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Oceanic and coastal waters are acidifying due to processes dominated in the open ocean by increasing atmospheric CO2 and dominated in estuaries and some coastal waters by nutrient-fueled respiration. The patterns and severity of acidification, as well as its effects, are modified by the host of stressors related to human activities that also influence these habitats. Temperature, deoxygenation, and changes in food webs are particularly important co-stressors because they are pervasive, and both their causes and effects are often mechanistically linked to acidification. Development of a theoretical underpinning to multiple stressor research that considers physiological, ecological, and evolutionary perspectives is needed because testing all combinations of stressors and stressor intensities experimentally is impossible. Nevertheless, use of a wide variety of research approaches is a logical and promising strategy for improving understanding of acidification and its effects. Future research that focuses on spatial and temporal patterns of stressor interactions and on identifying mechanisms by which multiple stressors affect individuals, populations, and ecosystems is critical. It is also necessary to incorporate consideration of multiple stressors into management, mitigation, and adaptation to acidification and to increase public and policy recognition of the importance of addressing acidification in the context of the suite of other stressors with which it potentially interacts.

Oceanic and coastal waters are acidifying due to processes dominated in the open ocean by increasing atmospheric CO2 and dominated in estuaries and some coastal waters by nutrient-fueled respiration. The patterns and severity of acidification, as well as its effects, are modified by the host of stressors related to human activities that also influence these habitats. Temperature, deoxygenation, and changes in food webs are particularly important co-stressors because they are pervasive, and both their causes and effects are often mechanistically linked to acidification. Development of a theoretical underpinning to multiple stressor research that considers physiological, ecological, and evolutionary perspectives is needed because testing all combinations of stressors and stressor intensities experimentally is impossible. Nevertheless, use of a wide variety of research approaches is a logical and promising strategy for improving understanding of acidification and its effects. Future research that focuses on spatial and temporal patterns of stressor interactions and on identifying mechanisms by which multiple stressors affect individuals, populations, and ecosystems is critical. It is also necessary to incorporate consideration of multiple stressors into management, mitigation, and adaptation to acidification and to increase public and policy recognition of the importance of addressing acidification in the context of the suite of other stressors with which it potentially interacts.

Carbon uptake and partitioning of two globally abundant diatom species, Thalassiosira weissflogii and Dactyliosolen fragilissimus, was investigated in batch culture experiments under four conditions: ambient (15 degrees C, 400 atm), high CO2 (15 degrees C, 1000 atm), high temperature (20 degrees C, 400 atm), and combined (20 degrees C, 1000 atm). The experiments were run from exponential growth into the stationary phase (six days after nitrogen depletion), allowing us to track biogeochemical dynamics analogous to bloom situations in the ocean. Elevated CO2 had a fertilizing effect and enhanced uptake of dissolved inorganic carbon (DIC) by about 8% for T. weissflogii and by up to 39% for D. fragilissimus. This was also reflected in higher cell numbers, build-up of particulate and dissolved organic matter, and transparent exopolymer particles. The CO2 effects were most prominent in the stationary phase when nitrogen was depleted and CO2(aq) concentrations were low. This indicates that diatoms in the high CO2 treatments could take up more DIC until CO2 concentrations in seawater became so low that carbon limitation occurs. These results suggest that, contrary to common assumptions, diatoms could be highly sensitive to ongoing changes in oceanic carbonate chemistry, particularly under nutrient limitation. Warming from 15 to 20 degrees C had a stimulating effect on one species but acted as a stressor on the other species, highlighting the importance of species-specific physiological optima and temperature ranges in the response to ocean warming. Overall, these sensitivities to CO2 and temperature could have profound impacts on diatoms blooms and the biological pump.

Carbon uptake and partitioning of two globally abundant diatom species, Thalassiosira weissflogii and Dactyliosolen fragilissimus, was investigated in batch culture experiments under four conditions: ambient (15 degrees C, 400 atm), high CO2 (15 degrees C, 1000 atm), high temperature (20 degrees C, 400 atm), and combined (20 degrees C, 1000 atm). The experiments were run from exponential growth into the stationary phase (six days after nitrogen depletion), allowing us to track biogeochemical dynamics analogous to bloom situations in the ocean. Elevated CO2 had a fertilizing effect and enhanced uptake of dissolved inorganic carbon (DIC) by about 8% for T. weissflogii and by up to 39% for D. fragilissimus. This was also reflected in higher cell numbers, build-up of particulate and dissolved organic matter, and transparent exopolymer particles. The CO2 effects were most prominent in the stationary phase when nitrogen was depleted and CO2(aq) concentrations were low. This indicates that diatoms in the high CO2 treatments could take up more DIC until CO2 concentrations in seawater became so low that carbon limitation occurs. These results suggest that, contrary to common assumptions, diatoms could be highly sensitive to ongoing changes in oceanic carbonate chemistry, particularly under nutrient limitation. Warming from 15 to 20 degrees C had a stimulating effect on one species but acted as a stressor on the other species, highlighting the importance of species-specific physiological optima and temperature ranges in the response to ocean warming. Overall, these sensitivities to CO2 and temperature could have profound impacts on diatoms blooms and the biological pump.

Carbon uptake and partitioning of two globally abundant diatom species, Thalassiosira weissflogii and Dactyliosolen fragilissimus, was investigated in batch culture experiments under four conditions: ambient (15 degrees C, 400 atm), high CO2 (15 degrees C, 1000 atm), high temperature (20 degrees C, 400 atm), and combined (20 degrees C, 1000 atm). The experiments were run from exponential growth into the stationary phase (six days after nitrogen depletion), allowing us to track biogeochemical dynamics analogous to bloom situations in the ocean. Elevated CO2 had a fertilizing effect and enhanced uptake of dissolved inorganic carbon (DIC) by about 8% for T. weissflogii and by up to 39% for D. fragilissimus. This was also reflected in higher cell numbers, build-up of particulate and dissolved organic matter, and transparent exopolymer particles. The CO2 effects were most prominent in the stationary phase when nitrogen was depleted and CO2(aq) concentrations were low. This indicates that diatoms in the high CO2 treatments could take up more DIC until CO2 concentrations in seawater became so low that carbon limitation occurs. These results suggest that, contrary to common assumptions, diatoms could be highly sensitive to ongoing changes in oceanic carbonate chemistry, particularly under nutrient limitation. Warming from 15 to 20 degrees C had a stimulating effect on one species but acted as a stressor on the other species, highlighting the importance of species-specific physiological optima and temperature ranges in the response to ocean warming. Overall, these sensitivities to CO2 and temperature could have profound impacts on diatoms blooms and the biological pump.

Transparent exopolymer particles (TEP) are the most ubiquitous gel particles in the ocean and form abiotically from dissolved precursors. Although these particles can accumulate at the ocean surface, being thus exposed to intense sunlight, the role of solar radiation for the assembly and degradation of TEP is unknown. In this study, we experimentally determined the effects of visible and ultraviolet B (UVB) radiation on (1) TEP degradation (photolysis experiments), (2) TEP assembly from dissolved polymers (photoinhibition experiments) and (3) TEP release by microorganisms. Solar radiation, particularly in the UVB range, caused significant TEP photolysis, with loss rates from 27 to 34% per day. Dissolved polysaccharides did not increase in parallel. No TEP were formed under UVB, visible or dark conditions, indicating that light does not promote TEP assembly. UVB radiation enhanced TEP release by microorganisms, possibly due to cell deaths, or as a protective measure. Increases in UVB may lead to enhanced TEP photolysis in the ocean, with further consequences for TEP dynamics and, ultimately, sea-air gas exchange.

The late stage of the North East Atlantic (NEA) spring bloom was investigated during June 2005 along a transect section from 45 to 66° N between 15 and 20° W in order to characterize the contribution of siliceous and calcareous phytoplankton groups and describe their distribution in relation to environmental factors. We measured several biogeochemical parameters such as nutrients, surface trace metals, algal pigments, biogenic silica (BSi), particulate inorganic carbon (PIC) or calcium carbonate, particulate organic carbon, nitrogen and phosphorus (POC, PON and POP, respectively), as well as transparent exopolymer particles (TEP). Results were compared with other studies undertaken in this area since the JGOFS NABE program. Characteristics of the spring bloom generally agreed well with the accepted scenario for the development of the autotrophic community. The NEA seasonal diatom bloom was in the late stages when we sampled the area and diatoms were constrained to the northern part of our transect, over the Icelandic Basin (IB) and Icelandic Shelf (IS). Coccolithophores dominated the phytoplankton community, with a large distribution over the Rockall-Hatton Plateau (RHP) and IB. The Porcupine Abyssal Plain (PAP) region at the southern end of our transect was the region with the lowest biomass, as demonstrated by very low Chla concentrations and a community dominated by picophytoplankton. Early depletion of dissolved silicic acid (DSi) and increased stratification of the surface layer most likely triggered the end of the diatom bloom, leading to coccolithophore dominance. The chronic Si deficiency observed in the NEA could be linked to moderate Fe limitation, which increases the efficiency of the Si pump. TEP closely mirrored the distribution of both biogenic silica at depth and prymnesiophytes in the surface layer suggesting the sedimentation of the diatom bloom in the form of aggregates, but the relative contribution of diatoms and coccolithophores to carbon export

The late stage of the North East Atlantic (NEA) spring bloom was investigated during June 2005 along a transect section from 45 to 66° N between 15 and 20° W in order to characterize the contribution of siliceous and calcareous phytoplankton groups and describe their distribution in relation to environmental factors. We measured several biogeochemical parameters such as nutrients, surface trace metals, algal pigments, biogenic silica (BSi), particulate inorganic carbon (PIC) or calcium carbonate, particulate organic carbon, nitrogen and phosphorus (POC, PON and POP, respectively), as well as transparent exopolymer particles (TEP). Results were compared with other studies undertaken in this area since the JGOFS NABE program. Characteristics of the spring bloom generally agreed well with the accepted scenario for the development of the autotrophic community. The NEA seasonal diatom bloom was in the late stages when we sampled the area and diatoms were constrained to the northern part of our transect, over the Icelandic Basin (IB) and Icelandic Shelf (IS). Coccolithophores dominated the phytoplankton community, with a large distribution over the Rockall-Hatton Plateau (RHP) and IB. The Porcupine Abyssal Plain (PAP) region at the southern end of our transect was the region with the lowest biomass, as demonstrated by very low Chla concentrations and a community dominated by picophytoplankton. Early depletion of dissolved silicic acid (DSi) and increased stratification of the surface layer most likely triggered the end of the diatom bloom, leading to coccolithophore dominance. The chronic Si deficiency observed in the NEA could be linked to moderate Fe limitation, which increases the efficiency of the Si pump. TEP closely mirrored the distribution of both biogenic silica at depth and prymnesiophytes in the surface layer suggesting the sedimentation of the diatom bloom in the form of aggregates, but the relative contribution of diatoms and coccolithophores to carbon export

The late stage of the North East Atlantic (NEA) spring bloom was investigated during June 2005 along a transect section from 45 to 66° N between 15 and 20° W in order to characterize the contribution of siliceous and calcareous phytoplankton groups and describe their distribution in relation to environmental factors. We measured several biogeochemical parameters such as nutrients, surface trace metals, algal pigments, biogenic silica (BSi), particulate inorganic carbon (PIC) or calcium carbonate, particulate organic carbon, nitrogen and phosphorus (POC, PON and POP, respectively), as well as transparent exopolymer particles (TEP). Results were compared with other studies undertaken in this area since the JGOFS NABE program. Characteristics of the spring bloom generally agreed well with the accepted scenario for the development of the autotrophic community. The NEA seasonal diatom bloom was in the late stages when we sampled the area and diatoms were constrained to the northern part of our transect, over the Icelandic Basin (IB) and Icelandic Shelf (IS). Coccolithophores dominated the phytoplankton community, with a large distribution over the Rockall-Hatton Plateau (RHP) and IB. The Porcupine Abyssal Plain (PAP) region at the southern end of our transect was the region with the lowest biomass, as demonstrated by very low Chla concentrations and a community dominated by picophytoplankton. Early depletion of dissolved silicic acid (DSi) and increased stratification of the surface layer most likely triggered the end of the diatom bloom, leading to coccolithophore dominance. The chronic Si deficiency observed in the NEA could be linked to moderate Fe limitation, which increases the efficiency of the Si pump. TEP closely mirrored the distribution of both biogenic silica at depth and prymnesiophytes in the surface layer suggesting the sedimentation of the diatom bloom in the form of aggregates, but the relative contribution of diatoms and coccolithophores to carbon export

Transparent exopolymer particles (TEP) are the most ubiquitous gel particles in the ocean and form abiotically from dissolved precursors. Although these particles can accumulate at the ocean surface, being thus exposed to intense sunlight, the role of solar radiation for the assembly and degradation of TEP is unknown. In this study, we experimentally determined the effects of visible and ultraviolet B (UVB) radiation on (1) TEP degradation (photolysis experiments), (2) TEP assembly from dissolved polymers (photoinhibition experiments) and (3) TEP release by microorganisms. Solar radiation, particularly in the UVB range, caused significant TEP photolysis, with loss rates from 27 to 34% per day. Dissolved polysaccharides did not increase in parallel. No TEP were formed under UVB, visible or dark conditions, indicating that light does not promote TEP assembly. UVB radiation enhanced TEP release by microorganisms, possibly due to cell deaths, or as a protective measure. Increases in UVB may lead to enhanced TEP photolysis in the ocean, with further consequences for TEP dynamics and, ultimately, sea-air gas exchange.

Surface active substances (SAS) in the water column were measured by voltammetry using the electrochemical probe o-nitrophenol (ONP) during EIFEX, a mesoscale open ocean iron enrichment experiment in the Southern Ocean. SAS levels were low throughout the experiment (<0.005 – 0.03 mg L−1 Triton X-100 equivalents). Initially SAS was extremely low in the photic zone, but as the phytoplankton bloom developed concentrations markedly increased throughout the upper 100 m (∼0.02 mg L−1 Triton X-100 equivalents). Highest concentrations of SAS (>0.02 mg L−1 Triton X-100 equivalents) were found at the end of the bloom particularly at density discontinuities where organic material may accumulate. Exudates from diatoms appeared to be the major source of SAS during EIFEX, either from direct extracellular release or in the action of being grazed upon by zooplankton.

The effect of phytoplankton on the other compartments of a simple microbial loop consisting of dissolved matter, bacteria and nanoflagellates was investigated in three 1000 l mesocosms. These mesocosms were inoculated with a natural bacterial community and (1) no other additions (No Addition tank), (2) Phaeocystis globosa (Phaeocystis tank), or (3) 2 diatom species (Diatom tank). During the 20 d experiment, autotrophic activity was negligible in the No Addition tank. In contrast a small P. globosa bloom developed in the Phaeocystis tank and a large diatom bloom dominated the Diatom tank. In this paper we describe the experiment, the changes in chlorophyll a and heterotrophic nanoflagellate concentrations, as well as the cycling of nitrogen, phosphorus, and silica. Then we provide a synthesis of the structure and functioning of the microbial loops in these 3 systems using cluster analysis, a statistical pattern recognition tool. The goal was to test the hypothesis that differences in the resident phytoplankton populations would be reflected in (1) the composition and concentration of dissolved organic matter, (2) the composition of the bacterial community, (3) the food web, and (4) the cycling of elements and organic matter. In all 3 mesocosms, nitrate and silicic acid remained abundant. Orthophosphate was preferred by diatoms, whereas Phaeocystis appeared to prefer dissolved organic phosphorus. The hypothesis that phytoplankton composition shapes the structure and functioning of the microbial loop was partially supported: 6 d after inoculation each mesocosm exhibited a distinct organic matter signature. After 10 to 12 d, concentrations of heterotrophic nanoflagellates were high enough to exert significant grazing pressure in all 3 mesocosms. A parallel shift in bacterial community composition was visible in all mesocosms at this time, possibly reflecting grazing pressure. The food-web structure developed divergently in the 3 mesocosms during the second half of the e

We monitored the dynamics of extracellular organic matter in 3 mesocosms: one dominated by a heterotrophic (microbial) community with negligible autotrophic activity (net heterotrophic system), a second where a small Phaeocystis bloom developed (production and loss almost balanced), and a third harboring a large diatom bloom (net autotrophic system). In all mesocosms, meso- and macroscopic heterotrophic organisms were excluded to primarily study extracellular organic matter production and turnover by specific algae and microbial loop organisms, respectively. Concentration and composition of dissolved organic matter (DOM), i.e. dissolved organic carbon (DOC), monosaccharides and total carbohydrates (MCHO and TCHO), free and combined neutral carbohydrates (DFCHO and DCCHO), as well as free and combined amino acids (DFAA and DCAA) were measured. In addition, net and gross community production rates were determined to calculate C-budgets. Whereas concentrations and composition of MCHO differed very little among the 3 mesocosms, dynamics of TCHO, DFCHO, and DCCHO differed significantly. Concentrations of DFAA were higher in both algal mesocosms compared to the heterotrophic system, and composition of DFAA was significantly different in the Phaeocystis and Diatom tanks. The composition and concentration of DCAA, however, were similar in all 3 mesocosms. Total dissolved carbohydrates and amino acids comprised a substantial fraction of the DOC pool. Dynamics of these DOC fractions, however, could only partly explain those of DOC, implying either that other dissolved compounds were important for overall C-cycling, or that microbial degradation of DOM affects the detection of carbohydrates and protein components.

Surface active substances (SAS) in the water column were measured by voltammetry using the electrochemical probe o-nitrophenol (ONP) during EIFEX, a mesoscale open ocean iron enrichment experiment in the Southern Ocean. SAS levels were low throughout the experiment (<0.005 – 0.03 mg L−1 Triton X-100 equivalents). Initially SAS was extremely low in the photic zone, but as the phytoplankton bloom developed concentrations markedly increased throughout the upper 100 m (∼0.02 mg L−1 Triton X-100 equivalents). Highest concentrations of SAS (>0.02 mg L−1 Triton X-100 equivalents) were found at the end of the bloom particularly at density discontinuities where organic material may accumulate. Exudates from diatoms appeared to be the major source of SAS during EIFEX, either from direct extracellular release or in the action of being grazed upon by zooplankton.

We monitored the dynamics of extracellular organic matter in 3 mesocosms: one dominated by a heterotrophic (microbial) community with negligible autotrophic activity (net heterotrophic system), a second where a small Phaeocystis bloom developed (production and loss almost balanced), and a third harboring a large diatom bloom (net autotrophic system). In all mesocosms, meso- and macroscopic heterotrophic organisms were excluded to primarily study extracellular organic matter production and turnover by specific algae and microbial loop organisms, respectively. Concentration and composition of dissolved organic matter (DOM), i.e. dissolved organic carbon (DOC), monosaccharides and total carbohydrates (MCHO and TCHO), free and combined neutral carbohydrates (DFCHO and DCCHO), as well as free and combined amino acids (DFAA and DCAA) were measured. In addition, net and gross community production rates were determined to calculate C-budgets. Whereas concentrations and composition of MCHO differed very little among the 3 mesocosms, dynamics of TCHO, DFCHO, and DCCHO differed significantly. Concentrations of DFAA were higher in both algal mesocosms compared to the heterotrophic system, and composition of DFAA was significantly different in the Phaeocystis and Diatom tanks. The composition and concentration of DCAA, however, were similar in all 3 mesocosms. Total dissolved carbohydrates and amino acids comprised a substantial fraction of the DOC pool. Dynamics of these DOC fractions, however, could only partly explain those of DOC, implying either that other dissolved compounds were important for overall C-cycling, or that microbial degradation of DOM affects the detection of carbohydrates and protein components.

The effect of phytoplankton on the other compartments of a simple microbial loop consisting of dissolved matter, bacteria and nanoflagellates was investigated in three 1000 l mesocosms. These mesocosms were inoculated with a natural bacterial community and (1) no other additions (No Addition tank), (2) Phaeocystis globosa (Phaeocystis tank), or (3) 2 diatom species (Diatom tank). During the 20 d experiment, autotrophic activity was negligible in the No Addition tank. In contrast a small P. globosa bloom developed in the Phaeocystis tank and a large diatom bloom dominated the Diatom tank. In this paper we describe the experiment, the changes in chlorophyll a and heterotrophic nanoflagellate concentrations, as well as the cycling of nitrogen, phosphorus, and silica. Then we provide a synthesis of the structure and functioning of the microbial loops in these 3 systems using cluster analysis, a statistical pattern recognition tool. The goal was to test the hypothesis that differences in the resident phytoplankton populations would be reflected in (1) the composition and concentration of dissolved organic matter, (2) the composition of the bacterial community, (3) the food web, and (4) the cycling of elements and organic matter. In all 3 mesocosms, nitrate and silicic acid remained abundant. Orthophosphate was preferred by diatoms, whereas Phaeocystis appeared to prefer dissolved organic phosphorus. The hypothesis that phytoplankton composition shapes the structure and functioning of the microbial loop was partially supported: 6 d after inoculation each mesocosm exhibited a distinct organic matter signature. After 10 to 12 d, concentrations of heterotrophic nanoflagellates were high enough to exert significant grazing pressure in all 3 mesocosms. A parallel shift in bacterial community composition was visible in all mesocosms at this time, possibly reflecting grazing pressure. The food-web structure developed divergently in the 3 mesocosms during the second half of the e

Surface active substances (SAS) in the water column were measured by voltammetry using the electrochemical probe o-nitrophenol (ONP) during EIFEX, a mesoscale open ocean iron enrichment experiment in the Southern Ocean. SAS levels were low throughout the experiment (<0.005 – 0.03 mg L−1 Triton X-100 equivalents). Initially SAS was extremely low in the photic zone, but as the phytoplankton bloom developed concentrations markedly increased throughout the upper 100 m (∼0.02 mg L−1 Triton X-100 equivalents). Highest concentrations of SAS (>0.02 mg L−1 Triton X-100 equivalents) were found at the end of the bloom particularly at density discontinuities where organic material may accumulate. Exudates from diatoms appeared to be the major source of SAS during EIFEX, either from direct extracellular release or in the action of being grazed upon by zooplankton.

We monitored the dynamics of extracellular organic matter in 3 mesocosms: one dominated by a heterotrophic (microbial) community with negligible autotrophic activity (net heterotrophic system), a second where a small Phaeocystis bloom developed (production and loss almost balanced), and a third harboring a large diatom bloom (net autotrophic system). In all mesocosms, meso- and macroscopic heterotrophic organisms were excluded to primarily study extracellular organic matter production and turnover by specific algae and microbial loop organisms, respectively. Concentration and composition of dissolved organic matter (DOM), i.e. dissolved organic carbon (DOC), monosaccharides and total carbohydrates (MCHO and TCHO), free and combined neutral carbohydrates (DFCHO and DCCHO), as well as free and combined amino acids (DFAA and DCAA) were measured. In addition, net and gross community production rates were determined to calculate C-budgets. Whereas concentrations and composition of MCHO differed very little among the 3 mesocosms, dynamics of TCHO, DFCHO, and DCCHO differed significantly. Concentrations of DFAA were higher in both algal mesocosms compared to the heterotrophic system, and composition of DFAA was significantly different in the Phaeocystis and Diatom tanks. The composition and concentration of DCAA, however, were similar in all 3 mesocosms. Total dissolved carbohydrates and amino acids comprised a substantial fraction of the DOC pool. Dynamics of these DOC fractions, however, could only partly explain those of DOC, implying either that other dissolved compounds were important for overall C-cycling, or that microbial degradation of DOM affects the detection of carbohydrates and protein components.

The effect of phytoplankton on the other compartments of a simple microbial loop consisting of dissolved matter, bacteria and nanoflagellates was investigated in three 1000 l mesocosms. These mesocosms were inoculated with a natural bacterial community and (1) no other additions (No Addition tank), (2) Phaeocystis globosa (Phaeocystis tank), or (3) 2 diatom species (Diatom tank). During the 20 d experiment, autotrophic activity was negligible in the No Addition tank. In contrast a small P. globosa bloom developed in the Phaeocystis tank and a large diatom bloom dominated the Diatom tank. In this paper we describe the experiment, the changes in chlorophyll a and heterotrophic nanoflagellate concentrations, as well as the cycling of nitrogen, phosphorus, and silica. Then we provide a synthesis of the structure and functioning of the microbial loops in these 3 systems using cluster analysis, a statistical pattern recognition tool. The goal was to test the hypothesis that differences in the resident phytoplankton populations would be reflected in (1) the composition and concentration of dissolved organic matter, (2) the composition of the bacterial community, (3) the food web, and (4) the cycling of elements and organic matter. In all 3 mesocosms, nitrate and silicic acid remained abundant. Orthophosphate was preferred by diatoms, whereas Phaeocystis appeared to prefer dissolved organic phosphorus. The hypothesis that phytoplankton composition shapes the structure and functioning of the microbial loop was partially supported: 6 d after inoculation each mesocosm exhibited a distinct organic matter signature. After 10 to 12 d, concentrations of heterotrophic nanoflagellates were high enough to exert significant grazing pressure in all 3 mesocosms. A parallel shift in bacterial community composition was visible in all mesocosms at this time, possibly reflecting grazing pressure. The food-web structure developed divergently in the 3 mesocosms during the second half of the e

Surface active substances (SAS) in the water column were measured by voltammetry using the electrochemical probe o-nitrophenol (ONP) during EIFEX, a mesoscale open ocean iron enrichment experiment in the Southern Ocean. SAS levels were low throughout the experiment (<0.005 – 0.03 mg L−1 Triton X-100 equivalents). Initially SAS was extremely low in the photic zone, but as the phytoplankton bloom developed concentrations markedly increased throughout the upper 100 m (∼0.02 mg L−1 Triton X-100 equivalents). Highest concentrations of SAS (>0.02 mg L−1 Triton X-100 equivalents) were found at the end of the bloom particularly at density discontinuities where organic material may accumulate. Exudates from diatoms appeared to be the major source of SAS during EIFEX, either from direct extracellular release or in the action of being grazed upon by zooplankton.

info:eu-repo/semantics/published

Particulate organic matter which displays a fairly constant C:N ratio (Redfield ratio) consists of different particle types, including organisms, detritus and transparent exopolymer particles (TEP). Whereas the C:N-ratio of cells frequently lies below Redfield, the C:N ratio of TEP, which are formed abiotically from exuded polysaccharides is higher. The mean molar C:N ratios of TEP derived from cultures and natural diatom populations grown under different conditions exhibited a mean value of 26. This is considered a low estimate, as the nitrogen fraction can be explained through adsorption of amino acids onto TEP. The TEP-derived carbon concentration was calculated in 6 marine environments, based on the derived relationship between carbon and TEP. The contribution of TEP-carbon to the total carbon pool was large, in some cases as high as the POC contribution. TEP are also an essential component of marine snow and contribute appreciably to carbon flux. Because TEP are enriched in carbon over nitrogen, an increase in the formation rate of TEP due to global warming would provide a pathway for the sequestration of excess carbon to the deeper water column.

Although detailed temporal studies of marine planktonic cyanobacteria have been reported, relatively little is known about variation of other marine picoplankton groups on time scales on the order of months to years. In this study, we followed variation in relative abundance of planktonic archaeal rRNA at 5 depths in the upper 300 m of the Santa Barbara Channel (SBC) over 32 mo. Small subunit rRNA targeted oligonucleotide probes were used to quantify archaeal, bacterial, and eucaryal rRNA relative abundance. Archaea subgroups were quantified using probes specific to 2 planktonic archaeal groups, termed GI and GII. The archaeal rRNA signal in the upper 20 m was characterized by intermittent 'blooms' that coincided with increases in the relative abundance of GII rRNA as well as decreases in chlorophyll a (chl a). At greater depths, archaeal rRNA abundance was consistently elevated, approaching bacterial rRNA abundance. The GI rRNA accounted for the majority of the deeper archaeal rRNA signal throughout the time series. Consistent with a previous report, the 2 groups of archaea had maximal rRNA abundance at different depths. The majority of the variability in the GI archaeal signal in the upper 75 m (77%) could be attributed to a positive relationship with nutrients and negative relationships with prokaryotic abundance, chl a, and temperature; although, at most, 41% of the variability at each depth could be explained. Only 21% of the variability in the GII rRNA signal could be explained by the variables included in the model. Leucine incorporation rates and, to a lesser extent, prokaryote abundance, were highly correlated with chl a and POC levels. Only weak relationships were observed between prokaryote abundance or leucine incorporation and increases in temperature. The results of linear regressions between prokaryote abundance and leucine incorporation suggest that top-down ecological controls may regulate prokaryotic assemblages in this dynamic coastal system. The

The development and decline of a phytoplankton spring bloom dominated by the prymnesiophyte Phaeocystis pouchetii were studied in Balsfjord, northern Norway between 30 March and 27 May 1992. At a fixed station, the concentration and composition of suspended particulate matter was monitored and compared to the particulate matter collected in sediment traps at six different depths. Direct sedimentation of phytoplankton contributed a minor fraction to particle flux and was confined to a few diatom genera. No evidence was found for pronounced aggregation of Phaeocystis colonies during bloom decline or direct sedimentation of either Phaeocystis colonies or single cells, Particle flux was dominated by faecal-pellet sedimentation during most of the study period, suggesting zooplankton grazing to be a main loss factor. Despite an abrupt decrease in faecal-pellet sedimentation after the decline of the bloom, particulate-carbon sedimentation rates remained high. High post-bloom sedimentation rates were characterized by elevated C/N and C/Chl a ratios of largely amorphous sedimented material. Post-bloom sedimentation coincided with a decrease in transparent exopolymeric particles (TEP) in the surface layer, suggesting that this change resulted from aggregation and sedimentation of carbon-rich exopolymeric material accumulated in the surface layer in the course of the bloom. While organic-carbon accumulation indicates the significance of disintegration of Phaeocystis colonies, post-bloom mucilage sedimentation could be a secondary pathway for the vertical flux of Phaeocystis-derived organic matter.

While the aggregation and mass settlement of diatoms at the termination of blooms results in significant export of carbon from the surface ocean, the mechanisms of bloom aggregation have been poorly understood. The aggregation of a multispecies diatom bloom was investigated under controlled conditions in a 1200 liter, nutrient-enriched, laboratory mesocosm in order to elucidate the parameters sufficient to accurately predict bloom aggregation. A diverse bloom of diatoms dominated by several species of Chaetoceros and Thalassiosira progressed through a classic pattern of exponential, stationary, and senescent phases in the mesocosm. Aggregates larger than 0.5 mm became detectable on the eighth day after inoculation, and aggregates >1 mm increased exponentially from Day 10 onward producing the appearance of a mass aggregation event late on Day 10. The bloom aggregated sequentially with Thalassiosira dominating early aggregates and Chaetoceros dominating later ones. Chaetoceros resting spores formed only in aggregates. Aggregation was not linked to nutrient depletion or to the physiological state of the cells since the onset of aggregation and the mass aggregation event occurred 1 to 3 days prior to nutrient depletion and while carbon:nitrogen ratios of cells were still very low and growth rates high. Moreover, visible aggregates did not form in the mesocosm until cell abundances were considerably higher than abundances observed to aggregate in nature, suggesting that aggregation was not strongly linked to phytoplankton cell concentration. Complementary studies in this volume clarify the role of non-phytoplankton particles in aggregation of the mesocosm bloom. The mesocosm approach proved highly effective in producing an aggregating diatom bloom under controlled conditions.

The development and decline of a phytoplankton spring bloom dominated by the prymnesiophyte Phaeocystis pouchetii were studied in Balsfjord, northern Norway between 30 March and 27 May 1992. At a fixed station, the concentration and composition of suspended particulate matter was monitored and compared to the particulate matter collected in sediment traps at six different depths. Direct sedimentation of phytoplankton contributed a minor fraction to particle flux and was confined to a few diatom genera. No evidence was found for pronounced aggregation of Phaeocystis colonies during bloom decline or direct sedimentation of either Phaeocystis colonies or single cells, Particle flux was dominated by faecal-pellet sedimentation during most of the study period, suggesting zooplankton grazing to be a main loss factor. Despite an abrupt decrease in faecal-pellet sedimentation after the decline of the bloom, particulate-carbon sedimentation rates remained high. High post-bloom sedimentation rates were characterized by elevated C/N and C/Chl a ratios of largely amorphous sedimented material. Post-bloom sedimentation coincided with a decrease in transparent exopolymeric particles (TEP) in the surface layer, suggesting that this change resulted from aggregation and sedimentation of carbon-rich exopolymeric material accumulated in the surface layer in the course of the bloom. While organic-carbon accumulation indicates the significance of disintegration of Phaeocystis colonies, post-bloom mucilage sedimentation could be a secondary pathway for the vertical flux of Phaeocystis-derived organic matter.

While the aggregation and mass settlement of diatoms at the termination of blooms results in significant export of carbon from the surface ocean, the mechanisms of bloom aggregation have been poorly understood. The aggregation of a multispecies diatom bloom was investigated under controlled conditions in a 1200 liter, nutrient-enriched, laboratory mesocosm in order to elucidate the parameters sufficient to accurately predict bloom aggregation. A diverse bloom of diatoms dominated by several species of Chaetoceros and Thalassiosira progressed through a classic pattern of exponential, stationary, and senescent phases in the mesocosm. Aggregates larger than 0.5 mm became detectable on the eighth day after inoculation, and aggregates >1 mm increased exponentially from Day 10 onward producing the appearance of a mass aggregation event late on Day 10. The bloom aggregated sequentially with Thalassiosira dominating early aggregates and Chaetoceros dominating later ones. Chaetoceros resting spores formed only in aggregates. Aggregation was not linked to nutrient depletion or to the physiological state of the cells since the onset of aggregation and the mass aggregation event occurred 1 to 3 days prior to nutrient depletion and while carbon:nitrogen ratios of cells were still very low and growth rates high. Moreover, visible aggregates did not form in the mesocosm until cell abundances were considerably higher than abundances observed to aggregate in nature, suggesting that aggregation was not strongly linked to phytoplankton cell concentration. Complementary studies in this volume clarify the role of non-phytoplankton particles in aggregation of the mesocosm bloom. The mesocosm approach proved highly effective in producing an aggregating diatom bloom under controlled conditions.

The development and decline of a phytoplankton spring bloom dominated by the prymnesiophyte Phaeocystis pouchetii were studied in Balsfjord, northern Norway between 30 March and 27 May 1992. At a fixed station, the concentration and composition of suspended particulate matter was monitored and compared to the particulate matter collected in sediment traps at six different depths. Direct sedimentation of phytoplankton contributed a minor fraction to particle flux and was confined to a few diatom genera. No evidence was found for pronounced aggregation of Phaeocystis colonies during bloom decline or direct sedimentation of either Phaeocystis colonies or single cells, Particle flux was dominated by faecal-pellet sedimentation during most of the study period, suggesting zooplankton grazing to be a main loss factor. Despite an abrupt decrease in faecal-pellet sedimentation after the decline of the bloom, particulate-carbon sedimentation rates remained high. High post-bloom sedimentation rates were characterized by elevated C/N and C/Chl a ratios of largely amorphous sedimented material. Post-bloom sedimentation coincided with a decrease in transparent exopolymeric particles (TEP) in the surface layer, suggesting that this change resulted from aggregation and sedimentation of carbon-rich exopolymeric material accumulated in the surface layer in the course of the bloom. While organic-carbon accumulation indicates the significance of disintegration of Phaeocystis colonies, post-bloom mucilage sedimentation could be a secondary pathway for the vertical flux of Phaeocystis-derived organic matter.

While the aggregation and mass settlement of diatoms at the termination of blooms results in significant export of carbon from the surface ocean, the mechanisms of bloom aggregation have been poorly understood. The aggregation of a multispecies diatom bloom was investigated under controlled conditions in a 1200 liter, nutrient-enriched, laboratory mesocosm in order to elucidate the parameters sufficient to accurately predict bloom aggregation. A diverse bloom of diatoms dominated by several species of Chaetoceros and Thalassiosira progressed through a classic pattern of exponential, stationary, and senescent phases in the mesocosm. Aggregates larger than 0.5 mm became detectable on the eighth day after inoculation, and aggregates >1 mm increased exponentially from Day 10 onward producing the appearance of a mass aggregation event late on Day 10. The bloom aggregated sequentially with Thalassiosira dominating early aggregates and Chaetoceros dominating later ones. Chaetoceros resting spores formed only in aggregates. Aggregation was not linked to nutrient depletion or to the physiological state of the cells since the onset of aggregation and the mass aggregation event occurred 1 to 3 days prior to nutrient depletion and while carbon:nitrogen ratios of cells were still very low and growth rates high. Moreover, visible aggregates did not form in the mesocosm until cell abundances were considerably higher than abundances observed to aggregate in nature, suggesting that aggregation was not strongly linked to phytoplankton cell concentration. Complementary studies in this volume clarify the role of non-phytoplankton particles in aggregation of the mesocosm bloom. The mesocosm approach proved highly effective in producing an aggregating diatom bloom under controlled conditions.

Marine species and ecosystems are exposed to a wide range of environmental change – both detrimental (threats) and beneficial – due to human activities. Some of the changes are global, whereas others are regional or local. It is important to distinguish the scale of each threat, as the solutions will differ as they are scale dependent. For example, the mitigation of a global problem requires a global response, which is more difficult to achieve than addressing a local problem with a local response. These wide-ranging changes are often referred to drivers or stressors.