Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, w... more Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, western Baltic Sea. Diatoms were dominant competitors at high Si:N ratios (under N-limited conditions) and high Si:P ratios (under P-limited conditions) Under lo\v silicate supply, cyanobacteria became dominant at low N:P ratios and chlorophytes at h~g h ones. Changes in light intensity ~nfluenced the outcome of competition at the species level but not at the level of higher taxa. The nitrogen source (ammonium or nitrate) had no impact on the outcome of competition.
Seawater samples were collected biweekly from the northern Gulf of Aqaba, Red Sea, for Phytoplank... more Seawater samples were collected biweekly from the northern Gulf of Aqaba, Red Sea, for Phytoplankton analysis during the period May 1998 to October 1999. Microscopic counts and HPLC methods were employed. Procaryotic and eucaryotic ultraplankton dominated throughout most of the year, with larger nano-and microplankton making up only 5% of the photosynthetic biomass. Moderate seasonal variations in the 0-125 m integrated Chl a contrasted with a pronounced seasonal succession of the major taxonomic groups, reflecting the changes in the density stratification of the water column: Prochlorococcus dominated during the stratified summer period and were almost absent in winter. Chlorophyceae and Cryptophyceae were dominant during winter mixing but scarce or absent during summer. Diatoms and Synechococcus showed sharp and moderate biomass peaks in late winter and spring respectively, but remained at only low Chl a levels for the rest of the year. Chrysophyceae, Prymnesiophyceae and the scarce Dinophyceae showed no clear seasonal distribution pattern. The implications of alternating procaryotic and eucaryote dominated algal communities for the Red Sea pelagic food web are discussed.
The impacts of biochemicals driving food web processes are under investigation for just the last ... more The impacts of biochemicals driving food web processes are under investigation for just the last few decades. In addition, as jellyfish are drawing increasing attentions because of their mass developments and of their potential capacity of driving food web structures and energy flow by "top-down" and "bottom-up" controls. We here show that the provision with the biochemical complex thiamin (vitamin B 1) to the common phytoplankton Rhodomonas baltica altered its fatty acid (FA) pattern toward ω3-highly-unsaturated FAs (ω3-HUFA) and that this pattern was further transferred up to the zooplankton consumer, the copepod Acartia tonsa. However, polyps of the Jellyfish Aurelia aurita feeding on A. tonsa only had a low relative ω3-HUFA content, especially due to a reduction in 22:6ω3 (DHA), but elevated levels of 20:4ω6 (ARA). The high proportion of the ω-6 HUFA, ARA in polyps may provide evidence for preferential conversion of ARA in polyps, eventually from DHA in a so far unknown pathway. In contrast to A. tonsa, newly hatched A. salina nauplii used as food for A. aurita polyps were almost devoid of HUFA, but contained high levels of C 18 polyunsaturated FAs (C 18-PUFA). Consequently, polyps feeding on them contained few HUFA, while high levels of C 18-PUFA predominated. This suggests that A. aurita polyps cannot efficiently convert ω3 C 18-PUFA to ω3-HUFA. In addition, besides a decrease in saturated FAs, especially an increase in HUFA in A. aurita polyps with decreasing temperature was observed, for which the dietary provision with HUFA seemed to be critical. Altering the FA pattern as a response of temperature reflects an adaptation to seasonal changes and may be related to their life history plasticity.
Brownification reduces oxygen gross primary production and community respiration and changes the ... more Brownification reduces oxygen gross primary production and community respiration and changes the phytoplankton community composition: an in situ mesocosm experiment with high-frequency sensor measurements in a North Atlantic bay.
Aquatic ecosystems face a multitude of environmental stressors, including warming and acidificati... more Aquatic ecosystems face a multitude of environmental stressors, including warming and acidification. While warming is expected to have a pronounced effect on plankton communities, many components of the plankton seem fairly robust towards realistic end-of-century acidification conditions. However, interactions of the two stressors and the inclusion of further factors such as nutrient concentration and trophic interactions are expected to change this outcome. We investigated the effects of warming and high CO2 on a nutrient-deplete late summer plankton community from the Kiel Fjord, Baltic Sea, using a mesocosm setup crossing two temperatures with a gradient of CO2. Phytoplankton and microzooplankton (MZP) growth rates as well as biomass, taxonomic composition, and grazing rates of MZP were analysed. We observed effects of high CO2, warming, and their interactions on all measured parameters. The occurrence and direction of the effects were dependent on the phytoplankton or MZP commun...
Aquatic ecosystems face a multitude of environmental stressors, including warming and acidificati... more Aquatic ecosystems face a multitude of environmental stressors, including warming and acidification. While warming is expected to have a pronounced effect on plankton communities, many components of the plankton seem fairly robust towards realistic end-of-century acidification conditions. However, interactions of the two stressors and the inclusion of further factors such as nutrient concentration and trophic interactions are expected to change this outcome. We investigated the effects of warming and high CO2 on a nutrient-deplete late summer plankton community from the Kiel Fjord, Baltic Sea, using a mesocosm setup crossing two temperatures with a gradient of CO2. Phytoplankton and microzooplankton (MZP) growth rates as well as biomass, taxonomic composition, and grazing rates of MZP were analysed. We observed effects of high CO2, warming, and their interactions on all measured parameters. The occurrence and direction of the effects were dependent on the phytoplankton or MZP commun...
'greenhouse' world. Instead, many studies have addressed the biological effects of either ocean a... more 'greenhouse' world. Instead, many studies have addressed the biological effects of either ocean acidification or warming in particular on phytoplankton species composition and biomass. These studies indicate that CO 2 can act as a stressor, in particular for calcifiers, but in cases where it is a limiting resource,
Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are ex... more Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are con [...]
Bacterial and archaeal diversity and succession were studied during a mesocosm experiment that in... more Bacterial and archaeal diversity and succession were studied during a mesocosm experiment that investigated whether changing light regimes could affect the onset of phytoplankton blooms. For this, 454-pyrosequencing of the bacterial V1-V3 and archaeal V3-V9 16S rRNA regions was performed in samples collected from four mesocosms receiving different light irradiances at the beginning and the end of the experiment and during phytoplankton growth. In total, 46 bacterial operational taxonomic units (OTUs) with ≥1% relative abundance occurred (22–34 OTUs per mesocosm). OTUs were affiliated mainly with Rhodobacteraceae, Flavobacteriaceae and Alteromonadaceae. The four mesocosms shared 11 abundant OTUs. Dominance increased at the beginning of phytoplankton growth in all treatments and decreased thereafter. Maximum dominance was found in the mesocosms with high irradiances. Overall, specific bacterial OTUs had different responses in terms of relative abundance under in situ and high light in...
, and the Publisher, the British Institute of Radiology, most gratefully acknowledge the contribu... more , and the Publisher, the British Institute of Radiology, most gratefully acknowledge the contribution of the following colleagues who have acted unstintingly as referees.
In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically d... more In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment ''bottleneck'' at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (Ͻ4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size.
While the isolated responses of marine phytoplankton to climate warming and to ocean acidificatio... more While the isolated responses of marine phytoplankton to climate warming and to ocean acidification have been studied intensively, studies on the combined effect of both aspects of Global Change are still scarce. Therefore, we performed a mesocosm experiment with a factorial combination of temperature (9 and 15°C) and pCO 2 (means: 439 ppm and 1040 ppm) with a natural autumn plankton community from the western Baltic Sea. Temporal trajectories of total biomass and of the biomass of the most important higher taxa followed similar patterns in all treatments. When averaging over the entire time course, phytoplankton biomass decreased with warming and increased with CO 2 under warm conditions. The contribution of the two dominant higher phytoplankton taxa (diatoms and cryptophytes) and of the 4 most important species (3 diatoms, 1 cryptophyte) did not respond to the experimental treatments. Taxonomic composition of phytoplankton showed only responses at the level of subdominant and rare species. Phytoplankton cell sizes increased with CO 2 addition and decreased with warming. Both effects were stronger for larger species. Warming effects were stronger than CO 2 effects and tended to counteract each other. Phytoplankton communities without calcifying species and exposed to short-term variation of CO 2 seem to be rather resistant to ocean acidification.
Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, w... more Nutrient competition experiments were performed with periphytic microalgae from the Kiel Fjord, western Baltic Sea. Diatoms were dominant competitors at high Si:N ratios (under N-limited conditions) and high Si:P ratios (under P-limited conditions) Under lo\v silicate supply, cyanobacteria became dominant at low N:P ratios and chlorophytes at h~g h ones. Changes in light intensity ~nfluenced the outcome of competition at the species level but not at the level of higher taxa. The nitrogen source (ammonium or nitrate) had no impact on the outcome of competition.
Seawater samples were collected biweekly from the northern Gulf of Aqaba, Red Sea, for Phytoplank... more Seawater samples were collected biweekly from the northern Gulf of Aqaba, Red Sea, for Phytoplankton analysis during the period May 1998 to October 1999. Microscopic counts and HPLC methods were employed. Procaryotic and eucaryotic ultraplankton dominated throughout most of the year, with larger nano-and microplankton making up only 5% of the photosynthetic biomass. Moderate seasonal variations in the 0-125 m integrated Chl a contrasted with a pronounced seasonal succession of the major taxonomic groups, reflecting the changes in the density stratification of the water column: Prochlorococcus dominated during the stratified summer period and were almost absent in winter. Chlorophyceae and Cryptophyceae were dominant during winter mixing but scarce or absent during summer. Diatoms and Synechococcus showed sharp and moderate biomass peaks in late winter and spring respectively, but remained at only low Chl a levels for the rest of the year. Chrysophyceae, Prymnesiophyceae and the scarce Dinophyceae showed no clear seasonal distribution pattern. The implications of alternating procaryotic and eucaryote dominated algal communities for the Red Sea pelagic food web are discussed.
The impacts of biochemicals driving food web processes are under investigation for just the last ... more The impacts of biochemicals driving food web processes are under investigation for just the last few decades. In addition, as jellyfish are drawing increasing attentions because of their mass developments and of their potential capacity of driving food web structures and energy flow by "top-down" and "bottom-up" controls. We here show that the provision with the biochemical complex thiamin (vitamin B 1) to the common phytoplankton Rhodomonas baltica altered its fatty acid (FA) pattern toward ω3-highly-unsaturated FAs (ω3-HUFA) and that this pattern was further transferred up to the zooplankton consumer, the copepod Acartia tonsa. However, polyps of the Jellyfish Aurelia aurita feeding on A. tonsa only had a low relative ω3-HUFA content, especially due to a reduction in 22:6ω3 (DHA), but elevated levels of 20:4ω6 (ARA). The high proportion of the ω-6 HUFA, ARA in polyps may provide evidence for preferential conversion of ARA in polyps, eventually from DHA in a so far unknown pathway. In contrast to A. tonsa, newly hatched A. salina nauplii used as food for A. aurita polyps were almost devoid of HUFA, but contained high levels of C 18 polyunsaturated FAs (C 18-PUFA). Consequently, polyps feeding on them contained few HUFA, while high levels of C 18-PUFA predominated. This suggests that A. aurita polyps cannot efficiently convert ω3 C 18-PUFA to ω3-HUFA. In addition, besides a decrease in saturated FAs, especially an increase in HUFA in A. aurita polyps with decreasing temperature was observed, for which the dietary provision with HUFA seemed to be critical. Altering the FA pattern as a response of temperature reflects an adaptation to seasonal changes and may be related to their life history plasticity.
Brownification reduces oxygen gross primary production and community respiration and changes the ... more Brownification reduces oxygen gross primary production and community respiration and changes the phytoplankton community composition: an in situ mesocosm experiment with high-frequency sensor measurements in a North Atlantic bay.
Aquatic ecosystems face a multitude of environmental stressors, including warming and acidificati... more Aquatic ecosystems face a multitude of environmental stressors, including warming and acidification. While warming is expected to have a pronounced effect on plankton communities, many components of the plankton seem fairly robust towards realistic end-of-century acidification conditions. However, interactions of the two stressors and the inclusion of further factors such as nutrient concentration and trophic interactions are expected to change this outcome. We investigated the effects of warming and high CO2 on a nutrient-deplete late summer plankton community from the Kiel Fjord, Baltic Sea, using a mesocosm setup crossing two temperatures with a gradient of CO2. Phytoplankton and microzooplankton (MZP) growth rates as well as biomass, taxonomic composition, and grazing rates of MZP were analysed. We observed effects of high CO2, warming, and their interactions on all measured parameters. The occurrence and direction of the effects were dependent on the phytoplankton or MZP commun...
Aquatic ecosystems face a multitude of environmental stressors, including warming and acidificati... more Aquatic ecosystems face a multitude of environmental stressors, including warming and acidification. While warming is expected to have a pronounced effect on plankton communities, many components of the plankton seem fairly robust towards realistic end-of-century acidification conditions. However, interactions of the two stressors and the inclusion of further factors such as nutrient concentration and trophic interactions are expected to change this outcome. We investigated the effects of warming and high CO2 on a nutrient-deplete late summer plankton community from the Kiel Fjord, Baltic Sea, using a mesocosm setup crossing two temperatures with a gradient of CO2. Phytoplankton and microzooplankton (MZP) growth rates as well as biomass, taxonomic composition, and grazing rates of MZP were analysed. We observed effects of high CO2, warming, and their interactions on all measured parameters. The occurrence and direction of the effects were dependent on the phytoplankton or MZP commun...
'greenhouse' world. Instead, many studies have addressed the biological effects of either ocean a... more 'greenhouse' world. Instead, many studies have addressed the biological effects of either ocean acidification or warming in particular on phytoplankton species composition and biomass. These studies indicate that CO 2 can act as a stressor, in particular for calcifiers, but in cases where it is a limiting resource,
Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are ex... more Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N:P Redfield ratio of 16:1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N:P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N:P supply ratio, although PON:POP of accumulated biomass generally exceeded the supply ratio. Microbial N:P composition was constrained by a general lower limit of 5:1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N:P supply. Low nutrient N:P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are con [...]
Bacterial and archaeal diversity and succession were studied during a mesocosm experiment that in... more Bacterial and archaeal diversity and succession were studied during a mesocosm experiment that investigated whether changing light regimes could affect the onset of phytoplankton blooms. For this, 454-pyrosequencing of the bacterial V1-V3 and archaeal V3-V9 16S rRNA regions was performed in samples collected from four mesocosms receiving different light irradiances at the beginning and the end of the experiment and during phytoplankton growth. In total, 46 bacterial operational taxonomic units (OTUs) with ≥1% relative abundance occurred (22–34 OTUs per mesocosm). OTUs were affiliated mainly with Rhodobacteraceae, Flavobacteriaceae and Alteromonadaceae. The four mesocosms shared 11 abundant OTUs. Dominance increased at the beginning of phytoplankton growth in all treatments and decreased thereafter. Maximum dominance was found in the mesocosms with high irradiances. Overall, specific bacterial OTUs had different responses in terms of relative abundance under in situ and high light in...
, and the Publisher, the British Institute of Radiology, most gratefully acknowledge the contribu... more , and the Publisher, the British Institute of Radiology, most gratefully acknowledge the contribution of the following colleagues who have acted unstintingly as referees.
In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically d... more In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment ''bottleneck'' at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (Ͻ4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size.
While the isolated responses of marine phytoplankton to climate warming and to ocean acidificatio... more While the isolated responses of marine phytoplankton to climate warming and to ocean acidification have been studied intensively, studies on the combined effect of both aspects of Global Change are still scarce. Therefore, we performed a mesocosm experiment with a factorial combination of temperature (9 and 15°C) and pCO 2 (means: 439 ppm and 1040 ppm) with a natural autumn plankton community from the western Baltic Sea. Temporal trajectories of total biomass and of the biomass of the most important higher taxa followed similar patterns in all treatments. When averaging over the entire time course, phytoplankton biomass decreased with warming and increased with CO 2 under warm conditions. The contribution of the two dominant higher phytoplankton taxa (diatoms and cryptophytes) and of the 4 most important species (3 diatoms, 1 cryptophyte) did not respond to the experimental treatments. Taxonomic composition of phytoplankton showed only responses at the level of subdominant and rare species. Phytoplankton cell sizes increased with CO 2 addition and decreased with warming. Both effects were stronger for larger species. Warming effects were stronger than CO 2 effects and tended to counteract each other. Phytoplankton communities without calcifying species and exposed to short-term variation of CO 2 seem to be rather resistant to ocean acidification.
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Papers by Ulrich Sommer