Predictable ecological response to rising CO2 of a community of marine phytoplankton

Rising atmospheric CO₂ and ocean acidification are fundamentally altering conditions for life of all marine organisms, including phytoplankton. Differences in CO₂ related physiology between major phytoplankton taxa lead to differences in their ability to take up and utilize CO₂. These differences may cause predictable shifts in the composition of marine phytoplankton communities in response to rising atmospheric CO₂. We report an experiment in which seven species of marine phytoplankton, belonging to four major taxonomic groups (cyanobacteria, chlorophytes, diatoms, and coccolithophores), were grown at both ambient (500 μatm) and future (1,000 μatm) CO₂ levels. These phytoplankton were grown as individual species, as cultures of pairs of species and as a community assemblage of all seven species in two culture regimes (high‐nitrogen batch cultures and lower‐nitrogen semicontinuous cultures, although not under nitrogen limitation). All phytoplankton species tested in this study increased their growth rates under elevated CO₂ independent of the culture regime. We also find that, despite species‐specific variation in growth response to high CO₂, the identity of major taxonomic groups provides a good prediction of changes in population growth and competitive ability under high CO₂. The CO₂‐induced growth response is a good predictor of CO₂‐induced changes in competition (R² > .93) and community composition (R² > .73). This study suggests that it may be possible to infer how marine phytoplankton communities respond to rising CO₂ levels from the knowledge of the physiology of major taxonomic groups, but that these predictions may require further characterization of these traits across a diversity of growth conditions. These findings must be validated in the context of limitation by other nutrients. Also, in natural communities of phytoplankton, numerous other factors that may all respond to changes in CO2, including nitrogen fixation, grazing, and variation in the limiting resource will likely complicate this prediction.     

The Effect of Atmospheric Carbon Dioxide Elevation on Plant Growth in Freshwater Ecosystems

We developed a dynamic model to investigate the effect of atmospheric carbon dioxide (CO₂) increase on plant growth in freshwater ecosystems. Steady-state simulations were performed to analyze the response of phytoplankton and submerged macrophytes to atmospheric CO₂ elevation from 350 to 700 ppm. We studied various conditions that may affect this response, such as alkalinity, the air–water exchange rate of CO₂, the community respiration rate, and the phosphorus (P) supply rate. The increase in atmospheric CO₂ could affect submerged plant growth only under relatively eutrophic conditions and at a low community respiration rate. Alkalinity had little effect on the response of the different species. When the air–water exchange was low, the proportional effect of the CO₂ increase on plant growth was higher. Under eutrophic conditions, algae and macrophytes using CO₂ and HCO₃^– may double their growth rate due to atmospheric CO₂ elevation, while the growth of macrophytes restricted to CO₂ assimilation may be threefold. The differences in response of the species under various conditions indicate that the elevation of atmospheric CO₂ may induce drastic changes in the productivity and species dominance in freshwater systems.     

Long-term culture at elevated atmospheric CO2 fails to evoke specific adaptation in seven freshwater phytoplankton species

The concentration of CO₂ in the atmosphere is expected to double by the end of the century. Experiments have shown that this will have important effects on the physiology and ecology of photosynthetic organisms, but it is still unclear if elevated CO₂ will elicit an evolutionary response in primary producers that causes changes in physiological and ecological attributes. In this study, we cultured lines of seven species of freshwater phytoplankton from three major groups at current (approx. 380 ppm CO₂) and predicted future conditions (1000 ppm CO₂) for over 750 generations. We grew the phytoplankton under three culture regimes: nutrient-replete liquid medium, nutrient-poor liquid medium and solid agar medium. We then performed reciprocal transplant assays to test for specific adaptation to elevated CO₂ in these lines. We found no evidence for evolutionary change. We conclude that the physiology of carbon utilization may be conserved in natural freshwater phytoplankton communities experiencing rising atmospheric CO₂ levels, without substantial evolutionary change.     

Subtropical freshwater phytoplankton show a greater response to increased temperature than to increased pCO2

Global increases in atmospheric CO₂ and temperatures will impact aquatic systems, with freshwater habitats being affected. Some studies suggest that these conditions will promote cyanobacterial dominance. There is a need for a clearer picture of how algal species and strains within species will respond to higher temperatures and CO₂, especially in combination. This study examined two chlorophytes (Monoraphidium and Staurastrum), and two strains of the cyanobacterium Raphidiopsis raciborskii (straight S07 and coiled C03), to determine how the combination of higher temperature and CO₂ levels will affect their growth and maximum cell concentrations. Continuous cultures were used to compare the steady state cell concentrations at 28 °C and 30 °C, and CO₂ partial pressures (pCO₂), 400 and 750 ppm for all cultures, and in addition 1000 ppm at 28 °C for R. raciborskii strains. This study showed that, for all species, water temperature had a greater effect than higher pCO₂ on cell concentrations. There were clear differences in response between the chlorophyte species, with Monoraphidium preferring 28 °C and Staurastrum preferring 30 °C. There were also differences in response of the R. raciborskii strains to increasing temperature and pCO₂, with S07 having a greater increase in cell concentration. Genome analysis of R. raciborskii showed that the straight strain has five additional carbon acquisition genes (β-CA, chpY, cmpB, cmpD and NdhD4), indicative of increased carbon metabolism. These differences in the strains’ response to elevated pCO₂ will lead to changes in the species population structure and distribution in the water column. This study shows that it is important to examine the effects of both pCO₂ and temperature, and to consider strain variation, to understand how species composition of natural systems may change under future climate conditions.     

Population parameters and growth of Myzus persicae Sulzer (Hemiptera: Aphididae) under elevated CO2 concentrations in the air

The effects of three different CO₂ concentrations (400, 600, and 1000 ppm) on the population parameters and growth of the green peach aphid, Myzus persicae, were examined. Raw life history data from M. persicae were analyzed using an age-stage, two-sex life table to take into account the viable development rate among individuals. The population projections of M. persicae indicate the stage structure and variability of the population growth under different CO₂ concentrations based on an age-stage, two-sex life table analysis. Significantly longer oviposition duration and higher fecundity were observed under elevated CO₂ (600 and 1000 ppm) than those under ambient CO₂ (400 ppm). Furthermore, the M. persicae population reared under elevated CO₂ concentrations showed higher intrinsic and finite rates of population increase than under ambient CO₂ concentrations. These results indicate that the population parameters and growth of M. persicae were positively influenced in the fecundity by elevated CO₂ concentrations relative to the ambient CO₂. These findings indicate that it is basically remained to understand the direct effects of CO₂ elevation on the host plants, and the interaction between the host plants and M. persicae in the same CO₂ concentration for establishing more realistic population growth model systems for M. persicae in the aerial environment rising CO₂ concentration level.     

Influence of enhanced CO2 on growth and photosynthesis of the red algaeGracilaria sp. andG. chilensis

The influence of elevated CO₂ concentrations on growth and photosynthesis ofGracilaria sp. andG. chilensis was investigated in order to procure information on the effective utilization of CO₂. Growth of both was enhanced by CO₂ enrichment (air + 650 ppm CO₂, air + 1250 ppm CO₂, the enhancement being greater inGracilaria sp. Both species increased uptake of NO₃ ^– with CO₂ enrichment. Photosynthetic inorganic carbon uptake was depressed inG. chilensis by pre-culture (15 days) with CO₂ enrichment, but little affected inGracilaria sp. Mass spectrometric analysis showed that O₂ uptake was higher in the light than in the dark for both species and in both cases was higher inGracilaria sp. The higher growth enhancement inGracilaria sp. was attributed to greater depression of photorespiration by the enrichment of CO₂ in culture.     

The effect of atmospheric carbon dioxide concentrations on the performance of the mangrove Avicennia germinans over a range of salinities

By increasing water use efficiency and carbon assimilation, increasing atmospheric CO₂ concentrations could potentially improve plant productivity and growth at high salinities. To assess the effect of elevated CO₂ on the salinity response of a woody halophyte, we grew seedlings of the mangrove Avicennia germinans under a combination of five salinity treatments [from 5 to 65 parts per thousand (ppt)] and three CO₂ concentrations (280, 400 and 800 ppm). We measured survivorship, growth rate, photosynthetic gas exchange, root architecture and foliar nutrient and ion concentrations. The salinity optima for growth shifted higher with increasing concentrations of CO₂, from 0 ppt at 280 ppm to 35 ppt at 800 ppm. At optimal salinity conditions, carbon assimilation rates were significantly higher under elevated CO₂ concentrations. However, at salinities above the salinity optima, salinity had an expected negative effect on mangrove growth and carbon assimilation, which was not alleviated by elevated CO₂, despite a significant improvement in photosynthetic water use efficiency. This is likely due to non‐stomatal limitations to growth at high salinities, as indicated by our measurements of foliar ion concentrations that show a displacement of K⁺ by Na⁺ at elevated salinities that is not affected by CO₂. The observed shift in the optimal salinity for growth with increasing CO₂ concentrations changes the fundamental niche of this species and could have significant effects on future mangrove distribution patterns and interspecific interactions.     

The response of fast- and slow-growing Acacia species to elevated atmospheric CO2: an analysis of the underlying components of relative growth rate

In this study we assessed the impact of elevated CO₂ with unlimited water and complete nutrient on the growth and nitrogen economy of ten woody Acacia species that differ in relative growth rate (RGR). Specifically, we asked whether fast- and slow-growing species systematically differ in their response to elevated CO₂. Four slow-growing species from semi-arid environments (Acacia aneura, A. colei, A. coriacea and A. tetragonophylla) and six fast-growing species from mesic environments (Acacia dealbata, A. implexa, A. mearnsii, A. melanoxylon, A. irrorata and A. saligna) were grown in glasshouses with either ambient (˜350 ppm) or elevated (˜700 ppm) atmospheric CO₂. All species reached greater final plant mass with the exception of A. aneura, and RGR, averaged across all species, increased by 10% over a 12-week period when plants were exposed to elevated CO₂. The stimulation of RGR was evident throughout the 12-week growth period. Elevated CO₂ resulted in less foliage area per unit foliage dry mass, which was mainly the result of an increase in foliage thickness with a smaller contribution from greater dry matter content per unit fresh mass. The net assimilation rate (NAR, increase in plant mass per unit foliage area and time) of the plants grown at elevated CO₂ was higher in all species (on average 30% higher than plants in ambient CO₂) and was responsible for the increase in RGR. The higher NAR was associated with a substantial increase in foliar nitrogen productivity in all ten Acacia species. Plant nitrogen concentration was unaltered by growth at elevated CO₂ for the slow-growing Acacia species, but declined by 10% for faster-growing species. The rate of nitrogen uptake per unit root mass was higher in seven of the species when grown under elevated CO₂, and leaf area per unit root mass was reduced by elevated CO₂ in seven of the species. The absolute increase in RGR due to growth under elevated CO₂ was greater for fast- than for slow-growing Acacia species. Received: 21 December 1998 / Accepted: 31 May 1999     

Population parameters and growth of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae) under elevated CO2 concentrations

We evaluated the direct effects of three different CO₂ concentrations (400, 600 and 1,000 ppm) on the population parameters and growth of the bean bug, Riptortus pedestris, while being fed on soybean. The raw life history data from R. pedestris was analyzed using an age‐stage, two‐sex life table to take the viable development rate among individuals into account. Based on the age‐stage, two‐sex life table analysis, the population projections of R. pedestris provide the stage structure and variability of the population growth under different CO₂ treatments. Our results showed significantly shorter immature durations and higher pre‐adult survival rate under elevated CO₂ (1,000 ppm) than those under ambient CO₂ (400 ppm). The population of R. pedestris reared under elevated CO₂ conditions showed higher intrinsic and finite rates of increase but a lower mean generation time than R. pedestris reared under ambient CO₂ conditions. Our results show the population parameters and growth of R. pedestris are influenced by increased CO₂ relative to ambient CO₂ treatment. Further studies on the long‐term direct effects of different CO₂ levels on R. pedestris are essential to understand their population dynamics and to establish appropriate management strategies.     

Increase of atmospheric CO2 promotes phytoplankton productivity

It is usually thought that unlike terrestrial plants, phytoplankton will not show a significant response to an increase of atmospheric CO₂. Here we suggest that this view may be biased by a neglect of the effects of carbon (C) assimilation on the pH and the dissociation of the C species. We show that under eutrophic conditions, productivity may double as a result of doubling of the atmospheric CO₂ concentration. Although in practice productivity increase will usually be less, we still predict a productivity increase of up to 40% in marine species with a low affinity for bicarbonate. In eutrophic freshwater systems doubling of atmospheric CO₂ may result in an increase of the productivity of more than 50%. Freshwaters with low alkalinity appeared to be very sensitive to atmospheric CO₂ elevation. Our results suggest that the aquatic C sink may increase more than expected, and that nuisance phytoplankton blooms may be aggravated at elevated atmospheric CO₂ concentrations.     

The influence of increased temperature and carbon dioxide levels on the benthic/sea ice diatom <Emphasis Type="Italic">Navicula</Emphasis><Emphasis Type="Italic">directa</Emphasis>

Polar oceans are very susceptible to increased levels of atmospheric CO₂ and may act as the world’s largest sink for anthropogenic CO₂. Simultaneously, as atmospheric CO₂ increases, sea surface temperature rises due to global warming. These two factors are important in regulating microalgal ecophysiology, and it has been suggested that future global changes may significantly alter phytoplankton species composition. This study aims to investigate potential consequences of global change in terms of increased temperature and CO₂ enrichment on the benthic/sea ice diatom Navicula directa. In a laboratory experiment, the physiological response to elevated temperature and partial pressure of CO₂ (pCO₂) was investigated in terms of growth, photosynthetic activity and photosynthetic pigment composition. The experiment was performed under manipulated levels of pCO₂ (380 and 960 ppm) and temperature (0.5 and 4.5°C) to simulate a change from present levels to predicted levels during a worst-case scenario by the year 2100. After 7 days of treatment, no synergetic effects between temperature and pCO₂ were detected. However, elevated temperature promoted effective quantum yield of photosynthesis (∆F/F￠_m F^\prime_{\rm m} ) and increased growth rates by approximately 43%. Increased temperature also resulted in an altered pigment composition. In addition, enrichment of CO₂ appeared to reduce specific growth rates of N. directa. Even though growth rates were only reduced by approximately 5%, we hereby report that increased pCO₂ levels might also have potential negative effects on certain diatom strains.     

Interacting effects of atmospheric CO2 enrichment and solar radiation on growth of the aquatic fern Azolla filiculoides

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The effects of elevated CO2 and eutrophication on surface elevation gain in a European salt marsh

Salt marshes can play a vital role in mitigating the effects of global environmental change by dissipating incident storm wave energy and, through accretion, tracking increasing water depths consequent upon sea level rise. Atmospheric CO₂ concentrations and nutrient availability are two key variables that can affect the biological processes that contribute to marsh surface elevation gain. We measured the effects of CO₂ concentrations and nutrient availability on surface elevation change in intact mixed‐species blocks of UK salt marsh using six open‐top chambers receiving CO₂‐enriched (800 ppm) or ambient (400 ppm) air. We found more rapid surface elevation gain in elevated CO₂ conditions: an average increase of 3.4 mm over the growing season relative to ambient CO₂. Boosted regression analysis to determine the relative influence of different parameters on elevation change identified that a 10% reduction in microbial activity in elevated CO₂‐grown blocks had a positive influence on elevation. The biomass of Puccinellia maritima also had a positive influence on elevation, while other salt marsh species (e.g. Suaeda maritima) had no influence or a negative impact on elevation. Reduced rates of water use by the vegetation in the high CO₂ treatment could be contributing to elevation gain, either directly through reduced soil shrinkage or indirectly by decreasing microbial respiration rates due to lower redox levels in the soil. Eutrophication did not influence elevation change in either CO₂ treatment despite doubling aboveground biomass. The role of belowground processes (transpiration, root growth and decomposition) in the vertical adjustment of European salt marshes, which are primarily minerogenic in composition, could increase as atmospheric CO₂ concentrations rise and should be considered in future wetland models for the region. Elevated CO₂ conditions could enhance resilience in vulnerable systems such as those with low mineral sediment supply or where migration upwards within the tidal frame is constrained.     

Changes in atmospheric CO2 influence the allergenicity of Aspergillus fumigatus

Increased susceptibility to allergies has been documented in the Western world in recent decades. However, a comprehensive understanding of its causes is not yet available. It is therefore essential to understand trends and mechanisms of allergy‐inducing agents, such as fungal conidia. In this study, we investigated the hypothesis that environmental conditions linked to global atmospheric changes can affect the allergenicity of Aspergillus fumigatus, a common allergenic fungal species in indoor and outdoor environments and in airborne particulate matter. We show that fungi grown under present‐day CO₂ levels (392 ppm) exhibit 8.5 and 3.5 fold higher allergenicity compared to fungi grown at preindustrial (280 ppm) and double (560 ppm) CO₂ levels, respectively. A corresponding trend is observed in the expression of genes encoding for known allergenic proteins and in the major allergen Asp f1 concentrations, possibly due to physiological changes such as respiration rates and the nitrogen content of the fungus, influenced by the CO₂ concentrations. Increased carbon and nitrogen levels in the growth medium also lead to a significant increase in the allergenicity. We propose that climatic changes such as increasing atmospheric CO₂ levels and changes in the fungal growth medium may impact the ability of allergenic fungi such as A. fumigatus to induce allergies.     

Response of the floating aquatic fern <Emphasis Type="Italic">Azolla filiculoides</Emphasis> to elevated CO<Subscript>2</Subscript>, temperature,and phosphorus levels

Azolla filiculoides is a floating aquatic fern growing in tropical and temperate freshwater ecosystems. As A. filiculoides has symbiotic nitrogen-fixing cyanobacteria (Anabaena azollae) within its leaf cavities, it is cultivated in rice paddies to improve N availability and suppress other wetland weeds. To understand how C assimilation and N accumulation in A. filiculoides respond to elevated atmospheric carbon dioxide concentration (CO₂) in combination with P addition and higher temperatures, we conducted pot experiments during the summer of 2007 and 2008. In 2007, we grew A. filiculoides in pots at two treatment levels of added P fertilizer and at two levels of [CO₂] (380 ppm for ambient and 680 ppm for elevated [CO₂]) in controlled-environment chambers. In 2008, we grew A. filiculoides in four controlled-environment chambers at two [CO₂] levels and two temperature levels (34/26°C (day/night) and 29/21°C). We found that biomass and C assimilation by A. filiculoides were significantly increased by elevated [CO₂], temperature, and P level (all P < 0.01), with a significant interaction between elevated [CO₂] and added P (P < 0.01). Tissue N content was decreased by elevated [CO₂] and increased by higher temperature and P level (all P < 0.01). The acetylene reduction assay showed that the N-fixation activity of A. filiculoides was not significantly different under ambient and elevated [CO₂] but was significantly stimulated by P addition. N-fixation activity decreased at higher temperatures (34/26°C), indicating that 29/21°C was more suitable for A. azollae growth. Therefore, we conclude that the N accumulation potential of A. filiculoides under future climate warming depends primarily on the temperature change and P availability, and C assimilation should be increased by elevated [CO₂].     

The performance of Brevicoryne brassicae on ornamental cabbages grown in CO2-enriched atmospheres

The effect of different atmospheric CO₂ concentrations on life table parameters and the biology of the cabbage aphid, Brevicoryne brassicae, when fed on two cultivars of ornamental cabbage, was studied in a greenhouse designed for CO₂ studies. Aphid performance was influenced by increasing atmospheric CO₂ levels, significantly affecting the intrinsic rate of increase (r_m), finite rate of increase (λ), mean generation time (T), doubling time (DT), and pre-reproductive period. The longest pre-reproductive period was observed for aphids grown at 380 ppm CO₂. The intrinsic rate of natural increase was highest for aphids at 1050 ppm CO₂, because of their faster development, high daily rate of progeny production, and higher survivorship. Future elevated CO₂ concentrations will enhance aphid population outbreaks and consequently increase the damage caused.     

Contrasting effects of rising CO2 on primary production and ecological stoichiometry at different nutrient levels

Although rising CO₂ concentrations are thought to promote the growth and alter the carbon : nutrient stoichiometry of primary producers, several studies have reported conflicting results. To reconcile these contrasting results, we tested the following hypotheses: rising CO₂ levels (1) will increase phytoplankton biomass more at high nutrient loads than at low nutrient loads, but (2) will increase their carbon : nutrient stoichiometry more at low than at high nutrient loads. We formulated a mathematical model to predict dynamic changes in phytoplankton population density, elemental stoichiometry and inorganic carbon chemistry in response to rising CO₂. The model was tested in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa. The model predictions and experimental results confirmed the hypotheses. Our findings provide a novel theoretical framework to understand and predict effects of rising CO₂ concentrations on primary producers and their nutritional quality as food for herbivores under different nutrient conditions.     

The effect of pre-industrial and predicted atmospheric CO2 concentrations on the development of diazotrophic and non-diazotrophic cyanobacterium: Dolichospermum circinale and Microcystis aeruginosa

Photoautotrophs are capable of consuming high quantities of CO₂, yet scant research exists examining the influence of different CO₂ concentrations on the growth of freshwater diazotrophic or non-diazotrophic cyanobacteria. In this study, we cultured two cyanobacteria taxa (Dolichospermum circinale and Microcystis aeruginosa) within controlled atmospheric CO₂ chambers at pre-industrial, and post-industrial concentrations. Biovolume and chlorophyll a (Chl-a) differed as a consequence of the adjusted CO₂ gradients. Significantly higher biovolume measurements were observed in the elevated CO₂ treatment for the diazotrophic species in the initial experiment. However, a follow-up experiment, with a corrected culture replenishment regime showed Chl-a measurements were greater for the diazotrophic and non-diazotrophic species in the elevated CO₂ treatment. Increasing CO₂ presents a risk to already compromised eutrophic and hyper-eutrophic ecosystems, and we reason increasing CO₂ concentrations could affect photosynthetic performance and CO₂ assimilation of surface dwelling cyanobacteria. Further experimental work is required to establish ecological thresholds for freshwater ecosystems, as pH levels showed a measurable reduction within the elevated CO₂ treatments. As cyanobacteria species may respond quite differently to future CO₂ concentrations similar comparative studies should be carried out that focus on CO₂ dynamics and pH. The findings of the study indicate diazotrophic cyanobacteria growth in particular may benefit from elevated atmospheric CO₂ concentrations.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.