CO2 Mesocosms Research Articles

Plankton Community Respiration and ETS Activity Under Variable CO2 and Nutrient Fertilization During a Mesocosm Study in the Subtropical North Atlantic

The enzymatic electron transport system (ETS) assay is frequently used as a proxy of respiratory activity in planktonic communities. It is thought to estimate the maximum overall activity of the enzymes associated with the respiratory ETS systems in both eukaryotic and prokaryotic organisms. Thus, in order to derive actual respiration rates (R) from ETS it is necessary to determine empirical R/ETS conversion algorithms. In this study we explore the temporal development of R and ETS activity in natural plankton communities (from bacteria to large phytoplankton) enclosed in mesocosms, treated with different CO2 concentrations. The experiment lasted 30 days, during which abrupt changes in community structure and biomass occurred through a sharp transition from oligotrophy (phase I) to highly eutrophic conditions (phase II) after nutrient-induced fertilization (day 18). R and ETS did not show any response to CO2 under oligotrophic conditions, but R increased significantly more in the two high CO2 mesocosms after fertilization, coinciding with a sharp rise in large phytoplankton (mostly diatoms). R and ETS were significantly correlated only during the eutrophic phase. The R/ETS ranged more than threefold in magnitude during the experiment, with phase-averaged values significantly higher under oligotrophic conditions (0.7-1.1) than after nutrient fertilization (0.5-0.7). We did not find any significant relationship between R/ETS and community structure or biomass, although R correlated significantly with total biomass after fertilization in the four mesocosms. Multiple stepwise regression models show that large phytoplankton explains most of the variance in R during phases I (86%) and II (53%) and of ETS (86%) during phase II, while picophytoplankton contributes up to 73% to explain the variance in the ETS model during phase I. Our results suggest that R/ETS may be too variable in the ocean as to apply constant values to different communities living under contrasting environmental conditions. Controlled experiments with natural communities, like the present one, would help to constrain the range of variability of the R/ETS ratio, and to understand the factors driving it.

Community barcoding reveals little effect of ocean acidification on the composition of coastal plankton communities: Evidence from a long-term mesocosm study in the Gullmar Fjord, Skagerrak.

The acidification of the oceans could potentially alter marine plankton communities with consequences for ecosystem functioning. While several studies have investigated effects of ocean acidification on communities using traditional methods, few have used genetic analyses. Here, we use community barcoding to assess the impact of ocean acidification on the composition of a coastal plankton community in a large scale, in situ, long-term mesocosm experiment. High-throughput sequencing resulted in the identification of a wide range of planktonic taxa (Alveolata, Cryptophyta, Haptophyceae, Fungi, Metazoa, Hydrozoa, Rhizaria, Straminipila, Chlorophyta). Analyses based on predicted operational taxonomical units as well as taxonomical compositions revealed no differences between communities in high CO2 mesocosms (~ 760 μatm) and those exposed to present-day CO2 conditions. Observed shifts in the planktonic community composition were mainly related to seasonal changes in temperature and nutrients. Furthermore, based on our investigations, the elevated CO2 did not affect the intraspecific diversity of the most common mesozooplankter, the calanoid copepod Pseudocalanus acuspes. Nevertheless, accompanying studies found temporary effects attributed to a raise in CO2. Differences in taxa composition between the CO2 treatments could, however, only be observed in a specific period of the experiment. Based on our genetic investigations, no compositional long-term shifts of the plankton communities exposed to elevated CO2 conditions were observed. Thus, we conclude that the compositions of planktonic communities, especially those in coastal areas, remain rather unaffected by increased CO2.

Competitive interactions between established grasses and woody plant seedlings under elevated CO₂ levels are mediated by soil water availability.

The expansion of woody plants into grasslands has been observed worldwide and is likely to have widespread ecological consequences. One proposal is that woody plant expansion into grasslands is driven in part by the rise in atmospheric CO2 concentrations. We have examined the effect of CO2 concentration on the competitive interactions between established C4 grasses and woody plant seedlings in a model grassland system. Woody plant seedlings were grown in mesocosms together with established C4 grasses in three competition treatments (root competition, shoot competition and root + shoot competition) under ambient and elevated CO2 levels. We found that the growth of the woody plant seedlings was suppressed by competition from grasses, with root and shoot competition having similar competitive effects on growth. In contrast to expectations, woody plant seedling growth was reduced at elevated CO2 levels compared to that at the ambient CO2 level across all competition treatments, with the most plausible explanation being reduced light and soil water availability in the elevated CO2 mesocosms. Reduced light and soil water availability in the elevated CO2 mesocosms was associated with an increased leaf area index of the grasses which offset the reductions in stomatal conductance and increased rainfall interception. The woody plant seedlings also had reduced 'escapability' (stem biomass and stem height) under elevated compared to ambient CO2 levels. Our results suggest that the expansion of woody plants into grasslands in the future will likely be context-dependent, with the establishment success of woody plant seedlings being strongly coupled to the CO2 response of competing grasses and to soil water availability.

Leaf area index drives soil water availability and extreme drought-related mortality under elevated CO2 in a temperate grassland model system.

The magnitude and frequency of climatic extremes, such as drought, are predicted to increase under future climate change conditions. However, little is known about how other factors such as CO2 concentration will modify plant community responses to these extreme climatic events, even though such modifications are highly likely. We asked whether the response of grasslands to repeat extreme drought events is modified by elevated CO2, and if so, what are the underlying mechanisms? We grew grassland mesocosms consisting of 10 co-occurring grass species common to the Cumberland Plain Woodland of western Sydney under ambient and elevated CO2 and subjected them to repeated extreme drought treatments. The 10 species included a mix of C3, C4, native and exotic species. We hypothesized that a reduction in the stomatal conductance of the grasses under elevated CO2 would be offset by increases in the leaf area index thus the retention of soil water and the consequent vulnerability of the grasses to extreme drought would not differ between the CO2 treatments. Our results did not support this hypothesis: soil water content was significantly lower in the mesocosms grown under elevated CO2 and extreme drought-related mortality of the grasses was greater. The C4 and native grasses had significantly higher leaf area index under elevated CO2 levels. This offset the reduction in the stomatal conductance of the exotic grasses as well as increased rainfall interception, resulting in reduced soil water content in the elevated CO2 mesocosms. Our results suggest that projected increases in net primary productivity globally of grasslands in a high CO2 world may be limited by reduced soil water availability in the future.

CO<sub>2</sub> increases <sup>14</sup>C primary production in an Arctic plankton community

Abstract. Responses to ocean acidification in plankton communities were studied during a CO2-enrichment experiment in the Arctic Ocean, accomplished from June to July 2010 in Kongsfjorden, Svalbard (78°56′ 2′′ N, 11°53′ 6′′ E). Enclosed in 9 mesocosms (volume: 43.9–47.6 m3), plankton was exposed to CO2 concentrations, ranging from glacial to projected mid-next-century levels. Fertilization with inorganic nutrients at day 13 of the experiment supported the accumulation of phytoplankton biomass, as indicated by two periods of high chl a concentration. This study tested for CO2 sensitivities in primary production (PP) of particulate organic carbon (PPPOC) and of dissolved organic carbon (PPDOC). Therefore, 14C-bottle incubations (24 h) of mesocosm samples were performed at 1 m depth receiving about 60% of incoming radiation. PP for all mesocosms averaged 8.06 ± 3.64 μmol C L−1 d−1 and was slightly higher than in the outside fjord system. Comparison between mesocosms revealed significantly higher PPPOC at elevated compared to low pCO2 after nutrient addition. PPDOC was significantly higher in CO2-enriched mesocosms before as well as after nutrient addition, suggesting that CO2 had a direct influence on DOC production. DOC concentrations inside the mesocosms increased before nutrient addition and more in high CO2 mesocosms. After addition of nutrients, however, further DOC accumulation was negligible and not significantly different between treatments, indicating rapid utilization of freshly produced DOC. Bacterial biomass production (BP) was coupled to PP in all treatments, indicating that 3.5 ± 1.9% of PP or 21.6 ± 12.5% of PPDOC provided on average sufficient carbon for synthesis of bacterial biomass. During the later course of the bloom, the response of 14C-based PP rates to CO2 enrichment differed from net community production (NCP) rates that were also determined during this mesocosm campaign. We conclude that the enhanced release of labile DOC during autotrophic production at high CO2 exceedingly stimulated activities of heterotrophic microorganisms. As a consequence, increased PP induced less NCP, as suggested earlier for carbon-limited microbial systems in the Arctic.

Effect of elevated CO<sub>2</sub> on the dynamics of particle-attached and free-living bacterioplankton communities in an Arctic fjord

Abstract. In the frame of the European Project on Ocean Acidification (EPOCA), the response of an Arctic pelagic community (<3 mm) to a gradient of seawater pCO2 was investigated. For this purpose 9 large-scale in situ mesocosms were deployed in Kongsfjorden, Svalbard (78°56.2´ N, 11°53.6´ E), in 2010. The present study investigates effects on the communities of particle-attached (PA; >3 μm) and free-living (FL; < 3 μm > 0.2 μm) bacteria by Automated Ribosomal Intergenic Spacer Analysis (ARISA) in 6 of the mesocosms, ranging from 185 to 1050 μatm initial pCO2, and the surrounding fjord. ARISA was able to resolve, on average, 27 bacterial band classes per sample and allowed for a detailed investigation of the explicit richness and diversity. Both, the PA and the FL bacterioplankton community exhibited a strong temporal development, which was driven mainly by temperature and phytoplankton development. In response to the breakdown of a picophytoplankton bloom, numbers of ARISA band classes in the PA community were reduced at low and medium CO2 (~ 185–685 μatm) by about 25%, while they were more or less stable at high CO2 (~ 820–1050 μatm). We hypothesise that enhanced viral lysis and enhanced availability of organic substrates at high CO2 resulted in a more diverse PA bacterial community in the post-bloom phase. Despite lower cell numbers and extracellular enzyme activities in the post-bloom phase, bacterial protein production was enhanced in high CO2 mesocosms, suggesting a positive effect of community richness on this function and on carbon cycling by bacteria.

Availability of phosphate for phytoplankton and bacteria and of glucose for bacteria at different <i>p</i>CO<sub>2</sub> levels in a mesocosm study

Abstract. Availability of phosphate for phytoplankton and bacteria and of glucose for bacteria at different pCO2 levels were studied in a mesocosm experiment (PeECE III). Using nutrient-depleted SW Norwegian fjord waters, three different levels of pCO2 (350 μatm: 1×CO2; 700 μatm: 2×CO2; 1050 μatm: 3×CO2) were set up, and nitrate and phosphate were added at the start of the experiment in order to induce a phytoplankton bloom. Despite similar responses of total particulate P concentration and phosphate turnover time at the three different pCO2 levels, the size distribution of particulate P and 33PO4 uptake suggested that phosphate transferred to the >10 μm fraction was greater in the 3×CO2 mesocosm during the first 6–10 days when phosphate concentration was high. During the period of phosphate depletion (after Day 12), specific phosphate affinity and specific alkaline phosphatase activity (APA) suggested a P-deficiency (i.e. suboptimal phosphate supply) rather than a P-limitation for the phytoplankton and bacterial community at the three different pCO2 levels. Specific phosphate affinity and specific APA tended to be higher in the 3×CO2 than in the 2×CO2 and 1×CO2 mesocosms during the phosphate depletion period, although no statistical differences were found. Glucose turnover time was correlated significantly and negatively with bacterial abundance and production but not with the bulk DOC concentration. This suggests that even though constituting a small fraction of the bulk DOC, glucose was an important component of labile DOC for bacteria. Specific glucose affinity of bacteria behaved similarly at the three different pCO2 levels with measured specific glucose affinities being consistently much lower than the theoretical maximum predicted from the diffusion-limited model. This suggests that bacterial growth was not severely limited by the glucose availability. Hence, it seems that the lower availability of inorganic nutrients after the phytoplankton bloom reduced the bacterial capacity to consume labile DOC in the upper mixed layer of the stratified mesocosms.

Build-up and decline of organic matter during PeECE III

Abstract. Increasing atmospheric carbon dioxide (CO2) concentrations due to anthropogenic fossil fuel combustion are currently changing the ocean's chemistry. Increasing oceanic [CO2] and consequently decreasing seawater pH have the potential to significantly impact marine life. Here we describe and analyze the build-up and decline of a natural phytoplankton bloom initiated during the 2005 mesocosm Pelagic Ecosystem CO2 Enrichment study (PeECE III). The draw-down of inorganic nutrients in the upper surface layer of the mesocosms was reflected by a concomitant increase of organic matter until day t11, the peak of the bloom. From then on, biomass standing stocks steadily decreased as more and more particulate organic matter was lost into the deeper layer of the mesocosms. We show that organic carbon export to the deeper layer was significantly enhanced at elevated CO2. This phenomenon might have impacted organic matter remineralization leading to decreased oxygen concentrations in the deeper layer of the high CO2 mesocosms as indicated by deep water ammonium concentrations. This would have important implications for our understanding of pelagic ecosystem functioning and future carbon cycling.

Effects of CO<sub>2</sub> on particle size distribution and phytoplankton abundance during a mesocosm bloom experiment (PeECE II)

Abstract. The influence of seawater carbon dioxide (CO2) concentration on the size distribution of suspended particles (2–60 μm) and on phytoplankton abundance was investigated during a mesocosm experiment at the large scale facility (LFS) in Bergen, Norway, in the frame of the Pelagic Ecosystem CO2 Enrichment study (PeECE II). In nine outdoor enclosures the partial pressure of CO2 in seawater was modified by an aeration system to simulate past (~190 μatm CO2), present (~370 μatm CO2) and future (~700 μatm CO2) CO2 conditions in triplicates. Due to the initial addition of inorganic nutrients, phytoplankton blooms developed in all mesocosms and were monitored over a period of 19 days. Seawater samples were collected daily for analysing the abundance of suspended particles and phytoplankton with the Coulter Counter and with Flow Cytometry, respectively. During the bloom period, the abundance of small particles (<4 μm) significantly increased at past, and decreased at future CO2 levels. At that time, a direct relationship between the total-surface-to-total-volume ratio of suspended particles and DIC concentration was determined for all mesocosms. Significant changes with respect to the CO2 treatment were also observed in the phytoplankton community structure. While some populations such as diatoms seemed to be insensitive to the CO2 treatment, others like Micromonas spp. increased with CO2, or showed maximum abundance at present day CO2 (i.e. Emiliania huxleyi). The strongest response to CO2 was observed in the abundance of small autotrophic nano-plankton that strongly increased during the bloom in the past CO2 mesocosms. Together, changes in particle size distribution and phytoplankton community indicate a complex interplay between the ability of the cells to physiologically respond to changes in CO2 and size selection. Size of cells is of general importance for a variety of processes in marine systems such as diffusion-limited uptake of substrates, resource allocation, predator-prey interaction, and gravitational settling. The observed changes in particle size distribution are therefore discussed with respect to biogeochemical cycling and ecosystem functioning.

Interactions among mycorrhizae, atmospheric CO2 and soil N impact plant community composition

AbstractWe examined plant community responses to interactions between arbuscular mycorrhizal (AM) fungi and availability of atmospheric CO2 and soil N. Communities of 14 plant species were grown in mesocosms containing living or killed AM fungal inoculum, ambient or elevated atmospheric CO2 and low or enriched soil N. After one growing season, significantly different plant communities existed in the different treatments. Plant species richness was lowest in +N mesocosms and highest in +AM + CO2 mesocosms. At ambient CO2, AM fungi reduced richness but at elevated CO2 they increased it. This was caused by changes in mortality rates of several C3 forbs and may suggest that CO2 enrichment ameliorates the carbon cost of some AM symbioses. Soil moisture was higher in +CO2 mesocosms but +AM counteracted this effect. These results suggest that AM symbioses may be important mediators of plant community responses to anthropogenic CO2 and N enrichment.