Description: Biogeochemical cycles of carbon, nutrients, and oxygen transmit mean states, trends and variations of the physical realm in coastal upwelling systems to their food webs and determine their role in regional budgets of greenhouse gases. This contribution focuses on biogeochemical processes in the northern Benguela Upwelling System (NBUS), where low oxygen levels in upwelling source water are a major influence on carbon and nutrient cycles. Based on measurements during numerous expeditions and results of 3-D regional ecosystem modeling (project GENUS; Geochemistry and Ecology of the Namibian Upwelling System) we here examine source water character, effects of low oxygen conditions on nutrient masses and ratios, and of diazotrophic N2-fixation on productivity of the system and its transition to the adjacent eastern South Atlantic. In available observations, the effects of denitrification in water and sediment and phosphate release from sediments are minor influences on nitrate:phosphate ratios of the system, and excess phosphate in aged upwelling water is inherited from upwelling source water. Contrary to expectation and model results, the low N:P ratios do not trigger diazotrophic N2-fixation in the fringes of the upwelling system, possibly due to a lack of seeding populations of Trichodesmium. We also examine the flux of carbon from the sea surface to either sediment, the adjacent sub-thermocline ocean, or to regenerated nutrients and CO2. Observed fluxes out of the surface mixed layer are significantly below modeled fluxes, and suggest that regeneration of nutrients and CO2 is unusually intense in the mixed layer. This contributes to very high fluxes of CO2 from the ocean to the regional atmosphere, which is not compensated for by N2-fixation. Based on observations, the NBUS thus is a significant net CO2 source (estimated at 14.8 Tg C a− 1), whereas the CO2 balance is closed by N2-fixation in the model. Methane concentrations were low in surface waters in on-line measurements during 1 expedition, and based on these our estimate for the emission of methane for the entire Benguela system is below 0.2 Tg CH4 a− 1.

Description: Literature data from 1905/06, 1912/13 and 1949/50 were compared with recent data (2001-2003) from Kiel Bight in order to investigate changes in phytoplankton composition and biomass, which may serve as indicators of environmental changes. In terms of biomass, diatomophyceae and dinophyceae are by far the most important groups. Their ratio is still close to unity. The share of diatomophyceae increased strongly in years with exceptionally high summer blooms (2001) or exceptionally early spring blooms (2003). The summer and autumn blooms of Chaetoceros and Skeletonema, detected in the early 20th century, are replaced by other diatoms (Cerataulina pelagica, Dactyliosolen fragilissimus, Proboscia alata, Pseudo-nitzschia spp.). Chaetoceros and Skeletonema are still important components of the spring blooms. Now as before, the autumn blooms are dominated by Ceratium spp., sometimes also by diatoms. Newly appearing bloom-forming species are mostly potentially toxic (Dicryocha speculum, Prorocentrum minimum, Pseudo-nitzschia spp.). The total phytoplankton biomass has roughly doubled in the course of the last century. The reference condition for phytoplankton biomass in Kiel Bight in the sense of the Water Framework Directive was defined at 55 mg C m(-3) (+/- 10%, annual mean). The mean annual biomass of diatomophyceae and dinophyceae was 25 mg C m-3 (+/- 40%) for each, indicating that the sum of their carbon biomass amounted to 90% (+/- 10%) of the total phytoplankton biomass on an annual average. Diatomophyceae represented at least 80% of carbon biomass in the spring bloom peak at the beginning of the 20th century

Description: In July 2007, phosphorus input by an upwelling event along the east coast of Gotland Island and the response of filamentous cyanobacteria were studied to determine whether introduced phosphorus can intensify cyanobacterial bloom formation in the eastern Gotland Basin. Surface temperature, nutrient concentrations, phytoplankton biomass and its stoichiometry, as well as phosphate uptake rates were determined in two transects between the coasts of Gotland and Latvia and in a short grid offshore of Gotland. In the upwelling area, surface temperatures of 11–12 °C and average dissolved inorganic phosphorus (DIP) concentrations of 0.26 μM were measured. Outside the upwelling, surface temperatures were higher (15.5–16.6 °C) and DIP supplies in the upper 10 m layer were exhausted. Nitrite and nitrate concentrations (0.01–0.22 μM) were very low within and outside the upwelling region. Abundances of filamentous cyanobacteria were highly reduced in the upwelling area, accounting for only 1.4–6.0% of the total phytoplankton biomass, in contrast to 18–20% outside the upwelling. The C:P ratio of filamentous cyanobacteria varied between 32.8 and 310 in the upwelling region, most likely due to the introduction of phosphorus-depleted organisms into the upwelling water. These organisms accumulate DIP in upwelling water and have lower C:P ratios as long as they remain in DIP-rich water. Thus, diazotrophic cyanobacteria benefit from phosphorus input directly in the upwelling region. Outside the upwelling region, the C:P ratios of filamentous cyanobacteria varied widely, between 240 and 463, whereas those of particulate material in the water ranged only between 96 and 224. To reduce their C:P ratio from 300 to 35, cyanobacteria in the upwelling region had to take up 0.05 mmol m−3 DIP, which is about 20% of the available DIP. Thus, a larger biomass of filamentous cyanobacteria may be able to benefit from a given DIP input. As determined from the DIP uptake rates measured in upwelling cells, the time needed to reduce the C:P ratio from 300 to 35 was too long to explain the huge bloom formations that typically occur in summer. However, phosphorus uptake rates increased significantly with increasing C:P ratios, allowing phosphorus accumulation within 4–5 days, a span of time suitable for bloom formation in July and August.

Description: Cyanobacteria blooms in the Baltic Sea appear after upwelling events, which transport phosphate-rich intermediate water to the surface. The growth potential of diazotrophic cyanobacteria in upwelled water was studied in a mesocosm (tank) experiment in summer 2007. An Anabaena bloom was only induced in the tanks filled with upwelled surface water but not in those filled with surface water from outside the upwelling cell and with intermediate water. The low initial cyanobacteria biomass in the intermediate water could not grow to bloom concentrations within three weeks. It is concluded that mixing of upwelled water with surrounding surface water forms a precondition for a cyanobacteria bloom. An additional mesocosm experiment conducted in 2009 revealed that mixing of intermediate water with surface water had the same stimulating effect on nitrogen fixation and cyanobacteria growth as artificial phosphate input. Phosphate input stimulates the growth of Nodularia and Anabaena more than that of Aphanizomenon. We suggest that the upwelled phosphate-rich intermediate water has to be mixed with the surface water containing physiologically “young” cyanobacteria biomass of at least 20 mg/m3 as an inoculum in order to initiate a cyanobacteria bloom.

Description: Massive cyanobacteria blooms occur almost every summer in the Baltic Sea but the capability to quantitatively predict their extent and intensity is poorly developed. Here we analyse statistical relationships between multi-decadal satellite-derived time series of the frequency of cyanobacteria surface accumulations (FCA) in the central Baltic Sea Proper and a suite of environmental variables. Over the decadal scale (∼5-20 years) FCA was highly correlated (R2 ∼ 0.69) with a set of biogeochemical variables related to the amount of phosphorus and hypoxia in bottom layers. Water temperature in the surface layer was also positively correlated with FCA at the decadal scale. In contrast, the inter-annual variations in FCA had no correlation with the biogeochemical variables. Instead, significant correlations were found with the solar shortwave direct flux in July and the sea-surface temperature, also in July. It thus appears that it is not possible to predict inter-annual fluctuations in cyanobacteria blooms from water chemistry. Moreover, environmental variables could only explain about 45% of the inter-annual variability in FCA, probably because year-to-year variations in FCA are significantly influenced by biological interactions.