Allelopathic effects of filamentous cyanobacteria on phytoplankton in the Baltic Sea

Eutrophication and enhanced internal nutrient loading of the Baltic Sea are most clearly reflected by increased late-summer cyanobacterial blooms, which often are toxic. In addition to their toxicity to animals, phytoplankton species can be allelopathic, which means that they produce chemicals that inhibit competing phytoplankton species. Such interspecific chemical warfare may lead to the formation of harmful phytoplankton blooms and the spread of exotic species into new habitats. This is the first report on allelopathic effects in brackish-water cyanobacteria. The experimental studies presented in this thesis showed that the filamentous cyanobacteria Anabaena sp., Aphanizomenon flos-aquae and Nodularia spumigena are capable of decreasing the growth of other phytoplankton species, especially cryptophytes, but also diatoms. The detected allelopathic effects are rather transitory, and some co-occurring species show tolerance to them. The allelochemicals are excreted during active growth and they decrease cell numbers, chlorophyll a content and carbon uptake of the target species. Although the more specific modes of action or chemical structures of the allelochemicals remain to be studied, the results clearly indicate that the allelopathic effects are not caused by the hepatotoxin, nodularin. On the other hand, cyanobacteria stimulated the growth of bacteria, other cyanobacteria, chlorophytes and flagellates in a natural phytoplankton community. In a long-term data analysis of phytoplankton abundances and hydrography of the northern Baltic Sea, a clear change was observed in phytoplankton community structure, together with a transition in environmental factors, between the late 1970s and early 2000s. Surface water salinity decreased, whereas water temperature and the concentration of dissolved inorganic nitrogen increased. In the phytoplankton community, the biomass of cyanobacteria, chrysophytes and chlorophytes significantly increased, and the late-summer phytoplankton community became increasingly cyanobacteria-dominated. In contrast, the biomass of cryptophytes decreased. The increased temperature and nutrient concentrations probably explain most of the changes in phytoplankton, but my results suggest that the possible effect of chemically mediated biological interactions should also be considered. Cyanobacterial allelochemicals can cause additional stress to other phytoplankton in the nutrient-depleted late-summer environment and thus contribute to the formation and persistence of long-lasting cyanobacterial mass occurrences. On the other hand, cyanobacterial blooms may either directly or indirectly promote the growth of some phytoplankton species. Therefore, a further increase in cyanobacteria will probably shape the late-summer pelagic phytoplankton community by stimulating some species, but inhibiting others.

The role of phosphorus as a regulator of bloom-forming diazotrophic cyanobacteria in the Baltic Sea

Eutrophication favours harmful algal blooms worldwide. The blooms cause toxic outbreaks and deteriorated recreational and aesthetic values, causing both economic loss and illness or death of humans and animals. The Baltic Sea is the world s only large brackish water habitat with recurrent blooms of toxic cyanobacteria capable of biological fixation of atmospheric nitrogen gas. Phosphorus is assumed to be the main limiting factor, along with temperature and light, for the growth of these cyanobacteria. This thesis evaluated the role of phosphorus nutrition as a regulating factor for the occurrence of nitrogen-fixing cyanobacteria blooms in the Baltic Sea, utilising experimental laboratory and field studies and surveys on varying spatial scales. Cellular phosphorus sources were found to be able to support substantial growth of the two main bloom forming species Aphanizomenon sp. and Nodularia spumigena. However, N. spumigena growth seemed independent of phosphorus source, whereas, Aphanizomenon sp. grew best in a phosphate enriched environment. Apparent discrepancies with field observations and experiments are explained by the typical seasonal temperature dependent development of Aphanizomenon sp. and N. spumigena biomass allowing the two species to store ambient pre-bloom excess phosphorus in different ways. Field experiments revealed natural cyanobacteria bloom communities to be predominantly phosphorus deficient during blooms. Phosphate additions were found to increase the accumulation of phosphorus relatively most in the planktonic size fraction dominated by the nitrogen-fixing cyanobacteria. Aphanizomenon sp. responded to phosphate additions whereas the phosphorus nutritive status of N. spumigena seemed independent of phosphate addition. The seasonal development of phosphorus deficiency is different for the two species with N. spumigena showing indications of phosphorus deficiency during a longer time period in the open sea. Coastal upwelling introduces phosphorus to the surface layer during nutrient deficient conditions in summer. The species-specific ability of Aphanizomenon sp. and N. spumigena to utilise phosphate enrichment of the surface layer caused by coastal upwelling was clarified. Typical bloom time vertical distributions of biomass maxima were found to render N. spumigena more susceptible to advection by surface currents caused by coastal upwellings. Aphanizomenon sp. populations residing in the seasonal thermocline were observed to be able to utilise the phosphate enrichment and a bloom was produced with a two to three week time lag subsequent to the relaxation of upwelling. Consistent high concentrations of dissolved inorganic phosphorus, caused by persistent internal loading of phosphorus, was found to be the main source of phosphorus for large-scale pelagic blooms. External loads were estimated to contribute with only a fraction of available phosphorus for open sea blooms. Remineralization of organic forms of phosphorus along with vertical mixing to the permanent halocline during winter set the level of available phosphorus for the next growth season. Events such as upwelling are important in replenishing phosphate concentrations during the nutrient deplete growth season. Autecological characteristics of the two main bloom forming species favour Aphanizomenon sp. populations in utilising the abundant excess phosphate concentrations and phosphate pulses mediated through upwelling. Whilst, N. spumigena displays predominant phosphorus limited growth mode and relies on more scarce cellular phosphorus stores and presumably dissolved organic phosphorus compounds for growth. The Baltic Sea is hypothesised to be in an inhibited state of recovery due to the extensive historical external nutrient loading, extensive internal phosphorus loading and the substantial nitrogen load caused by cyanobacteria nitrogen fixation. This state of the sea is characterised as a vicious circle .

Phosphorus and organic matter discharged by Finnish rivers to the Baltic Sea

Tiivistelmä: Suomen jokivesien Itämereen kuljettama fosfori ja orgaaninen aine

Optimal management of conflicting species: Grey seal (Halichoerus grypus) and Atlantic salmon (Salmo salar) in the Baltic Sea

The objective of this thesis is to examine the economic effects in the conflict between grey seal population and the salmon fishery in the Baltic Sea. We will formulate a bioeconomic model which provides new insights on the optimal management of Atlantic salmon with respect to the effects brought about by the grey seal population. As the catch losses caused by seals have an effect on salmon fishery in Baltic, we will study how seal population affects the present value of the salmon fishery. The study considers the Finnish coastal trap net fishery. The bioeconomic model considers a scenario of sole salmon fishery and a scenario of salmon fishery affected by the grey seal population. On the basis of these scenarios, a seal compensation scheme is introduced. We can observe a significant economic seal-induced effect on the salmon fishery. The results suggest that the present seal compensation scheme emploid by the Finnish government is suboptimal. This thesis is part of the TARMO –project, in which the conflict between grey seal population and salmon fishery is studied using the methods of environmental economics.

Baltic Sea : security tomorrow - three future scenarios

Aineisto on Keskustakampuksen kirjaston digitoimaa ja kirjasto vastaa aineiston käyttöluvista.

Host-parasitoid relationship in different Cotesia melitaearum and Melitaea cinxia populations around the Baltic Sea

The relationship between hosts and parasites is one of the most studied interactions between living organisms, and it is both universal and common in nature. Parasitoids are special type of parasites whose offspring develop attached to or within a single host organism that it ultimately consumes and kills. Hosts are arthropods and most parasitoids belong to the insect order Hymenoptera. For almost two decades metapopulation research on the Glanville fritillary butterfly (Melitaea cinxia) has been conducted in the Åland Islands, Finland. The studies have been concerned with the population dynamics, evolution, genetics, behavior, natural history and life history characteristics of M. cinxia, as well as with species interacting with the butterfly. The parasitoids of M. cinxia have been under long term studies and much has been learned about specific host-parasitoid interactions during the past decade. The research for this Master s thesis was done in the Åland Islands during summer 2010. I conducted a reciprocal transplant style experiment in order to compare the performance of host butterflies (M. cinxia) under attack by different parasitoid wasps (C. melitaearum). I used hosts and parasitoids from five origins around the Baltic Sea: Öland, Uppland, Åland, Saaremaa and Pikku-Tytärsaari. The host-parasitoid relationship was studied in terms of host susceptibility and parasitoid virulence, addressing specifically the possible effects of inbreeding and local adaptation of both parasitoids and their hosts. I compared various factors such as host defence ratio, parasitoid development rate, cocoon production rate etc. I also conducted a small scale C. melitaearum egg development experiment and C. melitaearum external morphology comparison between different parasitoid populations. The results show that host resistance and parasitoid virulence differ between both host and parasitoid populations. For example, Öland hosts were most susceptible to parasitoids and especially vulnerable to Pikku-Tytärsaari wasps. Pikku-Tytärsaari wasps were most successful in terms of parasitoids virulence and efficiency except in Saaremaa hosts, where the wasp did not succeed. Saaremaa hosts were resistant except towards Åland parasitoids. I did not find any simple pattern concerning host resistance and parasitoid virulence between inbred and outbred populations. Also, the effect of local adaptation was not detected, perhaps because metapopulation processes disturb local adaptation of the studied populations. Morphological comparisons showed differences between studied wasp populations and sexual dimorphism was obvious with females being bigger that males. There were also interesting differences among populations in male and female wing shapes. The results raise many further questions. Especially interesting were Pikku-Tytärsaari wasps that did well in terms of efficiency and virulence. Pikku-Tytärsaari is a small, isolated island in the Gulf of Finland and both the host and parasitoids are extremely inbred. For the host and parasitoid to persist in the island, the host has to have some mechanisms to escape the parasitoid. Further research will be done on the subject to discover the mechanisms of Pikku-Tytärsaari host s ability to escape parasitism. Also, genetic analyses will be conducted in the near future to determine the relatedness of used C. melitaearum populations.

Fast- and drift-ice communities in the Bothnian Bay and the impact of UVA radiation on the Baltic Sea ice ecology

The aim of this thesis was to study ecology of Baltic Sea ice from two perspectives. In the first two studies, sea-ice ecology from riverine-influenced fast ice to drift ice in the Bothnian Bay was investigated, whereas the last two studies focus on the sensitivity of sea-ice bacteria and algae to UVA examined in situ. The seasonal sea ice cover is one of the main characteristics of the Baltic Sea, and despite the brackish parental water, the ice structure is similar to polar ice with saline brine inclusions, the sea ice habitat. The decreasing seawater salinity from the northern Baltic Sea to the Bothnian Bay translates to decreasing brine volumes along the gradient, governing the size and community structure of the food webs in ice. However, the drift and fast ice in the Bothnian Bay may differ greatly in this sense, as drift ice may have been formed at more southern locations. Rafting and the formation of snow ice are common processes in the ice field of the Bothnian Bay. As evidenced in this thesis, rafting altered the vertical distribution of organisms and snow-ice formation provided habitable space in the better-illuminated, nitrogen-rich surface layer. The divergence between fast and drift ice became apparent at the more advanced stages, and chlorophyte biomass decreased from fast to drift ice, while the opposite held true for protozoan and metazoan biomass. The brine volumes affected the communities somewhat, and a higher percentage of flagellate species was generally linked to lower brine volumes, whereas chain-forming diatoms were mostly concentrated in layers with larger brine volumes. These results add to knowledge of the ecological significance of the ice cover lasting up to 7 months per year in this area. Sea-ice food webs are generally light-limited, but while increasing light irradiances typically enhance the primary production and further, the secondary production in sea ice, any increase in solar radiation also includes an increase in harmful UVA radiation. The Baltic Sea ice microbial communities were clearly sensitive to UVA and the responses were strongly linked to the earlier light history, as well as to the solar irradiances they were exposed to. The increased biomass of chlorophytes and pennate diatoms, when UVA was excluded, indicates that their normally minor contribution to the biomass in the upper layers of sea ice might be partly dictated by UVA. The effects of UVA on bacterial production in Baltic Sea ice mostly followed the responses in algal growth, but occasionally the exposure to UVA even enhanced the bacterial production. The dominant bacterial class, Flavobacteria, seemed to be UVA-tolerant, whereas all the Alpha-, Beta- and Gammaproteobacteria present in the surface layer showed UVA sensitivity. These results indicate that changes in the light field of ice may alter the community structure and affect the functioning of ice food webs, and are of importance when the effects of thinning of the ice cover are assessed.

Optimal multispecies fishery management in the Baltic Sea: A bioeconomic analysis

The aim of the thesis is to assess the fishery of Baltic cod, herring and sprat by simulation over 50 years time period. We form a bioeconomic multispecies model for the species. We include species interactions into the model because especially cod and sprat stocks have significant effects on each other. We model the development of population dynamics, catches and profits of the fishery with current fishing mortalities, as well as with the optimal profit maximizing fishing mortalities. Thus, we see how the fishery would develop with current mortalities, and how the fishery should be developed in order to yield maximal profits. Especially cod stock has been quite low recently and by optimizing the fishing mortality it could get recovered. In addition, we assess what would happen to the fisheries of the species if more favourable environmental conditions for cod recruitment dominate in the Baltic Sea. The results may yield new information for the fisheries management. According to the results the fishery of Baltic cod, herring and sprat are not at the most profitable level. The fishing mortalities of each species should be lower in order to maximize the profits. By fishing mortality optimizing the net present value would be almost three times higher in the simulation period. The lower fishing mortality of cod would result in a cod stock recovery. If the environmental conditions in the Baltic Sea improved, cod stock would recover even without a decrease in the fishing mortality. Then the increased cod stock would restrict herring and sprat stock remarkably, and harvesting of these species would not be as profitable anymore.

Denitrification in the River Estuaries of the Northern Baltic Sea

Estuaries have been suggested to have an important role in reducing the nitrogen load transported to the sea. We measured denitrification rates in six estuaries of the northern Baltic Sea. Four of them were river mouths in the Bothnian Bay (northern Gulf of Bothnia), and two were estuary bays, one in the Archipelago Sea (southern Gulf of Bothnia) and the other in the Gulf of Finland. Denitrification rates in the four river mouths varied between 330 and 905 mu mol N m(-2) d(-1). The estuary bays at the Archipelago Sea and the Gulf of Bothnia had denitrification rates from 90 mu mol N m(-2) d(-1) to 910 mu mol N m(-2) d(-1) and from 230 mu mol N m(-2) d(-1) to 320 mu mol N m(-2) d(-1), respectively. Denitrification removed 3.6-9.0% of the total nitrogen loading in the river mouths and in the estuary bay in the Gulf of Finland, where the residence times were short. In the estuary bay with a long residence time, in the Archipelago Sea, up to 4.5% of nitrate loading and 19% of nitrogen loading were removed before entering the sea. According to our results, the sediments of the fast-flowing rivers and them estuary areas with short residence times have a limited capacity to reduce the nitrogen load to the Baltic Sea.

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 N-2 production rates, 90-400 mu mol 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 N-2 production. The low percentage did, however, result in a significant error in the total N-2 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.

Aquaculture in the Baltic Sea area – A concept for joint research of Baltic Sea abutters

A concept for joint research on aquaculture in the Baltic Sea area is presented. It consists of three major parts, the promotion of an aquaculture-based-fisheries, the development of low-output land-based aquaculture systems, and the search for sustainably produced substances from aquatic organisms to be used for different processes. They include substitutes for feeding stuffs or products of importance for medical, biotechnological and industrial applications.

The BACOMA cod-end. History and recent developments in BACOMA cod-end regulations and a proposal for abetter specification of the BACOMA cod-end for Baltic Sea cod fishery

To improve the cod stocks in the Baltic Sea, a number of regulations have recently been established by the International Baltic Sea Fisheries Commission (IBSFC) and the European Commission. According to these, fishermen are obliged to use nets with escape windows (BACOMA nets) with a mesh size of the escape window of 120 mm until end of September 2003. These nets however, retain only fish much larger than the legal minimum landing size would al-low. Due to the present stock structure only few of such large fish are however existent. As a consequence fishermen use a legal alternative net. This is a conventional trawl with a cod-end of 130 mm diamond-shaped meshes (IBSFC-rules of 1st April 2002), to be increased to 140 mm on 1st September 2003, according to the mentioned IBSFC-rule. Due legal alterations of the net by the fishermen (e.g. use of extra stiff net material) these nets have acquired extremely low selective properties, i. e. they catch very small fish and produce great amounts of discards. Due to the increase of the minimum landing size from 35 to 38 cm for cod in the Baltic, the amount of discards has even increased since the beginning of 2003. Experiments have now been carried out with the BACOMAnet on German and Swedish commercial and research vessels since arguments were brought forward that the BACOMA net was not yet sufficiently tested on commercial vessels. The results of all experiments conducted so far, are compiled and evaluated here. As a result of the Swedish, Danish and German initiative and research the European Commission reacted upon this in June 2003 and rejected the increase of the diamond-meshed non-BACOMA net from 130 mm to 140mm in September 2003. To protect the cod stocks in the Baltic Sea more effectively the use of traditional diamond meshed cod-ends with-out escape window are prohibited in community waters without derogation, becoming effective 1st of September 2003. To enable more effective and simplified control of the bottom trawl fishery in the Baltic Sea the principle of a ”One-Net-Rule“ is enforced. This is going to be the BACOMA net, with the meshes of the escape window being 110 mm for the time being. The description of the BACOMA net as given in the IBSFC-rules no.10 (revision of the 28th session, Berlin 2002) concentrates on the cod-end and the escape window but only to a less extent on the design and mesh-composition of the remaining parts of the net, such as belly and funnel and many details. Thus, the present description is not complete and leaves, according to fishermen, ample opportunity for manipulation. An initiative has been started in Germany with joint effort from scientists and the fishery to better describe the entire net and to produce a proposal for a more comprehensive description, leaving less space for manipulation. A proposal in this direction is given here and shall be seen as a starting point for a discussion and development towards an internationally uniform net, which is agreed amongst the fishery, scientists and politicians. The Baltic Sea fishery is invited to comment on this proposal, and recommendations for further improvement and specifications are welcomed. Once the design is agreed by the Baltic Fishermen Association, it shall be proposed to the IBSFC and European Commission via the Baltic Fishermen Association.