No abstract

ABSTRACT-Vertical heterogeneity of pelagic organisms in a subsurface bloom of Gyrodinium aureolum (Hulburt) was studied on the decimeter scale by a high resolution sampler In the Kattegat, Denmark, durlng August 1990. Maximum concentrations of 3.25 X 106 G. aureolum I -' and 72 pg chlorophyll a I-' were recorded in the pycnocline at a depth of 14 m with 5 "10 of surface irrad~ance. Dinoflagellates were confined to a 1 m thick layer which contained more chlorophyll than the rest of the water column In situ primary production rates suggested a phytoplankton turnover tlme of l d. Bacterioplankton production and turnover rate and microzooplankton abundance were significantly reduced in the dinoflagellate layer

During 9 to 25 June 1987, carbon budgets were established for estuarine enclosures manipulated by additions of nutrients and suspension-feeding bivalves An intensive sampling program and a detailed examination of autotrophic and heterotrophic nlicroorganlsms enabled construction of carbon budgets of the microbial food web and comparison of flow rates through a number of microbial components. Phytoplankton biomass and production covaried, and, as expected, lowest values were recorded in enclosures with added mussels, and highest values in enclosures with added nutrients. Bacteria and heterotrophic nanoflagellates peaked a few days after maxima in phytoplankton biomass and production. In enclosures with added mussels, biomasses were lower for bacteria and microzooplankton, and mesozooplankton, but slightly higher for heterotrophic nanoflagellates. Bacteria, flagellates, and microzooplankton, mostly ciliates, dominated heterotrophic processes, whereas larger mesozooplankton ingestion did not exceed 5 % of phytoplankton primary production. Microzooplankton and flagellate clearances were higher in enclosures with added nutrients, whereas no such changes were found in the macrozooplankton, probably because the duration of the experiments did not allow full development of the macrozooplankton. The added mussels dominated heterotrophic consumption and controlled organisms > 20 Fm. Exclusion of mussels induced a primary dominance of microzooplankton followed by a subsequent increase of mesozooplankton. Additions of nutrients and filtration by suspension-feeding bivalves caused qualitative and quantitative changes at all levels in the microbial food web. These changes were measured from a large number of microbial components and allowed balances of the carbon budgets to be made as well as identification of factors controlling the structure and function of the pelagic carbon cycle.

During April, June/July and September 1986, phytoplankton biomass was followed in marine enclosures in the Roskilde Fjord, Denmark, manipulated by the addition of benthic suspension feeders, planktivorous fish, nutrients and contact to the sediment. During periods with high insolation and temperatures from 11 to 22 "C, the manipulations caused marked changes in the development of phytoplankton biomass. Additions of Mytilus edulis reduced phytoplankton biomass to between 10 and 59 % of controls, whereas addition of nutrients raised phytoplankton biomass to an average of 256 % of controls. Generally, low growth rates of mussels were found in enclosures containing mussels alone. Addition of planktivorous fish and nutrients increased growth rates of mussels. During June/July, when inorganic nitrogen limited phytoplankton growth, autotrophic picoplankton (1 to 2 pm cell diameter) constituted 70 to 93 % of phytoplankton biovolume in enclosures containing mussels compared to 4 to 20 % in controls. The mussels reduced phytoplankton biomass by only about 50 % during this period, presumably due to low retention efficiency of the small cells. In April and September, however, when nitrogen did not control phytoplankton growth, picoplankton comprised < 0.001 % of phytoplankton biovolume. During September (temperatures 11 to 13OC), M. edulis reduced chlorophyll levels to 10 O/ O of controls and the effects of nutrient additions were significantly reduced in enclosures containing mussels. The effect of fish additions revealed that zooplankton grazing removed (average for June and September) about 20 % of phytoplankton biomass. The sediment acted prin~arily as a nutrient source during summer However, in April at temperatures > 6 "C and in September, benthic suspension feeders maintained chlorophyll levels below those in enclosures with no sediment contact. During April, only minor changes were recorded in the chlorophyll levels in the various enclosures, presumably due to the low temperature (0 to 4.5OC) and a phytoplankton population of decaying diatoms. Thus, several factors or combinations of factors controlled the phytoplankton biomass in the enclosures. The most important of these factors was the balance between nutrient input, phytoplankton size structure and the physiological state of the mussels.

Subsurface maxima in the distribution of autotrophic flagellates and diatoms were found in the nutricline on 3 successive transects across the Skagerrak between Denmark and Norway, during June 1990. The subsurface phytoplankton maximum increased during the 6 d investigation period both in intensity and horizontal extension and followed the nutricline, which descended from ca 15 to ca 25 m depth, while the pycnocline remained at 5 to 10 m depth. The descent of the nutricline and the subsurface maximum is interpreted as the utilization of nutrients from the bottom water for new production. This interpretation is supported by a simple dynamic model simulating variations in biomass, irradiance and nutrient concentration with time and depth. About half of the primary production in the water column was associated with the subsurface maximum. Experiments using 15N-nitrate and 32P-phosphate confirmed that the subsurface maximum was partly supported by new production, whlle surface phytoplankton relied mainly on regenerated nutrients. Measurements of heterotrophic activity and vertical carbon flux suggest that the net primary production in the subsurface maximum was roughly evenly allocated to biomass build-up, heterotrophic remineralisation and sedimentation. The heterotrophic community responded to the subsurface phytoplankton maximum with corresponding maxima of bacterioplankton biomass and production and of biomasses of nano-and microzooplankton, whereas copepod biomass remained higher in the surface layer than in the subsurface phytoplankton maxlmum.