ALBERT

Description: Dissolved organic carbon (DOC) concentrations – mainly of terrestrial origin – are increasing worldwide in inland waters. The biodegradability of the DOC varies depending on quantity and chemical quality. Heterotrophic bacteria are the main consumers of DOC and thus determine DOC temporal dynamics and availability for higher trophic levels. It is therefore crucial to understand the processes controlling the bacterial turnover of additional allochthonous and autochthonous DOC in aquatic systems. Our aim was to study bacterial carbon (C) turnover with respect to DOC quantity and chemical quality using both allochthonous and autochthonous DOC sources. We incubated a natural bacterial community with allochthonous C (13C-labeled beech leachate) and increased concentrations and pulses (intermittent occurrence of organic matter input) of autochthonous C (algae lysate). We then determined bacterial carbon consumption, activities, and community composition together with the carbon flow through bacteria using stable C isotopes. The chemical analysis of single sources revealed differences in aromaticity and fractions of low and high molecular weight substances (LMWS and HMWS, respectively) between allochthonous and autochthonous C sources. In parallel to these differences in chemical composition, we observed a higher availability of allochthonous C as evidenced by increased DOC consumption and bacterial growth efficiencies (BGE) when solely allochthonous C was provided. In treatments with mixed sources, rising concentrations of added autochthonous DOC resulted in a further, significant increase in bacterial DOC consumption from 52 to 68% when nutrients were not limiting. This rise was accompanied by a decrease in the humic substances (HS) fraction and an increase in bacterial biomass. Stable C isotope analyses of phospholipid fatty acids (PLFA) and respired dissolved inorganic carbon (DIC) supported a preferential assimilation of autochthonous C and respiration of the allochthonous C. Changes in DOC concentration and consumption in mixed treatments did not affect bacterial community composition (BCC), but BCC differed in single vs. mixed incubations. Our study highlights that DOC quantity affects bacterial C consumption but not BCC in nutrient-rich aquatic systems. BCC shifted when a mixture of allochthonous and autochthonous C was provided simultaneously to the bacterial community. Our results indicate that chemical quality rather than source of DOC per se (allochthonous vs. autochthonous) determines bacterial DOC turnover.

Description: Heterocystous cyanobacteria of the genus Nodularia form extensive blooms in the Baltic Sea contributing substantially to the total annual primary production. Moreover, they dispense a large fraction of new nitrogen to the ecosystem, when inorganic nitrogen concentration in summer is low. Thus, it is of great ecological importance to know how Nodularia will react to future environmental changes, in particular to increasing carbon dioxide (CO2) concentrations and what consequences there might arise for cycling of organic matter in the ocean. Here, we determined carbon (C) and dinitrogen (N2) fixation rates, growth, elemental stoichiometry of particulate organic matter and nitrogen turnover during batch growth of the heterocystous cyanobacterium Nodularia spumigena under glacial (180 ppm), present (380 ppm), and future (780 ppm) CO2 concentrations. Our results demonstrate an overall stimulating effect of rising pCO2 on C and N2 fixation, as well as on cell growth. An increase in pCO2 resulted in an elevation in growth rate, C and N2 fixation by 23%, 36% and 25%, respectively (180 ppm vs. 380 ppm) and by 27%, 2% and 4%, respectively (380 ppm vs. 780 ppm). Additionally, elevation in the carbon and nitrogen to phosphorus quota of the particulate biomass formed (POC:POP and PON:POP) was observed at high pCO2. Our findings suggest that rising pCO2 stimulates the growth of heterocystous diazotrophic cyanobacteria, in a similar way as reported for non-heterocystous diazotrophs. Implications for biogeochemical cycling and food web dynamics, as well as ecological and socio-economical aspects in the Baltic Sea are discussed.

Description: Studies investigating the effect of increasing CO2 levels on the phosphorus cycle in natural waters are lacking although phosphorus often controls phytoplankton development in aquatic systems. The aim of our study was to analyze effects of elevated CO2 levels on phosphorus pool sizes and uptake. Therefore, we conducted a CO2-manipulation mesocosm experiment in the Storfjärden (western Gulf of Finland, Baltic Sea) in summer 2012. We compared the phosphorus dynamics in different mesocosm treatments but also studied them outside the mesocosms in the surrounding fjord water. In the mesocosms as well as in surface waters of Storfjärden, dissolved organic phosphorus (DOP) concentrations of 0.26 ± 0.03 and 0.23 ± 0.04 μmol L−1, respectively, formed the main fraction of the total P-pool (TP), whereas phosphate (PO4) constituted the lowest fraction with mean concentration of 0.15 ± 0.02 μmol L−1 and 0.17 ± 0.07 μmol L−1 in the mesocosms and in the fjord, respectively. Uptake of PO4 ranged between 0.6 and 3.9 nmol L−1 h−1 of which ~ 86 % (mesocosms) and ~ 72 % (fjord) were realized by the size fraction 〈 3 μm. Adenosine triphosphate (ATP) uptake revealed that additional P was supplied from organic compounds accounting for 25–27 % of P provided by PO4 only. CO2 additions did not cause significant changes in phosphorus (P) pool sizes, DOP composition, and uptake of PO4 and ATP when the whole study period was taken into account. About 18 % of PO4 was transformed into POP, whereby the major proportion (~ 82 %) was converted into DOP suggesting that the conversion of PO4 to DOP is the main pathway of the PO4 turnover. We observed that significant relationships (e.g., between POP and Chl a) in the untreated mesocosms vanished under increased fCO2 conditions. Consequently, it can be hypothesized that the relationship between POP formation and phytoplankton growth changed under elevated CO2 conditions. Significant short-term effects were observed for PO4 and particulate organic phosphorus (POP) pool sizes in CO2 treatments 〉 1000 μatm during periods when phytoplankton started to grow.

Description: Heterocystous cyanobacteria of the genus Nodularia form extensive blooms in the Baltic Sea and contribute substantially to the total annual primary production. Moreover, they dispense a large fraction of new nitrogen to the ecosystem when inorganic nitrogen concentration in summer is low. Thus, it is of ecological importance to know how Nodularia will react to future environmental changes, in particular to increasing carbon dioxide (CO2) concentrations and what consequences there might arise for cycling of organic matter in the Baltic Sea. Here, we determined carbon (C) and dinitrogen (N2) fixation rates, growth, elemental stoichiometry of particulate organic matter and nitrogen turnover in batch cultures of the heterocystous cyanobacterium Nodularia spumigena under low (median 315 μatm), mid (median 353 μatm), and high (median 548 μatm) CO2 concentrations. Our results demonstrate an overall stimulating effect of rising pCO2 on C and N2 fixation, as well as on cell growth. An increase in pCO2 during incubation days 0 to 9 resulted in an elevation in growth rate by 84 ± 38% (low vs. high pCO2) and 40 ± 25% (mid vs. high pCO2), as well as in N2 fixation by 93 ± 35% and 38 ± 1%, respectively. C uptake rates showed high standard deviations within treatments and in between sampling days. Nevertheless, C fixation in the high pCO2 treatment was elevated compared to the other two treatments by 97% (high vs. low) and 44% (high vs. mid) at day 0 and day 3, but this effect diminished afterwards. Additionally, elevation in carbon to nitrogen and nitrogen to phosphorus ratios of the particulate biomass formed (POC : POP and PON : POP) was observed at high pCO2. Our findings suggest that rising pCO2 stimulates the growth of heterocystous diazotrophic cyanobacteria, in a similar way as reported for the non-heterocystous diazotroph Trichodesmium. Implications for biogeochemical cycling and food web dynamics, as well as ecological and socio-economical aspects in the Baltic Sea are discussed.

Description: Dissolved organic carbon (DOC) concentrations – mainly of terrestrial origin – are increasing worldwide in inland waters. Heterotrophic bacteria are the main consumers of DOC and thus determine DOC temporal dynamics and availability for higher trophic levels. Our aim was to study bacterial carbon (C) turnover with respect to DOC quantity and chemical quality using both allochthonous and autochthonous DOC sources. We incubated a natural bacterial community with allochthonous C (13C-labeled beech leachate) and increased concentrations and pulses (intermittent occurrence of organic matter input) of autochthonous C (phytoplankton lysate). We then determined bacterial C consumption, activities, and community composition together with the C flow through bacteria using stable C isotopes. The chemical analysis of single sources revealed differences in aromaticity and low- and high-molecular-weight substance fractions (LMWS and HMWS, respectively) between allochthonous and autochthonous C sources. Both DOC sources (allochthonous and autochthonous DOC) were metabolized at a high bacterial growth efficiency (BGE) around 50%. In treatments with mixed sources, rising concentrations of added autochthonous DOC resulted in a further, significant increase in bacterial DOC consumption of up to 68% when nutrients were not limiting. This rise was accompanied by a decrease in the humic substance (HS) fraction and an increase in bacterial biomass. Changes in DOC concentration and consumption in mixed treatments did not affect bacterial community composition (BCC), but BCC differed in single vs. mixed incubations. Our study highlights that DOC quantity affects bacterial C consumption but not BCC in nutrient-rich aquatic systems. BCC shifted when a mixture of allochthonous and autochthonous C was provided simultaneously to the bacterial community. Our results indicate that chemical quality rather than source of DOC per se (allochthonous vs. autochthonous) determines bacterial DOC turnover.

Description: The predicted rise in anthropogenic CO2 emissions will increase CO2 concentrations and decrease seawater pH in the upper ocean. Recent studies have revealed effects of pCO2 induced changes in seawater chemistry on a variety of marine life forms, in particular calcifying organisms. To test whether the predicted increase in pCO2 will directly or indirectly (via changes in phytoplankton dynamics) affect abundance, activities, and community composition of heterotrophic bacteria during phytoplankton bloom development, we have aerated mesocosms with CO2 to obtain triplicates with three different partial pressures of CO2 (pCO2): 350 μatm (1×CO2), 700 μatm (2×CO2) and 1050 μatm (3×CO2). The development of a phytoplankton bloom was initiated by the addition of nitrate and phosphate. In accordance to an elevated carbon to nitrogen drawdown at increasing pCO2, bacterial production (BPP) of free-living and attached bacteria as well as cell-specific BPP (csBPP) of attached bacteria were related to the C:N ratio of suspended matter. These relationships significantly differed among treatments. However, bacterial abundance and activities were not statistically different among treatments. Solely community structure of free-living bacteria changed with pCO2 whereas that of attached bacteria seemed to be independent of pCO2 but tightly coupled to phytoplankton bloom development. Our findings imply that changes in pCO2, although reflected by changes in community structure of free-living bacteria, do not directly affect bacterial activity. Furthermore, bacterial activity and dynamics of heterotrophic bacteria, especially of attached bacteria, were tightly correlated to phytoplankton development and, hence, may also potentially depend on changes in pCO2.

Description: Using 15N and 33P, we measured the turnover of organic and inorganic nitrogen (N) and phosphorus (P) substrates, and the partitioning of N and P from these sources into two size fractions of marine osmotrophs during the course of a phytoplankton bloom in a nutrient manipulated mesocosm. The larger size fraction (〉0.8 μm), mainly consisting of the coccolithophorid Emiliania huxleyi, but also including an increasing amount of large particle-associated bacteria as the bloom proceeded, dominated uptake of the inorganic forms NH4+, NO3−, and PO43−. The uptake of N from leucine, and P from ATP and dissolved DNA (dDNA), was initially dominated by the 0.8–0.2 μm size fraction, but shifted towards dominance by the 〉0.8 μm size fraction as the system turned to an increasing degree of N-deficiency. Normalizing uptake to biomass of phytoplankton and heterotrophic bacteria revealed that organisms in the 0.8–0.2 μm size fraction had higher specific affinity for leucine-N than those in the 〉0.8 μm size fraction when N was deficient, whereas the opposite was the case for NH4+. There was no such difference regarding the specific affinity for P substrates. Since heterotrophic bacteria seem to acquire N from organic compounds like leucine more efficiently than phytoplankton, our results suggest different structuring of the microbial food chain in N-limited relative to P-limited environments.

Description: The predicted rise in anthropogenic CO2 emissions will increase CO2 concentrations and decrease seawater pH in the upper ocean. Recent studies have revealed effects of pCO2 induced changes in seawater chemistry on a variety of marine life forms, in particular calcifying organisms. To test whether the predicted increase in pCO2 will directly or indirectly (via changes in phytoplankton dynamics) affect abundance, activities, and community composition of heterotrophic bacteria during phytoplankton bloom development, we have aerated mesocosms with CO2 to obtain triplicates with three different partial pressures of CO2 (pCO2): 350 µatm (1×CO2), 700 µatm (2×CO2) and 1050 µatm (3×CO2). The development of a phytoplankton bloom was initiated by the addition of nitrate and phosphate. In accordance to an elevated carbon to nitrogen drawdown at increasing pCO2, bacterial production (BPP) of free-living and attached bacteria as well as cell-specific BPP (csBPP) of attached bacteria were related to the C:N ratio of suspended matter. These relationships significantly differed among treatments. However, bacterial abundance and activities were not statistically different among treatments. Solely community structure of free-living bacteria changed with pCO2 whereas that of attached bacteria seemed to be independent of pCO2 but tightly coupled to phytoplankton bloom development. Our findings imply that changes in pCO2, although reflected by changes in community structure of free-living bacteria, do not directly affect bacterial activity. Furthermore, bacterial activity and dynamics of heterotrophic bacteria, especially of attached bacteria, were tightly linked to phytoplankton development and, hence, may also potentially depend on changes in pCO2.

Description: Availability of phosphate for phytoplankton and bacteria and of labile organic carbon 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; 750 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) but not a P-limitation for the phytoplankton and bacterial community at the three different pCO2 levels. Although specific phosphate affinity and specific APA tended to be higher in 3×CO2 than in 2×CO2 and 1×CO2 mesocosms during the phosphate depletion period, no statistical differences were found. Responses of specific glucose affinity for bacteria were similar at the three different pCO2 levels. Measured specific glucose affinities were consistently much lower than the theoretical maximum predicted from the diffusion-limited model, suggesting that bacterial growth was not limited by the availability of labile dissolved organic carbon. These results suggest that availability of phosphate and glucose was similar at the three different pCO2 levels.