Evidence for active microbial nitrogen transformations in sea ice (Gulf of Bothnia, Baltic Sea) in midwinter

Nutrient concentrations, chlorophyll-a, bacterial biomass and relative activity of denitrifying organisms were investigated from ice-core, brine and underlying water samples in February 1998 in the Gulf of Bothnia, Baltic Sea. Examined sea ice was typical for the Baltic Sea; ice bulk salinity varied from 0.1 to 1.6 psu, and in underlying water salinity was from 4.2 to 4.7 psu. In 2- to 3-months-old sea ice (thickness 0.4–0.6 m), sea-ice communities were at the winter stage; chl-a concentrations were generally below 1 mg m⁻³ and heterotrophic organisms composed 7–20% of organism assemblage. In 1-month-old ice (thickness 0.2–0.25 m), an ice spring bloom was already developing and chl-a concentrations were up to 5.6 mg m⁻³. In relation to low salinity, high concentrations of NH⁺ ₄, NO⁻ ₂, PO³⁺ ₄ and SiOH₄ were found in the ice column. The results suggest that the upper part of ice accumulates atmospheric nutrient load during the ice season, and nutrients in the upper 10–20 cm of ice are mainly of atmospheric origin. The most important biological processes controlling the sea-ice nutrient status are nutrient regeneration, nutrient uptake and nitrogen transformations. Nutrient regeneration is specially active in the middle parts of the 50- to 60-cm-thick ice and subsequent accumulation of nutrients probably enhances the ice spring bloom. Nitrite accumulation and denitrifying activity were located in the same ice layers with nutrient regeneration, which together with the observed significant correlation between the concentrations of nitrogenous nutrients points to active nitrogen transformations occurring in the interior layers of sea ice in the Baltic Sea. Accepted: 12 June 2000     

Abiotic factors influencing cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Blooms of cyanobacteria are a recurrent phenomenon in the Baltic Sea, including the Gulf of Finland. The spatial extension, duration, intensity and species composition of these blooms varies widely between years. Alg@line data collected regularly from ferries as well as weather service and marine monitoring data from 1997 to 2005 are analysed to determine the main abiotic factors influencing the intensity and species composition of cyanobacterial blooms in the Gulf of Finland. It is demonstrated that the development of the Nodularia spumigena Mertens bloom is highly dependent on weather conditions such as photosynthetically active radiation and water temperature. Nutrient conditions, especially the surplus of phosphorus (according to Redfield ratio) related to the pre-bloom upwelling events in the Gulf, affect the intensity of Aphanizomenon sp. (L.) Ralfs blooms. Differences in bloom timing and duration indicate that, if the preconditions (like nutrient ratio/concentration and weather conditions) for bloom formation are favourable, then the Aphanizomenon bloom starts earlier, the overall bloom period is longer and the Nodularia peak might appear in a wider time window. Handling editor: K. Martens     

Diversity of Oil-Degrading Microorganisms in the Gulf of Finland (Baltic Sea) in Spring and in Summer

Diversity of the oil-degrading microbial strains isolated from the water and sediments of the Gulf of Finland (Baltic Sea) in winter and in summer was studied. Substrate specificity of the isolates for aliphatic and aromatic hydrocarbons was studied. The isolates belonged to 32 genera of the types Proteobacteria (alpha-, beta-, and gammaproteobacteria), Actinobacteria,Firmicutes, and Bacteroidetes. Seasonal variations of the oil-degrading microbial communities was revealed. The presence of the known genes responsible for the degradation of oil aliphatic and aromatic hydrocarbons was determined. The alkB sequence of the alkane hydroxylase gene was found in ~16% of the studied strains. The sequence of the phnAc phenanthrene 3,4- dioxygenase was found in Sphingobacterium sp. and Arthrobacter sp. isolates retrieved in winter and summer. In five Pseudomonas sp. strains from winter samples, the classical operons of naphthalene degradation (nah) were localized in catabolic plasmids, of which three belonged to IncР-9, one, to IncР-7, and two to an unidentified incompatibility group. Burkholderia and Delftia strains contained the operons for naphthalene degradation via salicylate and gentisate (nag). The presence of nag genes has not been previously reported for Delftia spp. strains. The sequences of the nagG salicylate 5-hydroxylase gene were also found in Achromobacter, Sphingobacterium, and Stenotrophomonas strains.     

Effects of nonindigenous species on diversity and community functioning in the eastern Gulf of Finland (Baltic Sea)

An increase of xenodiversity in plankton and benthos in the eastern Gulf of Finland was observed from 1998 to 2004. Nonindigenous species account for 4.8% of all species found and up to 96% of total biomass. Invasive benthic omnivores, the alien amphipods Gmelinoides fasciatus and Pontogammarus robustoides and the predaceous fish Perccottus glenii with their versatile diets strongly affect the community structure. Invasive sessile seston-feeders that directly (through grazing and water clearance) and indirectly (through recycling of nutrients) interact with other ecosystem components, are mainly represented by the zebra mussel Dreissena polymorpha, which affect the structure of benthic and planktonic communities as well as benthic-pelagic coupling. The invasive predatory cladocerans Cercopagis pengoi and Evadne anonyx and larvae of D. polymorpha are only temporary components in the zooplankton, which is limiting their overall effect. Alien benthic bioturbators, the polychaetes Marenzelleria neglecta and the oligochaete Tubificoides pseudogaster account for a high proportion of total abundance and biomass but their effects on native species need further research.     

Binding of phosphate in sediment accumulation areas of the eastern Gulf of Finland,Baltic Sea

The relationships between P and components binding P were studied by analysing the concentrations of N, P, Fe, Mn, Ca and Al in sediments and pore water along the estuarine transect of the River Neva in August 1995. The high sediment organic matter concentration resulted in low surface redox potential and high pore-water o-P concentration, whereas the abundance of amphipods resulted in high surface redox potentials and low pore-water o-P concentration. However, despite the variation in sediment organic matter and the abundance of amphipods, very reduced conditions and slightly variable concentrations of Tot-P (0.7–1.1 mg g^–1 DW) were observed in the 10–15 cm sediment depth along the estuarine gradient, indicating that the pools of mobile P were largely depleted within the depth of 0–15 cm. Multiple regression analysis demonstrated that organic matter and Tot-Fe concentration of the sediment were closely related to the variation in Tot-P concentration of the sediments (r ² = 0.817, n=32). In addition, the high total Fe:P ratio suggested that there is enough Fe to bind P in sediments along the estuarine gradient. However, low Fe_diss concentrations in the pore water of reduced sediment (redox-potential <–50 mV) indicated efficient precipitation of FeS (FeS and FeS₂), incapable to efficiently bind P. Consequently, the low Fe_diss:o-P ratio (< 1) recorded in pore water in late summer implied that Fe³⁺ oxides formed by diffusing Fe_diss in the oxic zone of the sediments were insufficient to bind the diffusing o-P completely. The measured high o-P concentrations in the near-bottom water are consistent with this conclusion. However, there was enough Fe_diss in pore water to form Fe³⁺ oxides to bind upwards diffusing P in the oxic sediment layer of the innermost Neva estuary and the areas bioturbated by abundant amphipods.     

The influence of weather conditions (temperature and wind) on cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Cyanobacterial blooms are common in the Baltic Sea. They are dominated by Aphanizomenon flos-aquae and Nodularia spumigena and take place in July–August. Investigations of bloom development using different approaches have been carried out in the Gulf of Finland during recent years. The ship-of-opportunity technique allows to observe the upper layer dynamics from meso- to basin-wide scale with high temporal and spatial frequency at low cost. Unattended measurements on board a commercial ferry along a transect between Tallinn and Helsinki have been conducted for 3 years (1997–1999). The influence of weather conditions—temperature and wind—on the cyanobacterial bloom development was investigated. The formation of cyanobacterial blooms was favoured by warm and calm weather, while in cold and windy conditions other species formed mass occurrences. Water temperature has been found to be the main factor controlling the initiation of the bloom, in general, while vertical stratification appeared to be the critical factor determining the intensity of the bloom at species level. The spatial distribution of the cyanobacterial bloom was determined rather by the wind-forced advection than by the possible vertical transport of nutrients in the areas of the observed upwelling events.     

Seasonal and short-term variation in denitrification and anammox at a coastal station on the Gulf of Finland,Baltic Sea

Benthic processes were measured at a coastal deposition area in the northern Baltic Sea, covering all seasons. The N₂ production rates, 90–400 μmol N m⁻² d⁻¹, were highest in autumn-early winter and lowest in spring. Heterotrophic bacterial production peaked unexpectedly late in the year, indicating that in addition to the temperature, the availability of carbon compounds suitable for the heterotrophic bacteria also plays a major role in regulating the denitrification rate. Anaerobic ammonium oxidation (anammox) was measured in spring and autumn and contributed 10% and 15%, respectively, to the total N₂ production. The low percentage did, however, result in a significant error in the total N₂ production rate estimate, calculated using the isotope pairing technique. Anammox must be taken into account in the Gulf of Finland in future sediment nitrogen cycling research. Handling editor: J. Cole     

Distribution,population structure and salinity tolerance of the invasive amphipod Gmelinoides fasciatus (Stebbing) in the Neva Estuary (Gulf of Finland,Baltic Sea)

In the early 1970s, the Baikalian amphipod Gmelinoides fasciatus (Stebbing) was intentionally introduced into several lakes in the Gulf of Finland basin in order to enhance fish production. By 1996, G. fasciatus successfully colonized the littoral zone of Lake Ladoga and, via the Neva River, invaded the Neva Bay, the freshwater part of the Neva Estuary. In 1999, G. fasciatus was first registered in the inner Neva Estuary, the very first record of the Baikalian amphipod in brackish waters of the Baltic Sea. Distribution, abundance, reproduction and population structure of G. fasciatus in the Neva Estuary were studied during 1998–2000. In some locations of the Neva Estuary, maximum densities of G. fasciatus reached 3500 ind. m⁻². In general, density and biomass of G. fasciatus in the freshwater part of the Neva Estuary were higher (around 1.5 fold) than in the brackish-water part. Fecundity of this amphipod averaged 10–20 eggs per female, depending on body size of females and season. In order to assess the possibility of further spread of G. fasciatus in the Baltic Sea, the salinity tolerance of this species was determined in a series of laboratory experiments. Our results showed that the invasive amphipod G. fasciatus is potentially able to colonize shallow coastal habitats of, for example, the Gulf of Bothnia, Gulf of Riga and other parts of the Baltic Sea with water salinities ranging from 1 to 5 psu.

     

Spring time production of bottom ice algae in the landfast sea ice zone at Barrow, Alaska

The primary production of bottom ice algae is an important food source for sympagic, pelagic and benthic organisms in the Arctic Ocean as well as Antarctic Ocean. Using ¹³C-¹⁵N isotope tracers, the recent ice algal production at Barrow during the spring season was lower in 2003 than three decades ago, although the maximum chlorophyll-a concentration for the bottom ice algae was similar to the values from previous studies. Estimated recent new and total production rates of the ice algae were 0.8 g C m^- 2 yr^- 1 and 2.0 g C m^- 2 yr^- 1 respectively, while the rates of water column phytoplankton were 0.2 g C m^- 2 yr^- 1 and 0.7 g C m^- 2 yr^- 1 for the spring sampling period in 2003. The ice algae contributed 74% of the pelagic primary production under the landfast sea ice at Barrow before the phytoplankton spring bloom. At the end of the season in 2003, a high carbon allocation of lipids in the ice algae was found. Three possible explanations- nutrient depletion, increasing light, and/or changes in species composition- were suggested for the high carbon incorporation into lipids. This high lipid synthesis of the bottom ice algae might be significant to zooplankton and benthic fauna grazers because lipids are the most energy dense biomolecules.     

Further observations on flagellates within sea ice in northern Bothnian Bay, the Baltic Sea

To improve our knowledge of flagellates inhabiting the Baltic Sea ice and water column during the winter, material was obtained from northern Bothnian Bay in March/April 1995. Light microscopical observations on live and fixed material and further transmission electron microscopy of whole mounts revealed 47 nanoflagellate taxa. In addition, detached scales of eight taxa were encountered. It is now evident that nearly all nanoflagellate classes are present within Bothnian Bay sea ice. The most common groups were cryptomonads, dinoflagellates, chrysophytes, prasinophytes, choanoflagellates and heterotrophic flagellates of unknown systematic position (Protista incertae sedis). Most flagellates in Baltic Sea ice biota apparently thrive in both the water column and the sea ice, while some, e.g. Paraphysomonas spp. (Chrysophyceae), heterotrophic euglenids, volvocalean chlorophytes, and some taxa of uncertain systematic affinity, are more frequently found within the sea ice. Received: 9 February 1997 / Accepted: 23 September 1997     

European smelt Osmerus eperlanus in the eastern Gulf of Finland,Baltic Sea: Stock status and fishery

In the eastern Gulf of Finland, European smelt Osmerus eperlanus occurs as an anadromous ecological form that spawns in coastal, low-salinity zones and in several rivers entering the gulf, most importantly in the River Neva. Osmerus eperlanus is a key commercial fish species for the population of St. Petersburg, and the city's fish symbol. However, the state of the smelt stock has considerably deteriorated over the past few decades. Monitoring shows that annual catches during 1965–1992 averaged 2274 t, compared to 292 t in 2002–2017, although some improvement is inidicated by a gradual increase in catches in recent years (e.g., to 595 t in 2017). This paper addresses long-term and recent changes in the fisheries for smelt in the eastern Gulf of Finland, including both commercial and recreational fisheries, and the range of fishing gears used. The commercial smelt fishery is mainly conducted during the spawning period (April–May) on migrating fish, using trap nets of various designs and beach seines. There has been an increase in recreational fisheries for smelt; in years when ice cover on the gulf during winter (December–April) is steady, recreational fishers from St. Petersburg, using hook and line, may catch quantities that are comparable to those of commercial catches. Several factors may account for the stock dynamics and associated changes in catches of smelt: these include a decrease in prey availability in the gulf; loss and degradation of spawning and nursery habitats, partly associated with large-scale hydroengineering operations in Neva Bay; and illegal, unreported and unregulated (IUU) fishing which became widespread in post-Soviet decades. In spite of a gradual increase in smelt abundance in the most recent 15 years, a failure to accurately monitor and manage the extent of IUU and recreational fishing for smelt, to assess the status of the stock, or to forecast catches, increases the risk of overfishing of the spawning stock.     

Community development and modes of phosphorus utilization in a late summer ecosystem in the central Gulf of Finland, the Baltic Sea

The development of a filamentous, nitrogen-fixing cyanobacterial bloom was followed during July–August 1990 in a stratified basin in the central Gulf of Finland, Baltic Sea. Hydrography, dissolved inorganic, particulate and total nutrients, chlorophyll a, alkaline phosphatase activity, ³²PO₄-uptake and phytoplankton species were measured. The study period was characterized by wind-induced mixing events, followed by marked nutrient pulses and plankton community responses. Phosphate uptake was highest throughout the study period in the size fraction dominated by bacteria and picocyanobacteria (< 2 µm) and the proportion of uptake in the size fraction 2–10 µm remained low (2–6%). Higher phosphate turnover times were observed in a community showing signs of enhanced heterotrophic activity. The bloom of filamentous, nitrogen-fixing cyanobacteria Aphanizomenon flos-aquae was promoted by a nutrient pulse with an inorganic nutrient ratio (DIN:DIP) of 15. The results show that the quality, frequency and magnitude of the physically forced nutrient pulses have an important role in determining the relative share of the different modes of phosphorus utilization and hence in determining the cyanobacterial bloom intensity and species composition in the Baltic Sea.     

Blue mussels (Mytilus edulis) in the diet of roach (Rutilus rutilus) in outer archipelago areas of the western Gulf of Finland,Baltic Sea

The blue mussel (Mytilus edulis) is one of the key species in the Baltic Sea ecosystem and it is living at the edge of its range in the western Gulf of Finland. Roach (Rutilus rutilus) is a freshwater fish species that has benefited from recent coastal eutrophication and is at present highly abundant in the outer archipelago of the Gulf of Finland. In 2000 and 2001, a total of 516 roach were sampled for diet analysis in three study areas. Shelled molluscs formed over 95% of the diet of roach, blue mussels being the dominant single species. The proportion of this species in the food of roach in the three study areas ranged between 38–61% for smaller roach (<225 mm) and 39–85% for larger (>225 mm) roach, indicating that blue mussel is a highly important food source for roach in outer archipelago areas of the western Gulf of Finland, in contrast to reports from other parts of the northern Baltic Sea. The scarcity of large blue mussels in mussel beds in the easternmost study area was reflected in the lower proportion of blue mussels in the diet of larger roach. However, the growth of roach was not affected by the availability of blue mussels. The twofold differences observed in the annual growth of roach between warm and cold years demonstrated that temperature is an important factor controlling the growth of roach in the western Gulf of Finland.     

Genetic diversity in strains of the genus Anabaena isolated from planktonic and benthic habitats of the Gulf of Finland (Baltic Sea)

Late summer cyanobacterial blooms in the Baltic Sea contain Anabaena sp. together with Nodularia spumigena and Aphanizomenon flos-aquae. Although Anabaena is common especially in the Gulf of Finland, very little is known about its genetic diversity. Here we undertook a molecular phylogenetic study of 68 Anabaena strains isolated from the brackish Gulf of Finland. We sequenced the 16S rRNA genes from 54 planktonic and 14 benthic Anabaena strains, and rbcL and rpoC1 genes from a subset of these strains. Phylogenetic trees showed that Anabaena strains, from both planktonic and benthic habitats, were genetically diverse. Although the Anabaena strains were morphologically diverse, in our study only one genetically valid species was found to exist in the plankton. Evolutionary distances between benthic Anabaena strains were greater than between planktonic strains, suggesting that benthic habitats allow for the maintenance of greater genetic diversity than planktonic habitats. A number of novel lineages containing only sequences obtained in this study were compiled in the phylogenetical analyses. Thus, it seemed that novel lineages of the genus Anabaena may be present in the Baltic Sea. Our results demonstrate that the Baltic Sea Anabaena strains show surprisingly high genetic diversity.     

Long-Term and Seasonal Dynamics of the Population of Cercopagis pengoi (Ostroumov 1891) (Cladocera,Onychopoda) in the Eastern Part of the Gulf of Finland,Baltic Sea

Biology Bulletin - Long-term seasonal studies of the population of Cercopagis pengoi (Ostroumov 1891) (Cladocera, Onychopoda) in the eastern Gulf of Finland were carried out for the first time....     

Drift-ice and under-ice water communities in the Gulf of Bothnia (Baltic Sea)

The algal, protozoan and metazoan communities within different drift-ice types (newly formed, pancake and rafted ice) and in under-ice water were studied in the Gulf of Bothnia in March 2006. In ice, diatoms together with unidentified flagellates dominated the algal biomass (226 ± 154 μg ww l⁻¹) and rotifers the metazoan and protozoan biomass (32 ± 25 μg ww l⁻¹). The under-ice water communities were dominated by flagellates and ciliates, which resulted in lower biomasses (97 ± 25 and 21 ± 14 μg ww l⁻¹, respectively). The under-ice water and newly formed ice separated from all other samples to their own cluster in hierarchical cluster analysis. The most important discriminating factors, according to discriminant analysis, were chlorophyll-a, phosphate and silicate. The under-ice water/newly formed ice cluster was characterized by high nutrient and low chlorophyll-a values, while the opposite held true for the ice cluster. Increasing trends in chlorophyll-a concentration and biomass were observed with increasing ice thickness. Within the thick ice columns (>40 cm), the highest chlorophyll-a concentrations (6.6–22.2 μg l⁻¹) were in the bottom layers indicating photoacclimation of the sympagic community. The ice algal biomass showed additional peaks in the centric diatom-dominated surface layers coinciding with the highest photosynthetic efficiencies [0.019–0.032 μg C (μg Chl-a ⁻¹ h⁻¹) (μE m⁻² s⁻¹)⁻¹] and maximum photosynthetic capacities [0.43-1.29 μg C (μg Chl-a ⁻¹ h⁻¹)]. Rafting and snow-ice formation, determined from thin sections and stable oxygen isotopic composition, strongly influenced the physical, chemical and biological properties of the ice. Snow-ice formation provided the surface layers with nutrients and possibly habitable space, which seemed to have favored centric diatoms in our study.     

Development of ice biota in a temperate sea area (Gulf of Bothnia)

A study of sea ice biota was carried out in the Gulf of Bothnia (northern Baltic Sea) during the winter of 1989–1990. Samples (ice cores) were taken at a coastal station at regular time intervals during the ice season. Chlorophyll a concentration, algal species distribution, bacterial numbers, and primary and bacterial production were measured. Colonization of the ice began in January when daylight was low. As the available light increased, the algae started to grow exponentially. The vertical chlorophyll a distribution changed and algal species composition and biomass changed during the season. During the initial and middle phase of colonization, ice-specific diatoms, Nitzschia frigida and Navicula pelagica, dominated the algal biomass. Nutrients (PO₄ ^3– and NO_3j) were found to be depleted during the time of algal exponential growth. The maximum algal biomass exceeded 800 g C 1^–1. The primary production supplied food for heterotrophic organisms. The presence of heterotrophic organisms of different trophic levels (bacteria, flagellates, ciliates and rotifers) indicated an active microbial food web.