Impacts of nutrient enrichment and sediment on phytoplankton community structure in the northern Baltic Sea

A three-week mesocosm experiment was conducted in order to study the effects of bottom sediment and nutrient enrichment on phytoplankton and zooplankton community structure in the Archipelago Sea, northern Baltic Sea. The transparent polyethylene enclosures included the whole water column and varied in volume from 30 to 40 m³. There were two types of enclosures: some with natural sediment as a bottom and others with a plastic bottom. The experiment was a 2 × 2 factorial design with presence of sediment and nutrient enrichment as treatment factors. Both the sediment presence and nutrient enrichment significantly increased water nutrient concentrations and the rate of primary production. However, external nutrient enrichment and the presence of sediment stimulated the growth of different phytoplankton groups, indicating that the effect of sediment was not related to nutrient fluxes alone, but involved more complex interactions. External nutrient enrichment was primarily channelled to picoplanktonic cyanobacteria, the biomass of which increased four- to fivefold due to enrichment. The presence of sediment increased the biomass of cryptophytes, chrysophytes and prasinophytes, but decreased the biomass of N₂-fixing cyanobacteria. Zooplankton biomass increased during the experiment, but was not affected by the treatments. The study shows that sediment plays a significant role in phytoplankton dynamics, underlining the importance of including sediment in shallow-water mesocosm experiments. Handling editor: J. Padisak     

Microbial community structure in polluted Baltic Sea sediments

Nearly half the seabed of the Baltic Proper is incapable of supporting life of higher organisms as a consequence of oxygen depletion resulting from eutrophication. However, these areas are actually teeming with microbial life. Here we used terminal-restriction fragment length polymorphism (T-RFLP) to investigate the dominant archaeal and bacterial groups, with respect to community structure, in surface layers of bottom sediments of the Baltic Sea along a coastal pollution gradient. Both archaeal and bacterial communities formed distinct clusters along the pollution gradient and the community compositions were different at the polluted sites compared with the relatively clean reference sites. The structures of the bacterial communities were most strongly correlated to water depth, followed by organic carbon, oxygen, salinity and silicate levels. In contrast, the structures of the archaeal communities were most strongly correlated to oxygen, salinity, organic carbon, silicate and nitrate levels. Some members of the microbial communities were identified using a combination of traditional and molecular approaches. Isolates obtained on different culture media were identified by partial sequencing of their 16S rRNA genes and some novel species were found. In addition, we developed a computer program, aplaus, to elucidate the putative identities of the most dominant community members by T-RFLP.     

Seasonal variability in biomarkers in the bivalves Mytilus edulis and Macoma balthica from the northern Baltic Sea

Metallothionein level (MT), and acetylcholinesterase (AChE), catalase (CAT) and glutathione-S-transferase (GST) enzyme activities in the bivalves Mytilus edulis and Macoma balthica were investigated for seasonal variations from an inshore and an offshore site in the northern Baltic Sea. All the biomarkers showed variability, following mostly a similar pattern at both sites. Relationships between biomarkers and environmental factors and protein concentration and weight of target tissues were examined. In M. edulis, GST activity was related to Secchi depth, while in M. balthica a correlation with near-bottom oxygen saturation was observed. AChE activity correlated with the weight of the foot tissue of M. balthica. In both species, an integrated biomarker index indicated a stressed condition during the spring/early summer period. Strong seasonal variability in temperature and a concentrated period of food availability in spring-both governing the reproductive cycle of the bivalves-probably explains most of the observed natural variability in biomarkers in this sea area.     

Effect of nutrient enrichment on bacterioplankton biomass and community composition in mesocosms in the Archipelago Sea, northern Baltic

Effects of nutrient enrichment on the biomass and communitycomposition of heterotrophic bacteria and picocyanobacteriawere studied in large (42 m³) mesocosms in the brackish-waterArchipelago Sea (Baltic Sea) in late summer 2000 using cellcounts and denaturing gradient gel electrophoresis (DGGE) ofpolymerase chain reaction (PCR)-amplified 16S rRNA gene fragments.The identity of the major DNA bands was determined by sequencing.The obtained sequences were related to - and -proteobacteria,actinobacteria, verrucomicrobia and cyanobacteria. Nitrogenand phosphorus additions increased the biomasses of heterotrophicbacteria and picocyanobacteria and caused significant changesin their community composition judging from the DGGE bandingpatterns. Most verrucomicrobial bands had their highest relativeintensity in the control treatment and their lowest in the highernutrient addition treatment, whereas most Synechococcus-relatedbands had their lowest relative intensity in the lower nutrientaddition treatment. The responses of proteobacteria and actinobacteriawere more variable. The presence of both freshwater and marinesequences among the closest relatives to our sequences highlightsthe intermediate character of the Archipelago Sea between afreshwater and truly marine environment.     

Seasonal variability in the individual carbon content of the calanoid copepod Acartia bifilosa from the northern Baltic Sea

The seasonal variation in the carbon content of the calanoid copepod Acartia bifilosa was studied in the northern Baltic Sea. Monthly length-weight relationships were followed from November 1990 to October 1991 by analysing the organic carbon content of individuals, using a high temperature combustion method. The monthly length (L) on carbon (C) regressions of copepodites were best described by power functions (C = aL^b), where 75% to 88% of the variation could be explained by length. Nauplii length explained less of their carbon content (63–71 %). The carbon content per length was highest in summer when the nutritional situation was good and temperature at its highest. However, the carbon-length regressions differed significantly between months, except in winter, when two subsequent months had similar regressions. The results pointed out the importance of seasonal variation as well as the risk of making errors if biomass estimates based on length on carbon regressions are used. Carbon analysis should, if possible, be done on every sample or should at least be tested as to whether the regressions are consistent with the study material before estimations are calculated.     

Spatio-temporal dynamics of chlorophyll in the open Baltic Sea

Five charts of the chlorophyll and hydrographic fields by verticalprofiles of in situ fluorescence and CTD were made on a stationarygrid of 20 ? 25 nautical miles with a 5-mile spacing in theopen Baltic Sea. Both chlorophyll levels and variability weremaximal close to the spring bloom. High chlorophyll levels insummer are sustained by recurrent nutrient injections from thedeep saline layer. Two of the surveys showed close couplingbetween the coarse-scale (10 km) chlorophyll distribution andthe hydrographic structure determining the intensity of nutrienttransfer. Vigorous advection, stirring and current shear, associatedwith a strong mesoscale eddy, probably dominated the chlorophyllpattern on three surveys. The upward velocities in the cycloniceddy resulted in accumulation of phytoplankton in the aphoticzone. Intensive heat input from the surface caused a suddensinking of the phytoplankton and the formation of a pronouncedsub-surface chlorophyll maximum.     

Linking denitrifier community structure and prevalent biogeochemical parameters in the pelagial of the central Baltic Proper (Baltic Sea)

The oxic-anoxic interface of the water column of the Gotland Basin (central Baltic Sea) is characterised by defined biogeochemical gradients and is hypothesised to be a zone of pronounced denitrification. Our aim was to analyse the composition and distribution of pelagic denitrifying microorganisms in relation to the physico-chemical gradients in the water column. PCR-amplified nirS genes--coding for dissimilatory nitrite reductase--were analysed as functional markers by terminal restriction fragment length polymorphism and cloning. The overall nirS diversity was low, with the lowest levels found at the oxic-anoxic interface. Only a few terminal restriction fragments dominated the denitrifier communities throughout the water column, and these could be assigned to several new Baltic Sea clusters that were revealed by phylogenetic analysis. The novel clusters were separated in two groups corresponding to the oxygen concentrations within specific layers of the water column. Gradients of prevalent biogeochemical parameters (H(2)S, NH(4) (+), NO(3) (-) and O(2)) largely determined the composition of the nirS-type denitrifier communities within the water column of the Gotland Basin.     

Subsidies to predators, apparent competition and the phylogenetic structure of prey communities

Ecosystems are fragmented by natural and anthropogenic processes that affect organism movement and ecosystem dynamics. When a fragmentation restricts predator but not prey movement, then the prey produced on one side of an ecosystem edge can subsidize predators on the other side. When prey flux is high, predator density on the receiving side increases above that possible by in situ prey productivity, and when low, the formerly subsidized predators can impose strong top-down control of in situ prey—in situ prey experience apparent competition from the subsidy. If predators feed on some evolutionary clades of in situ prey over others, then subsidy-derived apparent competition will induce phylogenetic structure in prey composition. Dams fragment the serial nature of river ecosystems by prohibiting movement of organisms and restricting flowing water. In the river tailwater just below a large central Mexican dam, fish density was high and fish gorged on reservoir-derived zooplankton. When the dam was closed, water flow and the zooplankton subsidy ceased, densely packed pools of fish formed, fish switched to feed on in situ prey, and the tailwater macroinvertebrate community was phylogenetic structured. We derived expectations of structure from trait-based community assembly models based on macroinvertebrate body size, tolerance to anthropogenic disturbance, and fish-diet selectivity. The diet-selectivity model best fit the observed tailwater phylogenetic structure. Thus, apparent competition from subsidies phylogenetically structures prey communities, and serial variation in phylogenetic community structure can be indicative of fragmentation in formerly continuous ecosystems.     

Spatial structure of eukaryotic ultraplankton community in the northern South China Sea

Spatial distribution, diversity and composition of eukaryotic ultraplankton community of the northern South China Sea (nSCS) surface water and the relationship with the in situ water environment were investigated by the method of polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE). A total of 18 DGGE intensive bands were detected and the sequence analysis of these DGGE bands revealed that Alveolata was the dominant eukaryotic ultraplankton group of surface water in the nSCS (50%). Other species belonged to Bicoecea, Bolidophyceae, Polycystinea and Chlorophyta, which accounted for less proportion of eukaryotic ultraplankton in the study area. Unweighted pair group method with arithmetic averages clustering of the sampling stations indicated that all stations were classified mainly based on geographical proximity. Redundancy analysis (RDA) was employed to further investigate the relationships between DGGE band pattern and the environmental variables. Based on the RDA analysis, temperature, salinity, phosphorus and silicate were the important factors to shape the eukaryotic ultraplankton community composition in the nSCS.     

Selective predation by herring and mysids, and zooplankton community structure in a Baltic Sea coastal area

In lakes, fish and invertebrate predation are recognized asstrong structuring forces on zooplankton communities. The objectof this investigation was to study whether predation has a similarpotential in a coastal area of the brackish Baltic Sea and ifit could explain observed differences in zooplankton communitystructure between a reference area and an eutrophied area. Speciescomposition and daily vertical migration of zooplankton andzooplanktivores, and the diets of the latter, were studied inJuly and August at two 30 m deep stations differing in primaryproductivity. The biomass of zooplankton >35 µm wasdominated by copepods, but cladocerans and rotifers also occurredin significant numbers. The dominating zooplanktivores wereherring (Clupea harengus) and the mysid shrimp Mysis mixta.They fed almost exclusively on zooplankton, mainly copepods,and their estimated food consumption equalled or exceeded thesummer copepodite production. The structure of the zooplanktoncommunity cannot be explained by effects of predation or byfeeding conditions alone. Increases in rotifer and cladoceranabundances with increased primary production suggests effectsof food supply. However, a generally rapid decline in the annualsummer peak of cladocerans may be caused by predation. The totalabundance of copepods did not increase with improved feedingconditions, but there was a shift in species dominance. Thecopepod Ewytemora affinis hirundoides, which was intensivelypreyed upon, increased with increased phytoplankton production,while Acartia bifilosa and/or A.longiremis, which was less eaten,decreased. Predation may explain a pronounced daily verticalmigration of the most predated copepods. They occurred in deeperwater during the day, when the visually feeding herring wereactive, and moved closer to the surface at night when M.mixtaleft the bottom, to forage in the water column.     

Ecological balance between ciliate plankton and its prey candidates,pico- and nanoplankton,in the East China Sea

Standing stocks of ciliate plankton and its prey candidates, both picoplankton and nanoplankton, were investigated in spring in the East China Sea. The former was 1.36 × 10⁵–1.54 × 10⁸ μm³ l⁻¹ in biovolume, and the latter was 5.33 × 10⁶–1.11 × 10⁸ μm³ l⁻¹. The biovolume ratio of ciliate plankton to prey candidates ranged from 1.31 × 10⁻² to 2.00 × 10⁰; it was larger in abundant prey conditions and smaller in sparse preys. Making some plausible assumptions about physiological activity on both organisms, every ratio meet the quantitative restriction that prey production should be equal to or larger than ciliate consumption. However, prey candidates would be so sparsely distributed that ciliate plankton could not capture sufficient prey organisms in its random filter-feeding manner. Even though planktonic ciliates must have some extraordinary mechanisms to capture preys efficiently, this quantitative imbalance might be one of the reasons for decreasing ciliate/prey ratio in sparse prey conditions. Handling editor: K. Martens     

Polychaetes associated with the sciaphilic alga community in the northern Aegean Sea: spatial and temporal variability

Polychaete biodiversity has received little attention despite its importance in biomonitoring. This study describes polychaete diversity, and its spatial and temporal variability in infralittoral, hard substrate assemblages. Seven stations were chosen in the central area of the northern Aegean Sea. At each station, one to three depth levels were set (15, 30 and 40 m). Five replicates were collected by scuba diving with a quadrat sampler (400 cm²) from each station and depth level during summer for the spatial analysis, and seasonally for the study of temporal changes. Common biocoenotic methods were employed (estimation of numerical abundance, mean dominance, frequency, Margalefs richness, Shannon-Weaver index and Pielous evenness). A total of 5,494 individuals, belonging to 79 species, were counted and classified. Diversity indices were always high. Clustering and multidimensional scaling techniques indicated a high heterogeneity of the stations, although these were all characterized by the sciaphilic alga community. A clear seasonal pattern was not detectable. Summer and autumn samples discriminate, while winter and spring form an even group. The abundance/biomass comparison indicated a dominance of k-strategy patterns, characteristic of stable communities.Communicated by H.-D. Franke     

Reproductive characteristics of the male herring in the northern Baltic Sea

The gonad weight and gonadosomatic index of the male Baltic herring Clupea harengus membras L., were highest at the beginning of the reproductive season (April/May), the values decreasing towards the end of it (July/August) during 1988–1991. The decline could not be explained by fish size but may have been due to fish condition. A high individual variation was typical for both gonad weights and gonadosomatic indices in fish of the same size and maturity stage. The mean density of sperm cells was significantly higher in June (34·9 × 10⁹ ml⁻¹) than in July (19·2 × 10⁹ ml⁻¹, Mann-Whitney U= 17; P<0·05), the variation among the males being high in both groups. Electron microscope analysis showed a severe disruption of the mitochondrial elements in males spawning at 22°C.     

Seasonal variability in the grazing potential of the invasive amphipod Gammarus tigrinus and the native amphipod Gammarus salinus (Amphipoda: Crustacea) in the northern Baltic Sea

Mesograzers are known to reduce the biomass of their host plant and modify the structure of the whole macrophyte community in many ecosystems. Thus, the introduction of an efficient mesograzer may destabilize macrophyte community and also affect the native grazers. We estimated how large proportion of macrophyte production are consumed by the alien gammarid G. tigrinus and the native gammarid G. salinus in the species poor ecosystem of the northern Baltic Sea. We analysed whether G. tigrinus consumes different diet as the native G. salinus and whether the effect of G. tigrinus on the survival of the native G. salinus is macrophyte species specific. Grazing experiments showed that there was a clear difference in the grazing rates of gammarids among the studied macrophyte species in summer and autumn but not in spring. The grazing rates were significantly higher in the prevailing macrophyte Pilayella littoralis as compared to other macrophytes. The grazing was inversely related to the diurnal net photosynthetic values of macrophytes. The gammarid amphipods potentially removed only a minor part of plant primary production except for summer and autumn when grazing of a few perennial species exceeded macrophyte production. Macrophyte species and presence of G. salinus had no effect on the survival of G. tigrinus. The presence of G. tigrinus, however, reduced the survival of the native gammarids within P. littoralis in summer. To conclude it is likely that both native and alien gammarid amphipods do not exert significant pressure on the macroalgal communities in the northern Baltic Sea. Competitive interactions between G. tigrinus and G. salinus within the prevailing macrophyte P. littoralis is the likely explanation of the decline of the native gammarid amphipods after the establishment of G. tigrinus in the northern Baltic Sea.     

Habitat and infauna prey availability for flatfishes in the northern Bering Sea

Yellowfin sole (Limanda aspera), northern rock sole (Lepidopsetta polyxystra), and Alaska plaice (Pleuronectes quadrituberculatus) are valuable flatfishes in the southeastern Bering Sea (EBS) bottom trawl fishery. The northern Bering Sea (NBS) is near their northern distribution limit. We conducted the first assessment of NBS habitat suitability for these benthivorous flatfishes from the perspective of prey availability in 2010. Benthic samples were collected at 12 trawl stations along a meridional transect extending from 60.5°N to 64.5°N east of St. Lawrence Island to characterize the prey environment. Stomach contents from the flatfishes were concomitantly collected to relate diets to prey fields. The diet compositions did not correspond spatially with the infauna communities. The flatfishes elected a prey group regardless of its relative availability. The spatial mismatch between diet and infauna compositions suggests that prey availability was high in the NBS. The flatfishes generally have versatile diets, but they were more selective of their prey here than in the EBS. The biomass and the abundance of the infauna along the transect were comparable with the EBS. Although niche overlap was high in the NBS, competition for food was likely lower than in the EBS because of the lower density of flatfish. The bottom temperatures in the NBS were in the same range as the EBS during the summer of 2010. The NBS appears to be suitable flatfish habitat at least during the summer ice-free period. The effects of climate warming on the prey environment and on the production and distribution of flatfish are complex and difficult to predict, but if the NBS were to shift over time toward milder winter conditions that more resemble the EBS, its suitability as flatfish habitat would presumably increase based on the present prey availability.     

Corynosoma acanthocephalans in their paratenic fish hosts in the northern Baltic Sea

Cystacanth stages of three Corynosoma (Acanthocephala) species, C. strumosum and C. semerme, and a new species in the Bothnian Bay, C. magdaleni, were studied in the fourhorn sculpin (Myoxocephalus quadricornis) in 1996-1997. The length of the trunk and proboscis differentiated the three species in the fish. The stability of the Corynosoma infection was studied by comparing the present results with those collected from the same areas (central and coastal) in 1977-1982 (Valtonen, 1983a). As C. magdaleni and C. strumosum were not separated at that time, the joint infections of these two species (called "C. strumosum") were compared. The stability of infection with C. semerme in the central Bothnian Bay was noticeable (prevalences were 82.9 in earlier period and 81.9% in later period), while the prevalences of "C. strumosum" in the same areas had decreased (21 and 13.5%, respectively). This is suggested to be due to the disappearance of the marine bull-rout, Myoxocephalus scorpius, from the Bothnian Bay during the 1990s due to an overall decline in salinity throughout the Baltic Sea. In the coastal area Corynosoma infection was clearly lower than in central area in both periods.