Understanding human impact on the Baltic ecosystem: changing views in recent decades.

Grave environmental problems, including contamination of biota by organochlorines and heavy metals, and increasing deep-water oxygen deficiency, were discovered in the Baltic Sea in the late 1960s. Toxic pollutants, including the newly discovered PCB, were initially seen as the main threat to the Baltic ecosystem, and the impaired reproduction found in Baltic seals and white-tailed eagles implied a threat also to human fish eaters. Countermeasures gradually gave results, and today the struggle to limit toxic pollution of the Baltic is an international environmental success story. Calculations showed that Baltic deep-water oxygen consumption must have increased, and that the Baltic nutrient load had grown about fourfold for nitrogen and 8 times for phosphorus. Evidence of increased organic production at all trophic levels in the ecosystem gradually accumulated. Phosphorus was first thought to limit Baltic primary production, but measurements soon showed that nitrogen is generally limiting in the open Baltic proper, except for nitrogen-fixing cyanobacteria. Today, the debate is concerned with whether phosphorus, by limiting nitrogen-fixers, can control open-sea ecosystem production, even where phytoplankton is clearly nitrogen limited. The Baltic lesson teaches us that our views of newly discovered environmental problems undergo repeated changes, and that it may take decades for scientists to agree on their causes. Once society decides on countermeasures, it may take decades for them to become effective, and for nature to recover. Thus, environmental management decisions can hardly wait for scientific certainty. We should therefore view environmental management decisions as experiments, to be monitored, learned from, and then modified as needed.

Nitrogen and the Baltic Sea: managing nitrogen in relation to phosphorus.

The Baltic is a large, brackish sea (4 x 10(5) km2) extending from 54N to approximately 66N, with a fourfold larger drainage area (population 8 x 10(7). Surface salinity (2 to 8 PSU) and hence biodiversity is low. In the last century, annual nutrient loads increased to 10(6) metric tons N and 5 x 10(4) ton P. Eutrophication is evident in the N-limited south, where cyanobacteria fix 2 to 4 x 10(5) ton N each summer, Secchi depths have been halved, and O2-deficient bottom areas have spread. Production remains low in the P-limited north. In nutrient-enriched coastal areas, phytoplankton blooms, toxic at times, and filamentous macroalgae reduce amenity values. Loads need to be reduced of both N, to reduce production, and P, to limit N-fixing cyanobacterial blooms. When large N-load reductions have been achieved locally, algal biomass has declined. So far, P loads have been reduced more than N loads. If this continues, a P-limited Baltic proper may result, very different from previous N-limited conditions. Reaching the management goal of halved anthropogenic N and P loads at minimum cost will require better understanding of biogeochemical nutrient cycles, economic evaluation of proposed measures, and improved stakeholder participation.

Benthic predator-prey interactions: evidence that adult Monoporeia affinis (Amphipoda) eat postlarval Macoma balthica (Bivalvia).

Predation by adults of the amphipod Monoporeia affinis on the plantigrade postlarval stage of the bivalve Macoma balthica was studied in the laboratory. We confirmed that M. affinis consumes small M. balthica. Amphipods offered 14C-labelled postlarvae took up the radioactive tracer, while those presented Rhodamine B-stained postlarvae acquired gut contents fluorescing strongly in orange, whereas control amphipods did not. Both labelling methods proved convenient to use in laboratory experiments, and are particularly useful when organisms lack structures that can be easily identified after being ingested, or when cross-over reactions may bias the results of immunoassays. The results reported here support the conclusion from earlier studies that predation by M. affinis on M. balthica can affect population dynamics of M. balthica and is likely to be an important structuring factor in the low-diversity benthic macrofauna community of the Baltic Sea.

Breakdown of phytoplankton pigments in Baltic sediments: effects of anoxia and loss of deposit-feeding macrofauna.

We examined the decay of chlorophyll a and the carotenoid fucoxanthin in oxic and anoxic sediment microcosms, with and without the deposit-feeding benthic amphipod Monoporeia affinis, over 57 days at 5 degrees C. Deep frozen phytoplankton from the Baltic Sea proper was added to all but a few microcosms. The range of chlorophyll a and fucoxanthin decay rate constants observed in microcosms with phytoplankton addition was 0.04-0.07 day(-1). The fastest pigment decay and build-up of chlorophyll breakdown products after phytoplankton addition were found in oxic treatments with amphipods. No effects of amphipods on pigment breakdown were found in anoxic treatments, or in treatments without phytoplankton addition. Greater losses of chlorophyll a in oxic (96%) than in anoxic (80%) treatments after 57 days indicates that preservation of sedimentary organic matter will be enhanced during periods of anoxia. Due to slow recruitment and recolonization in Baltic sediments, a single anoxic event may cause long-term (years) absence of significant macrobenthos. Anoxic events will thus not only reduce decay of plant pigments, and presumably other organic matter, while they last, but will also have longer-term effects, through elimination of macrofauna, which when present enhance organic matter decomposition.

Meiofaunal prominence and benthic seasonality in a coastal marine ecosystem.

The muds of a shallow (7 m) site in Narragansett Bay, Rhode Island contained higher abundances of meiofauna (averaging 17×106 individuals per m2 and ash free dry weight of 2.9 g/m2 during a 3 year period) than have been found in any other sediment. The majority of sublittoral muds, worldwide, have been reported to contain about 106 individuals per m2. This difference is attributed primarily to differences in sampling techniques and laboratory processing.Extremely high meiofaunal abundances may have also occurred because Narragansett Bay sediments were a foodrich environment. While the quantity of organic deposition in the bay is not unusually high for coastal waters, this input, primarily composed of diatom detritus, may contain an unusually high proportion of labile organics. Furthermore, meiofauna could have thrived because of spatial segregation of meiofauna and macrofauna. While meiofauna were concentrated at the sediment-water interface, most macrofauna were subsurface deposit feeders. Macrofaunal competition with, and ingestion of meiofauna may thus have been minimized.The seasonal cycles of meiofauna and macrofauna were similar. Highest abundances and biomass were observed in May and June and lowest values in the late summer and fall. Springtime increases of meiofaunal abundance were observed in all depth horizons, to 10 cm. We hypothesize that phytoplankton detritus accumulated in the sediment during the winter and early spring, and that the benthos responded to this store of food when temperatures rose rapidly in the late spring. By late summer, the stored detritus was exhausted and the benthos declined.