Trichodesmium cultures adapted to high-CO 2 conditions for 7 years, followed by additional exposure to iron and/or phosphorus (co)limitation. We show that "future ocean" conditions of high CO 2 to projected future ocean conditions include decreases in the nitrogen-fixing nitrogenase enzymes coupled with major increases in enzymes that oxidize the abundant organic nitrogen source trimethylamine (TMA). Such a shift toward organic nitrogen uptake and away from nitrogen fixation may substantially reduce new-nitrogen inputs by Trichodesmium to projected future ocean conditions include decreases in the nitrogen-fixing nitrogenase enzymes coupled with major increases in enzymes that oxidize the abundant organic nitrogen source trimethylamine (TMA). Such a shift toward organic nitrogen uptake and away from nitrogen fixation may substantially reduce new-nitrogen inputs by Trichodesmium to the rest of the microbial community in the future high-CO 2 ocean, with potential global implications for ocean carbon and nitrogen cycling., (Copyright © 2018 American Society for Microbiology.)

Trichodesmium is a globally distributed cyanobacterium whose nitrogen-fixing capability fuels primary production in warm oligotrophic oceans. Like many photoautotrophs, Trichodesmium serves as a host to various other microorganisms, yet little is known about how this associated community modulates fluxes of environmentally relevant chemical species into and out of the supraorganismal structure. Here, we utilized metatranscriptomics to examine gene expression activities of microbial communities associated with Trichodesmium erythraeum (strain IMS101) using laboratory-maintained enrichment cultures that have previously been shown to harbor microbial communities similar to those of natural populations. In enrichments maintained under two distinct CO 2 concentrations for ∼8 years, the community transcriptional profiles were found to be specific to the treatment, demonstrating a restructuring of overall gene expression had occurred. Some of this restructuring involved significant increases in community respiration-related transcripts under elevated CO 2 , potentially facilitating the corresponding measured increases in host nitrogen fixation rates. Particularly of note, in both treatments, community transcripts involved in the reduction of nitrate, nitrite, and nitrous oxide were detected, suggesting the associated organisms may play a role in colony-level nitrogen cycling. Lastly, a taxon-specific analysis revealed distinct ecological niches of consistently cooccurring major taxa that may enable, or even encourage, the stable cohabitation of a diverse community within Trichodesmium consortia. IMPORTANCE is a genus of globally distributed, nitrogen-fixing marine cyanobacteria. As a source of new nitrogen in otherwise nitrogen-deficient systems, these organisms help fuel carbon fixation carried out by other more abundant photoautotrophs and thereby have significant roles in global nitrogen and carbon cycling. Members of the Trichodesmium genus tend to form large macroscopic colonies that appear to perpetually host an association of diverse interacting microbes distinct from the surrounding seawater, potentially making the entire assemblage a unique miniature ecosystem. Since its first successful cultivation in the early 1990s, there have been questions about the potential interdependencies between Trichodesmium genus tend to form large macroscopic colonies that appear to perpetually host an association of diverse interacting microbes distinct from the surrounding seawater, potentially making the entire assemblage a unique miniature ecosystem. Since its first successful cultivation in the early 1990s, there have been questions about the potential interdependencies between Trichodesmium and its associated microbial community and whether the host's seemingly enigmatic nitrogen fixation schema somehow involved or benefited from its epibionts. Here, we revisit these old questions with new technology and investigate gene expression activities of microbial communities living in association with Trichodesmium ., (Copyright © 2017 American Society for Microbiology.)

A mesocosm experiment was conducted in Wuyuan Bay (Xiamen), China, to investigate the effects of elevated pCO 2 on bloom formation by phytoplankton species previously studied in laboratory-based ocean acidification experiments, to determine if the indoor-grown species performed similarly in mesocosms under more realistic environmental conditions. We measured biomass, primary productivity and particulate organic carbon (POC) as well as particulate organic nitrogen (PON). Phaeodactylum tricornutum outcompeted Thalassiosira weissflogii and Emiliania huxleyi, comprising more than 99% of the final biomass. Mainly through a capacity to tolerate nutrient-limited situations, P. tricornutum showed a powerful sustained presence during the plateau phase of growth. Significant differences between high and low CO 2 treatments were found in cell concentration, cumulative primary productivity and POC in the plateau phase but not during the exponential phase of growth. Compared to the low pCO 2 (LC) treatment, POC increased by 45.8-101.9% in the high pCO 2 (HC) treated cells during the bloom period. Furthermore, respiratory carbon losses of gross primary productivity were found to comprise 39-64% for the LC and 31-41% for the HC mesocosms (daytime C fixation) in phase II. Our results suggest that the duration and characteristics of a diatom bloom can be affected by elevated pCO 2 . Effects of elevated pCO 2 observed in the laboratory cannot be reliably extrapolated to large scale mesocosms with multiple influencing factors, especially during intense algal blooms., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

We conducted shipboard microcosm experiments at both off-shore (SEATS) and near-shore (D001) stations in the northern South China Sea (NSCS) under three treatments, low temperature and low pCO 2 (LTLC), high temperature and low pCO 2 (HTLC), and high temperature and high pCO 2 (HTHC). Biomass of phytoplankton at both stations were enhanced by HT. HTHC did not affect phytoplankton biomass at station D001 but decreased it at station SEATS. HT alone increased net primary productivity by 234% at station SEATS and by 67% at station D001 but the stimulating effect disappeared when HC was combined. HT also increased respiration rate by 236% at station SEATS and by 87% at station D001 whereas HTHC reduced it by 61% at station SEATS and did not affect it at station D001. Overall, our findings indicate that the positive effect of ocean warming on phytoplankton assemblages in NSCS could be damped or offset by ocean acidification., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

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 CO 2 for a marine nitrogen-fixer. To address this gap, we cultured the globally important cyanobacterium Trichodesmium at both low and high CO 2 for 4.5 y, followed by reciprocal transplantation experiments to test for adaptation. Intriguingly, fitness actually increased in all high-CO 2 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 CO 2 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., Competing Interests: The authors declare no conflict of interest.

The unabated rise in anthropogenic CO₂ emissions is predicted to strongly influence the ocean's environment, increasing the mean sea-surface temperature by 4°C and causing a pH decline of 0.3 units by the year 2100. These changes are likely to affect the nutritional value of marine food sources since temperature and CO₂ can influence the fatty (FA) and amino acid (AA) composition of marine primary producers. Here, essential amino (EA) and polyunsaturated fatty (PUFA) acids are of particular importance due to their nutritional value to higher trophic levels. In order to determine the interactive effects of CO₂ and temperature on the nutritional quality of a primary producer, we analyzed the relative PUFA and EA composition of the diatom Cylindrotheca fusiformis cultured under a factorial matrix of 2 temperatures (14 and 19°C) and 3 partial pressures of CO₂ (180, 380, 750 μatm) for >250 generations. Our results show a decay of ~3% and ~6% in PUFA and EA content in algae kept at a pCO₂ of 750 μatm (high) compared to the 380 μatm (intermediate) CO₂ treatments at 14°C. Cultures kept at 19°C displayed a ~3% lower PUFA content under high compared to intermediate pCO₂, while EA did not show differences between treatments. Algae grown at a pCO₂ of 180 μatm (low) had a lower PUFA and AA content in relation to those at intermediate and high CO₂ levels at 14°C, but there were no differences in EA at 19°C for any CO₂ treatment. This study is the first to report adverse effects of warming and acidification on the EA of a primary producer, and corroborates previous observations of negative effects of these stressors on PUFA. Considering that only ~20% of essential biomolecules such as PUFA (and possibly EA) are incorporated into new biomass at the next trophic level, the potential impacts of adverse effects of ocean warming and acidification at the base of the food web may be amplified towards higher trophic levels, which rely on them as source of essential biomolecules.

Increasing pCO2 (partial pressure of CO2 ) in an "acidified" ocean will affect phytoplankton community structure, but manipulation experiments with assemblages briefly acclimated to simulated future conditions may not accurately predict the long-term evolutionary shifts that could affect inter-specific competitive success. We assessed community structure changes in a natural mixed dinoflagellate bloom incubated at three pCO2 levels (230, 433, and 765 ppm) in a short-term experiment (2 weeks). The four dominant species were then isolated from each treatment into clonal cultures, and maintained at all three pCO2 levels for approximately 1 year. Periodically (4, 8, and 12 months), these pCO2 -conditioned clones were recombined into artificial communities, and allowed to compete at their conditioning pCO2 level or at higher and lower levels. The dominant species in these artificial communities of CO2 -conditioned clones differed from those in the original short-term experiment, but individual species relative abundance trends across pCO2 treatments were often similar. Specific growth rates showed no strong evidence for fitness increases attributable to conditioning pCO2 level. Although pCO2 significantly structured our experimental communities, conditioning time and biotic interactions like mixotrophy also had major roles in determining competitive outcomes. New methods of carrying out extended mixed species experiments are needed to accurately predict future long-term phytoplankton community responses to changing pCO2 ., (© 2013 The Author(s). Evolution © 2013 The Society for the Study of Evolution.)

Anthropogenic CO(2) is progressively acidifying the ocean, but the responses of harmful algal bloom species that produce toxins that can bioaccumulate remain virtually unknown. The neurotoxin domoic acid is produced by the globally-distributed diatom genus Pseudo-nitzschia. This toxin is responsible for amnesic shellfish poisoning, which can result in illness or death in humans and regularly causes mass mortalities of marine mammals and birds. Domoic acid production by Pseudo-nitzschia cells is known to be regulated by nutrient availability, but potential interactions with increasing seawater CO(2) concentrations are poorly understood. Here we present experiments measuring domoic acid production by acclimatized cultures of Pseudo-nitzschia fraudulenta that demonstrate a strong synergism between projected future CO(2) levels (765 ppm) and silicate-limited growth, which greatly increases cellular toxicity relative to growth under modern atmospheric (360 ppm) or pre-industrial (200 ppm) CO(2) conditions. Cellular Si:C ratios decrease with increasing CO(2), in a trend opposite to that seen for domoic acid production. The coastal California upwelling system where this species was isolated currently exhibits rapidly increasing levels of anthropogenic acidification, as well as widespread episodic silicate limitation of diatom growth. Our results suggest that the current ecosystem and human health impacts of toxic Pseudo-nitzschia blooms could be greatly exacerbated by future ocean acidification and 'carbon fertilization' of the coastal ocean.

Iglesias-Rodriguez et al. (Research Articles, 18 April 2008, p. 336) reported that the coccolithophore Emiliania huxleyi doubles its organic matter production and calcification in response to high carbon dioxide partial pressures, contrary to previous laboratory and field studies. We argue that shortcomings in their experimental protocol compromise the interpretation of their data and the resulting conclusions.

Anthropogenic greenhouse gas emissions are leading to global temperature increases, ocean acidification, and significant ecosystem impacts. Given current emissions trajectories, the IPCC's reports indicate that rapid abatement of CO 2 emissions and development of carbon dioxide removal (CDR) strategies are needed to address legacy and difficult-to-abate emissions sources. These CDR methods must efficiently and safely sequester gigatons of atmospheric CO 2. Coastal enhanced weathering (CEW) via the addition of the common mineral olivine to coastal waters is one promising approach to enhance ocean alkalinity for large-scale CDR. As olivine weathers, it releases several biologically active dissolution products, including alkalinity, trace metals, and the nutrient silicate. Released trace metals can serve as micronutrients but may also be toxic at high concentrations to marine biota, including phytoplankton, which lie at the base of marine food webs. We grew six species representing several globally important phytoplankton species under elevated concentrations of olivine dissolution products via a synthetic olivine leachate (OL) based on olivine's elemental composition. We monitored their physiological and biogeochemical responses, which allowed us to determine physiological impacts and thresholds at elevated olivine leachate concentrations, in addition to individual effects of specific constituents. We found both positive and neutral responses but no evident toxic effects for two silicifying diatoms, a calcifying coccolithophore, and three cyanobacteria. In both single and competitive co-cultures, silicifiers and calcifiers benefited from olivine dissolution products like iron and silicate or enhanced alkalinity, respectively. The non-N 2 -fixing picocyanobacterium could use synthetic olivine-derived iron for growth, while N 2 -fixing cyanobacteria could not. However, other trace metals like nickel and cobalt supported cyanobacterial growth across both groups. Growth benefits to phytoplankton groups in situ will depend on species-specific responses and ambient concentrations of other required nutrients. Results suggest olivine dissolution products appear unlikely to cause negative physiological effects for any of the phytoplankton examined, even at high concentrations, and may support growth of particular taxa under some conditions. Future studies can shed light on long-term eco-evolutionary responses to olivine exposure and on the potential effects that marine microbes may in turn have on olivine dissolution rates and regional biogeochemistry. [ABSTRACT FROM AUTHOR]

Diatoms are often considered to be a single functional group, yet there is a great deal of morphological, genetic and ecological diversity within the class. How these differences will translate into species-specific responses to rapid changes in the ocean environment resulting from climate change and eutrophication is currently poorly understood. We investigated the response of a natural diatom-dominated assemblage in coastal California waters to interactions between the variables nitrogen source (nitrate and urea), temperature (19 and 23°C) and CO2 (380 and 800 ppm) in a factorial experimental matrix using continuous culture (ecostat) methods. The community included diatoms of the cosmopolitan genera Pseudo-nitzschia and Chaetoceros, as well as Leptocylindrus and Cylindrotheca. Our results demonstrate strong interactive effects of these variables on community composition; notably, nitrogen source alone and nitrogen and CO2 together had a much greater influence on diatom community structure at 23°C compared with 19°C. In addition, warming and acidification interactions significantly increased cellular quotas of the neurotoxin domoic acid produced by Pseudo-nitzschia multiseries. In general, the effects observed for the factors tested differed significantly between the various diatom genera in this assemblage, suggesting potentially divergent responses of some of these ecologically and biogeochemically important phytoplankton taxa to interactions between global-scale and local-scale anthropogenic stressors in a changing ocean. [ABSTRACT FROM AUTHOR]

We investigated the effects of temperature and CO2 variation on the growth and elemental composition of cultures of the diatom Pseudo-nitzschia subcurvata and the prymnesiophyte Phaeocystis antarctica, two ecologically dominant phytoplankton species isolated from the Ross Sea, Antarctica. To obtain thermal functional response curves, cultures were grown across a range of temperatures from 0 to 14 °C. In addition, a co-culturing experiment examined the relative abundance of both species at 0 and 6 °C. CO2 functional response curves were conducted from 100 to 1730 ppm at 2 and 8 °C to test for interactive effects between the two variables. The growth of both phytoplankton was significantly affected by temperature increase, but with different trends. Growth rates of P. subcurvata increased with temperature from 0 °C to maximum levels at 8 °C, while the growth rates of P. antarctica only increased from 0 to 2 °C. The maximum thermal limits of P. subcurvata and P. antarctica where growth stopped completely were 14 and 10 °C, respectively. Although P. subcurvata outgrew P. antarctica at both temperatures in the co-incubation experiment, this happened much faster at 6 than at 0 °C. For P. subcurvata, there was a significant interactive effect in which the warmer temperature decreased the CO2 half-saturation constant for growth, but this was not the case for P. antarctica. The growth rates of both species increased with CO2 increases up to 425 ppm, and in contrast to significant effects of temperature, the effects of CO2 increase on their elemental composition were minimal. Our results suggest that future warming may be more favorable to the diatom than to the prymnesiophyte, while CO2 increases may not be a major factor in future competitive interactions between Pseudo-nitzschia subcurvata and Phaeocystis antarctica in the Ross Sea. [ABSTRACT FROM AUTHOR]

Biological effects of ultraviolet radiation (UVR; 280-400 nm) on marine primary producers are of general concern, as oceanic carbon fixers that contribute to the marine biological CO2 pump are being exposed to increasing UV irradiance due to global change and ozone depletion. We investigated the effects of UV-B (280-320 nm) and UV-A (320-400 nm) on the biogeochemically-critical filamentous marine N2-fixing cyanobacterium Trichodesmium (strain IMS101) using a solar simulator as well as under natural solar radiation. Short exposure to UV-B, UV-A, or integrated total UVR significantly reduced the effective quantum yield of photosystem II (PSII) and photosynthetic carbon and N2 fixation rates. Cells acclimated to low light were more sensitive to UV exposure compared to high-light grown ones, which had more UV absorbing compounds, most likely mycosporine-like amino acids (MAAs). After acclimation under natural sunlight, the specific growth rate was lower (by up to 44 %), MAAs content was higher, and average trichome length was shorter (by up to 22 %) in the full spectrum of solar radiation with UVR, than under a photosynthetically active radiation (PAR) alone treatment (400-700 nm). These results suggest that prior shipboard experiments in UV-opaque containers may have substantially overestimated in-situ nitrogen fixation rates by Trichodesmium, and that natural and anthropogenic elevation of UV radiation intensity could significantly inhibit this vital source of new nitrogen to the current and future oligotrophic oceans. [ABSTRACT FROM AUTHOR]

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 (longterm) 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. [ABSTRACT FROM AUTHOR]

Marine dinitrogen (N2)-fixing cyanobacteria have large impacts on global biogeochemistry as they fix carbon dioxide (CO2) and fertilize oligotrophic ocean waters with new nitrogen. Iron (Fe) and phosphorus (P) are the two most important limiting nutrients for marine biological N2 fixation, and their availabilities vary between major ocean basins and regions. A long-standing question concerns the ability of two globally dominant N2-fixing cyanobacteria, unicellular Crocosphaera and filamentous Trichodesmium, to maintain relatively high N2-fixation rates in these regimes where both Fe and P are typically scarce. We show that under P-deficient conditions, cultures of these two cyanobacteria are able to grow and fix N2 faster when Fe deficient than when Fe replete. In addition, growth affinities relative to P increase while minimum concentrations of P that support growth decrease at low Fe concentrations. In Crocosphaera, this effect is accompanied by a reduction in cell sizes and elemental quotas. Relatively high growth rates of these two biogeochemically critical cyanobacteria in low-P, low-Fe environments such as those that characterize much of the oligotrophic ocean challenge the common assumption that low Fe levels can have only negative effects on marine primary producers. The closely interdependent influence of Fe and P on N2-fixing cyanobacteria suggests that even subtle shifts in their supply ratio in the past, present and future oceans could have large consequences for global carbon and nitrogen cycles. [ABSTRACT FROM AUTHOR]

Much of the bioavailable nitrogen that supports open ocean food webs and biogeochemical cycles is fixed from the atmosphere by marine cyanobacteria of the genera Trichodesmium and Crocosphaera. In previous experiments carried out with a limited set of cyanobacterial isolates, rates of cyanobacterial nitrogen fixation were shown to increase with carbon dioxide concentrations. Here, we report results from a series of laboratory experiments in which we grew seven strains of Trichodesmium and Crocosphaera from the Atlantic and Pacific oceans under a wide range of carbon dioxide concentrations, and monitored rates of nitrogen fixation and growth. We document large, strain-specific differences in the relationship between nitrogen fixation and carbon dioxide concentration, suggesting that individual strains within each genus are adapted to grow and fix nitrogen at different concentrations of carbon dioxide. We apply kinetic constants from the individual carbon dioxide response curves to an illustrative biogeochemical model of the ocean in 2100, which suggests that strains adapted to high carbon dioxide concentrations could potentially be favoured in a future acidified ocean. We suggest that surface ocean carbon dioxide concentrations could constitute a previously unrecognized selective force that shapes the community composition and diversity of nitrogen-fixing cyanobacteria. [ABSTRACT FROM AUTHOR]

Coastal upwelling regimes associated with eastern boundary currents are the most biologically productive ecosystems in the ocean. As a result, they play a disproportionately important role in the microbially mediated cycling of marine nutrients. These systems are characterized by strong natural variations in carbon dioxide concentrations, pH, nutrient levels and sea surface temperatures on both seasonal and interannual timescales. Despite this natural variability, changes resulting from human activities are starting to emerge. Carbon dioxide derived from fossil fuel combustion is adding to the acidity of upwelled low-pH waters. Low-oxygen waters associated with coastal upwelling systems are growing in their extent and intensity as a result of a rise in upper ocean temperatures and productivity. And nutrient inputs to the coastal ocean continue to grow. Coastal upwelling systems may prove more resilient to changes resulting from human activities than other ocean ecosystems because of their ability to function under extremely variable conditions. Nevertheless, shifts in primary production, fish yields, nitrogen gain and loss, and the flux of climate-relevant gases could result from the perturbation of these highly productive and dynamic ecosystems. [ABSTRACT FROM AUTHOR]

We describe interactive effects of total phosphorus (total P = 0.1-4.0 µmol L-1; added as H2NaPO4), irradiance (40 and 150 µmol quanta m-2 s-1), and the partial pressure of carbon dioxide (PCO2 19 and 81 Pa, i.e., 190 and 800 ppm) on growth and CO2- and dinitrogen (N2)-fixation rates of the unicellular N2-fixing cyanobacterium Crocosphaera watsonii (WH0003) isolated from the Pacific Ocean near Hawaii. In semicontinuous cultures of C. watsonii, elevated positively affected growth and CO2- and N2-fixation rates under high light. Under low light, elevated positively affected growth rates at all concentrations of P, but CO2- and N2-fixation rates were affected by elevated only when P was low. In both high-light and low-light cultures, the total P requirements for growth and CO2- and N2-fixation declined as increased. The minimum concentration (Cmin) of total P and half-saturation constant (K1/2) for growth and CO2- and N2-fixation rates with respect to total P were reduced by 0.05 µmol L-1 as a function of elevated . We speculate that low P requirements under high resulted from a lower energy demand associated with carbon-concentrating mechanisms in comparison with low-cultures. There was also a 0.10 µmol L-1 increase in Cmin and K1/2 for growth and N2 fixation with respect to total P as a function of increasing light regardless of concentration. We speculate that cellular P concentrations are responsible for this shift through biodilution of cellular P and possibly cellular P uptake systems as a function of increasing light. Changing concentrations of P, CO2, and light have both positive and negative interactive effects on growth and CO2-, and N2-fixation rates of unicellular oxygenic diazotrophs like C. watsonii. [ABSTRACT FROM AUTHOR]

The diazotrophic cyanobacteria Trichodesmium spp. contribute approximately half of the known marine dinitrogen (N2) fixation. Rapidly changing environmental factors such as the rising atmospheric partial pressure of carbon dioxide (pCO2) and shallower mixed layers (higher light intensities) are likely to affect N2-fixation rates in the future ocean. Several studies have documented that N2 fixation in laboratory cultures of T. erythraeum increased when pCO2 was doubled from present-day atmospheric concentrations (∼380 ppm) to projected future levels (∼750 ppm). We examined the interactive effects of light and pCO2 on two strains of T. erythraeum Ehrenb. (GBRTRLI101 and IMS101) in laboratory semicontinuous cultures. Elevated pCO2 stimulated gross N2-fixation rates in cultures growing at 38 μmol quanta · m−2 · s−1 (GBRTRLI101 and IMS101) and 100 μmol quanta · m−2 · s−1 (IMS101), but this effect was reduced in both strains growing at 220 μmol quanta · m−2 · s−1. Conversely, CO2-fixation rates increased significantly ( P < 0.05) in response to high pCO2 under mid- and high irradiances only. These data imply that the stimulatory effect of elevated pCO2 on CO2 fixation and N2 fixation by T. erythraeum is correlated with light. The ratio of gross:net N2 fixation was also correlated with light and trichome length in IMS101. Our study suggests that elevated pCO2 may have a strong positive effect on Trichodesmium gross N2 fixation in intermediate and bottom layers of the euphotic zone, but perhaps not in light-saturated surface layers. Climate change models must consider the interactive effects of multiple environmental variables on phytoplankton and the biogeochemical cycles they mediate. [ABSTRACT FROM AUTHOR]

Phytoplankton growth can be limited by numerous inorganic nutrients and organic growth factors. Using the subarctic diatom Attheya sp. in culture studies, we examined how the availability of vitamin B12 and carbon dioxide partial pressure (pCO2) influences growth rate, primary productivity, cellular iron (Fe), cobalt (Co), zinc (Zn) and cadmium (Cd) quotas, and the net use efficiencies (NUEs) of these bioactive trace metals (mol C fixed per mol cellular trace metal per day). Under B12-replete conditions, cells grown at high pCO2 had lower Fe, Zn and Cd quotas, and used those trace metals more efficiently in comparison with cells grown at low pCO2. At high pCO2, B12-limited cells had ∼50% lower specific growth and carbon fixation rates, and used Fe ∼15-fold less efficiently, and Zn and Cd ∼3-fold less efficiently, in comparison with B12-replete cells. The observed higher Fe, Zn and Cd NUE under high pCO2/B12-replete conditions are consistent with predicted downregulation of carbon-concentrating mechanisms. Co quotas of B12-replete cells were ∼5- to 14-fold higher in comparison with B12-limited cells, suggesting that >80% of cellular Co of B12-limited cells was likely from B12. Our results demonstrate that CO2 and vitamin B12 interactively influence growth, carbon fixation, trace metal requirements and trace metal NUE of this diatom. This suggests the need to consider complex feedback interactions between multiple environmental factors for this biogeochemically critical group of phytoplankton in the last glacial maximum as well as the current and future changing ocean. [ABSTRACT FROM AUTHOR]

The potential impact of elevated sea surface temperature (SST) and pCO2 on algal community structure and particulate dimethylsulfoniopropionate (DMSPp) concentrations in the southeastern Bering Sea was examined using a shipboard 'Ecostat' continuous culture system. The ecostat system was used to mimic the conditions projected to exist in the world's oceans by the end of this century (i.e. elevated pCO2 (750 ppm) and elevated SST (ambient + 4°C). Two experiments were conducted using natural phytoplankton assemblages from the high-nutrient low-chlorophyll (HNLC) central basin and from the middle domain of the southeastern continental shelf. At the HNLC site, the relative abundances of haptophytes and pelagophytes were higher and the relative abundance of diatoms lower under 'greenhouse' conditions (i.e. combined 750 ppm CO2 and elevated temperature) than control conditions (380 ppm CO2 and ambient temperature). This shift in algal community structure was accompanied by increases in DMSPp (2-3 fold), DMSPp:Chl a (2-3 fold) and DMSP:PON (2 fold). At the continental shelf site, the changes in the relative abundances of haptophytes, pelagophytes and diatoms under 'greenhouse' conditions were similar to those observed at the HNLC site, with 2.5 fold increases in DMSPp, 50-100% increases in DMSPp:Chl a and 1.8 fold increases in DMSP:PON. At both locations, changes in community structure and the DMSPp parameters were largely driven by increasing temperature. The observed changes were also consistent with the phytoplankton-DMS-albedo climate feedback mechanism proposed in the Charlson-Lovelock-Andreae-Warren (CLAW) hypothesis. [ABSTRACT FROM AUTHOR]

Ocean acidification produced by dissolution of anthropogenic carbon dioxide (CO2) emissions in seawater has profound consequences for marine ecology and biogeochemistry. The oceans have absorbed one-third of CO2 emissions over the past two centuries, altering ocean chemistry, reducing seawater pH, and affecting marine animals and phytoplankton in multiple ways. Microbially mediated ocean biogeochemical processes will be pivotal in determining how the earth system responds to global environmental change; however, how they may be altered by ocean acidification is largely unknown. We show here that microbial nitrification rates decreased in every instance when pH was experimentally reduced (by 0.05-0.14) at multiple locations in the Atlantic and Pacific Oceans. Nitrification is a central process in the nitrogen cycle that produces both the greenhouse gas nitrous oxide and oxidized forms of nitrogen used by phytoplankton and other microorganisms in the sea; at the Bermuda Atlantic Time Series and Hawaii Ocean Time-series sites, experimental acidification decreased ammonia oxidation rates by 38% and 36%. Ammonia oxidation rates were also strongly and inversely correlated with pH along a gradient produced in the oligotrophic Sargasso Sea (r² = 0.87, P < 0.05). Across all experiments, rates declined by 8-38% in low pH treatments, and the greatest absolute decrease occurred where rates were highest off the California coast. Collectively our results suggest that ocean acidification could reduce nitrification rates by 3-44% within the next few decades, affecting oceanic nitrous oxide production, reducing supplies of oxidized nitrogen in the upper layers of the ocean, and fundamentally altering nitrogen cycling in the sea. [ABSTRACT FROM AUTHOR]

The article details the findings made during an incubation experiment and discusses the changes in dimethylsulfoniopropionate (DMSP) concentrations and DMSP-lyase activity (DLA) that result from changes in algal community composition in the North Atlantic spring bloom. It describes the effect of both atmospheric temperature and carbon dioxide on biogeochemical processes and algal community structure. It also illustrates changes in and connections between microzooplankton grazing dynamics and algal community structure.

The article focuses on a study which investigates the effects of atmospheric carbon dioxide and temperature on algal community, phytoplankton succession and elemental cycling in the North Atlantic spring bloom area. The study conducts a shipboard continuous culture incubation with the use of a bloom phytoplankton community. Findings of this study are presented in three papers, one on phytoplankton community and biogeochemical responses, another on microzooplankton dynamics and one on dimethylsulfoniopropionate. It concludes that atmospheric carbon dioxide enrichment effects the phytoplankton community of the spring bloom area.

We examined the effects of increased temperature, pCO2, and irradiance on a calcifying strain of the marine coccolithophore Emiliania huxleyi in semi-continuous laboratory cultures. Emiliania huxleyi CCMP 371 was cultured in four temperature and pCO2 treatments at both low and high irradiance (50 and 400 µmol photons m-2 s-1): (i) 20°C and 375 ppm CO2 (ambient control); (ii) 20°C and 750 ppm CO2 (high pCO2); (iii) 24°C and 375 ppm CO2 (high temperature); and (iv) 24°C and 750 ppm CO2 ('greenhouse'). The growth of E. huxleyi was greatly accelerated by elevated temperature at low irradiance. Photosynthesis was significantly promoted by increases in both pCO2 and temperature at both irradiances. Higher cellular C/P ratios were found in the higher CO2 treatments at high irradiance, indicating a reduced requirement for P. The PIC/POC (particulate inorganic to organic carbon) ratio remained constant at low light, regardless of CO2 or temperature conditions. However, both the cellular PIC content and PIC/POC ratio were greatly decreased by elevated irradiance, and were further decreased by increased pCO2 only at high light, indicating a combined effect of CO2 and light on calcification. These results suggest that future trends of CO2 enrichment, sea-surface warming and exposure to higher mean irradiances from intensified stratification will have a large influence on the growth of Emiliania huxleyi, and potentially on the PIC/POC 'rain ratio'. Our study demonstrates that it is possible to obtain a more complete picture of global change impacts on marine phytoplankton by designing experiments that consider multiple global change variables and their mutual interactions. [ABSTRACT FROM AUTHOR]

Little is known about the combined impacts of future CO2 and temperature increases on the growth and physiology of marine picocyanobacteria. We incubated Synechococcus and Prochlorococcus under present-day (380 ppm) or predicted year-2100 CO2 levels (750 ppm), and under normal versus elevated temperatures (+4°C) in semicontinuous cultures. Increased temperature stimulated the cell division rates of Synechococcus but not Prochlorococcus. Doubled CO2 combined with elevated temperature increased maximum chl a–normalized photosynthetic rates of Synechococcus four times relative to controls. Temperature also altered other photosynthetic parameters (α, Φmax, Ek, and ) in Synechococcus, but these changes were not observed for Prochlorococcus. Both increased CO2 and temperature raised the phycobilin and chl a content of Synechococcus, while only elevated temperature increased divinyl chl a in Prochlorococcus. Cellular carbon (C) and nitrogen (N) quotas, but not phosphorus (P) quotas, increased with elevated CO2 in Synechococcus, leading to ∼20% higher C:P and N:P ratios. In contrast, Prochlorococcus elemental composition remained unaffected by CO2, but cell volume and elemental quotas doubled with increasing temperature while maintaining constant stoichiometry. Synechococcus showed a much greater response to CO2 and temperature increases for most parameters measured, compared with Prochlorococcus. Our results suggest that global change could influence the dominance of Synechococcus and Prochlorococcus ecotypes, with likely effects on oligotrophic food-web structure. However, individual picocyanobacteria strains may respond quite differently to future CO2 and temperature increases, and caution is needed when generalizing their responses to global change in the ocean. [ABSTRACT FROM AUTHOR]

The performance of capacitive micromachined ultrasonic transducers (cMUTs) was investigated at low pressures in various gases such as air, carbon dioxide, and helium. The aim was to replicate the pressure conditions likely to meet on the surface of other planets such as Mars, where ultrasonic wind velocity measurements might be possible. It is demonstrated that cMUTs are capable of operating at low pressures, and the response to pressures below terrestrial atmospheric values is observed experimentally and compared to theoretical predictions. The center frequency of Operation and sensitivity are both observed to be affected by changing pressures. [ABSTRACT FROM AUTHOR]

The article focuses on a study published in the periodical "Nature Climate Change" on the responses of marine phytoplankton to increasing carbon dioxide (CO2) and acidification. It states that the study used Ostreococcus tauri ecotypes, a globally distributed green alga. It notes that the ecotypes showed increased growth and photosynthetic rates. It mentions that the ecotypes may achieve competitive dominance of less responsive ones in acidified ocean.

Highlights: [•] Conditioning of a clonal culture of Alexandrium catenella to a factorial combination of environmental change variables; CO2, temperature, phosphorus. [•] Measurements of growth and cellular toxins after eight months of conditioning. [•] Increased CO2 led to enhanced cellular toxicity. [•] Combined high CO2, low temperature and phosphate-limitation were the ‘most toxic’. [Copyright &y& Elsevier]