Conservation Genetics of Exploited Finnish Salmonid Fishes

Genetic diversity is one of the levels of biodiversity that the World Conservation Union (IUCN) has recognized as being important to preserve. This is because genetic diversity is fundamental to the future evolution and to the adaptive flexibility of a species to respond to the inherently dynamic nature of the natural world. Therefore, the key to maintaining biodiversity and healthy ecosystems is to identify, monitor and maintain locally-adapted populations, along with their unique gene pools, upon which future adaptation depends. Thus, conservation genetics deals with the genetic factors that affect extinction risk and the genetic management regimes required to minimize the risk. The conservation of exploited species, such as salmonid fishes, is particularly challenging due to the conflicts between different interest groups. In this thesis, I conduct a series of conservation genetic studies on primarily Finnish populations of two salmonid fish species (European grayling, Thymallus thymallus, and lake-run brown trout, Salmo trutta) which are popular recreational game fishes in Finland. The general aim of these studies was to apply and develop population genetic approaches to assist conservation and sustainable harvest of these populations. The approaches applied included: i) the characterization of population genetic structure at national and local scales; ii) the identification of management units and the prioritization of populations for conservation based on evolutionary forces shaping indigenous gene pools; iii) the detection of population declines and the testing of the assumptions underlying these tests; and iv) the evaluation of the contribution of natural populations to a mixed stock fishery. Based on microsatellite analyses, clear genetic structuring of exploited Finnish grayling and brown trout populations was detected at both national and local scales. Finnish grayling were clustered into three genetically distinct groups, corresponding to northern, Baltic and south-eastern geographic areas of Finland. The genetic differentiation among and within population groups of grayling ranged from moderate to high levels. Such strong genetic structuring combined with low genetic diversity strongly indicates that genetic drift plays a major role in the evolution of grayling populations. Further analyses of European grayling covering the majority of the species’ distribution range indicated a strong global footprint of population decline. Using a coalescent approach the beginning of population reduction was dated back to 1 000-10 000 years ago (ca. 200-2 000 generations). Forward simulations demonstrated that the bottleneck footprints measured using the M ratio can persist within small populations much longer than previously anticipated in the face of low levels of gene flow. In contrast to the M ratio, two alternative methods for genetic bottleneck detection identified recent bottlenecks in six grayling populations that warrant future monitoring. Consistent with the predominant role of random genetic drift, the effective population size (N_e) estimates of all grayling populations were very low with the majority of N_e estimates below 50. Taken together, highly structured local populations, limited gene flow and the small N_e of grayling populations indicates that grayling populations are vulnerable to overexploitation and, hence, monitoring and careful management using the precautionary principles is required not only in Finland but throughout Europe. Population genetic analyses of lake-run brown trout populations in the Inari basin (northernmost Finland) revealed hierarchical population structure where individual populations were clustered into three population groups largely corresponding to different geographic regions of the basin. Similar to my earlier work with European grayling, the genetic differentiation among and within population groups of lake-run brown trout was relatively high. Such strong differentiation indicated that the power to determine the relative contribution of populations in mixed fisheries should be relatively high. Consistent with these expectations, high accuracy and precision in mixed stock analysis (MSA) simulations were observed. Application of MSA to indigenous fish caught in the Inari basin identified altogether twelve populations that contributed significantly to mixed stock fisheries with the Ivalojoki river system being the major contributor (70%) to the total catch. When the contribution of wild trout populations to the fisheries was evaluated regionally, geographically nearby populations were the main contributors to the local catches. MSA also revealed a clear separation between the lower and upper reaches of Ivalojoki river system – in contrast to lower reaches of the Ivalojoki river that contributed considerably to the catch, populations from the upper reaches of the Ivalojoki river system (>140 km from the river mouth) did not contribute significantly to the fishery. This could be related to the available habitat size but also associated with a resident type life history and increased cost of migration. The studies in my thesis highlight the importance of dense sampling and wide population coverage at the scale being studied and also demonstrate the importance of critical evaluation of the underlying assumptions of the population genetic models and methods used. These results have important implications for conservation and sustainable fisheries management of Finnish populations of European grayling and brown trout in the Inari basin.

Swan Lake ballet

Äänitetty: 14.-15.12.1954, University of Minnesota, Northrop Memorial Auditorium.

Holocene Varved Lake Sediments as Records of Palaeoenvironmental Change and Geomagnetic Palaeosecular Variation in Eastern Finland

This dissertation discusses Holocene palaeoenvironmental and palaeomagnetic secular variation (PSV) records reconstructed from sediments preserved in Lake Lehmilampi (63º37´N, 29º06´E) and Lake Kortejärvi (63º37´N, 28º56´E) in eastern Finland. Several piston and freeze cores were obtained from both lakes for varve and magnetic analyses. Sediment samples were impregnated in low-viscosity epoxy and physical parameters of varves, including varve thickness and relative grey-scale values, were recorded using x-ray densitometry combined with semiautomatic digital image analysis. On average, varve records of Lehmilampi and Kortejärvi cover 5122 and 3902 years, respectively. Past solar activity, as estimated by residual ₁₄C data, compares favourably with varve thicknesses from Lehmilampi during the last 2000 years. This indicates the potential of clastic-organic varves to record sensitively climatic variations. Bulk magnetic parameters, including magnetic susceptibility together with natural, anhysteretic and isothermal remanent magnetizations, were measured to describe mineral magnetic properties and geomagnetic palaeosecular variation recorded in the sediments. Main stages in the development of the investigated lakes are reflected in the variations in the mineral magnetic records, sediment lithology and composition. Similar variations in magnetic parameters and sediment organic matter suggest contribution of bacterial magnetite in the magnetic assemblages of Lehmilampi. Inclination and relative declination records yielded largely consistent results, attesting to the great potential of these sediments to preserve directional palaeosecular variation in high resolution. The PSV data from Lehmilampi and Kortejärvi were stacked into North Karelian PSV stack, which may be used for dating homogenous lake sediments in the same regional context. Reconstructed millennial variations in relative palaeointensity results are approximately in agreement with those seen in the absolute palaeointensity data from Europe. Centennial variations in the relative palaeointensity, however, are influenced by environmental changes. Caution is recommended when using varved lake sediments in reconstructing relative palaeointensity.

Identification and application of bile metabolites to assess the exposure of fish to pharmaceuticals in the environment

Thousands of tons of pharmaceuticals are consumed yearly worldwide. Due to the continuous and increasing consumption and their incomplete elimination in wastewater treatment plants (WWTP), pharmaceuticals and their metabolites can be detected in receiving waters, although at low concentrations (ng to low μg/L). As bioactive molecules the presence of pharmaceuticals in the aquatic environment must be considered potentially hazardous for the aquatic organisms. In this thesis, the biotransformation and excretion of pharmaceuticals in fish was studied. The main biotransformation pathways of three anti‐inflammatory drugs, diclofenac, naproxen and ibuprofen, in rainbow trout were glucuronidation and taurine conjugation of the parent compounds and their phase I metabolites. The same metabolites were present in fish bile in aquatic exposures as in fish dosed with intraperitoneal injection. Higher bioconcentration factor in bile (BCFbile) was found for ibuprofen when compared to diclofenac and naproxen. Laboratory exposure studies were followed by a study of uptake of pharmaceuticals in a wild fish population living in lake contaminated with WWTP effluents. Of the analyzed 17 pharmaceuticals and six phase I metabolites, only diclofenac, naproxen and ibuprofen was present in bream and roach bile. It was shown, that diclofenac, naproxen and ibuprofen excreted by the liver can be found in rainbow trout and in two native fish species living in the receiving waters. In the bream and roach bile, the concentrations of diclofenac, naproxen and ibuprofen were roughly 1000 times higher than those found in the lake water, while in the laboratory exposures, the bioconcentration of the compounds and their metabolites in rainbow trout bile were at the same level as in wild fish or an order of magnitude higher. Thus, the parent compounds and their metabolites in fish bile can be used as a reliable biomarker to monitor the exposure of fish to environmental pharmaceuticals present in water receiving discharges from WWTPs.

Effects of anthropogenic and natural hydrological changes on the behavior of the acidic metal discharge from acid sulfate soils in a river- and lake system in western Finland

Larsmo-Öjasjön i Österbotten skapades genom invallningar på 1960-talet pga. industrins behov av sötvatten. Sedan dess har vattenområdet drabbats av återkommande försurning och fiskdöd, och invallningen har ofta beskyllts för problemen. Avhandlingen undersöker syrabelastningen i området; bl.a. hur markanvändning, hydrologi och klimatförändringen påverkar belastningen. Konsekvenserna undersöks med fiskyngel som bioindikator, och olika miljömetoder testas och diskuteras. Ökad kunskap om försurningen hjälper oss att tillämpa effektiva miljömetoder och få förbättrad vattenkvalitet i framtiden. Den primära orsaken till den försämrade vattenkvaliteten under de senaste 40 åren är intensiv dikning av svavelrika sediment. Detta leder till oxidering av svavlet till svavelsyra och uppkomst av sura sulfatjordar. Syran löser upp mängder med toxiska metaller som spolas ut i vattendragen. Undersökningen visar att tiotusentals ton svavelsyra tillsammans med stora mängder metaller rinner till Larsmo-Öjasjön per år från sura sulfatjordar. Åarna bidrar med mest belastning, men den sammanlagda belastningen från de otaliga dikena och bäckarna är oväntat stor. Andra potentiella källor till försurningen, t.ex. muddringar och humussyror, beräknas vara obetydliga. Syra- och metallbelastningen varierar kraftigt med hydrologin, dvs. störst belastning sker under vår- och höstflöden. En eventuell klimatförändring kan ändra på avrinningsmönstret och orsaka mera belastning vintertid. Den årligt återkommande syra- och metallbelastningen kan ofta hindra lakens förökning, vilket kan ha större långtgående konsekvenser för fiskpopulationerna än de relativt sällsynta stora surchockerna med synlig fiskdöd. För att förebygga skador på vattendragen bör man undvika att dränera svavelrika sedimenten. På redan existerande sura sulfatjordar visade sig kontroll av grundvattennivån kunna möjliggöra en effektiverad markanvändning utan märkbart ökade miljökonsekvenser.

Long-term nutrient load management and lake restoration: Case of Säkylän Pyhäjärvi (SW Finland)

Eutrophication caused by anthropogenic nutrient pollution has become one of the most severe threats to water bodies. Nutrients enter water bodies from atmospheric precipitation, industrial and domestic wastewaters and surface runoff from agricultural and forest areas. As point pollution has been significantly reduced in developed countries in recent decades, agricultural non-point sources have been increasingly identified as the largest source of nutrient loading in water bodies. In this study, Lake Säkylän Pyhäjärvi and its catchment are studied as an example of a long-term, voluntary-based, co-operative model of lake and catchment management. Lake Pyhäjärvi is located in the centre of an intensive agricultural area in southwestern Finland. More than 20 professional fishermen operate in the lake area, and the lake is used as a drinking water source and for various recreational activities. Lake Pyhäjärvi is a good example of a large and shallow lake that suffers from eutrophication and is subject to measures to improve this undesired state under changing conditions. Climate change is one of the most important challenges faced by Lake Pyhäjärvi and other water bodies. The results show that climatic variation affects the amounts of runoff and nutrient loading and their timing during the year. The findings from the study area concerning warm winters and their influences on nutrient loading are in accordance with the IPCC scenarios of future climate change. In addition to nutrient reduction measures, the restoration of food chains (biomanipulation) is a key method in water quality management. The food-web structure in Lake Pyhäjärvi has, however, become disturbed due to mild winters, short ice cover and low fish catch. Ice cover that enables winter seining is extremely important to the water quality and ecosystem of Lake Pyhäjärvi, as the vendace stock is one of the key factors affecting the food web and the state of the lake. New methods for the reduction of nutrient loading and the treatment of runoff waters from agriculture, such as sand filters, were tested in field conditions. The results confirm that the filter technique is an applicable method for nutrient reduction, but further development is needed. The ability of sand filters to absorb nutrients can be improved with nutrient binding compounds, such as lime. Long-term hydrological, chemical and biological research and monitoring data on Lake Pyhäjärvi and its catchment provide a basis for water protection measures and improve our understanding of the complicated physical, chemical and biological interactions between the terrestrial and aquatic realms. In addition to measurements carried out in field conditions, Lake Pyhäjärvi and its catchment were studied using various modelling methods. In the calibration and validation of models, long-term and wide-ranging time series data proved to be valuable. Collaboration between researchers, modellers and local water managers further improves the reliability and usefulness of models. Lake Pyhäjärvi and its catchment can also be regarded as a good research laboratory from the point of view of the Baltic Sea. The main problem in both of them is eutrophication caused by excess nutrients, and nutrient loading has to be reduced – especially from agriculture. Mitigation measures are also similar in both cases.

Design, implementation, and evaluation of an online water quality monitoring system in Lake Saimaa, Finland

Environmental threats are growing nowadays, they became global issues. People around the world try to face these issues by two means: solving the current affected environs and preventing non-affected environs. This thesis describes the design, implementation, and evaluation of online water quality monitoring system in Lake Saimaa, Finland. The water quality in Lake Saimaa needs to be monitored in order to provide responsible bodies with valuable information which allows them to act fast in order to prevent any negative impact on the lake's environment. The objectives were to design a suitable system, implement the system in Lake Saimaa, and then to evaluate the applicability and reliability of such systems for this environment. The needs for the system were first isolated, and then the design, needed modifications, and the construction of the system took place. After that was the testing of the system in Lake Saimaa in two locations nearby Mikkeli city. The last step was to evaluate the whole system. The main results were that the application of online water quality monitoring systems in Lake Saimaa can benefit of many advantages such as reducing the required manpower, time and running costs. However, the point of unreliability of the exact measured values of some parameters is still the drawback of such systems which can be developed by using more advanced equipments with more sophisticated features specifically for the purpose of monitoring in the predefined location.

Persistent organic pollutants in sediments and fish from Lake Victoria, Uganda

This thesis describes the occurrence and sources of selected persistent organic pollutants (POPs) such as polychlorinated dibenzo-p-dioxins (PCDDs), polychlorinated dibenzofurans (PCDFs), polychlorinated biphenyls (PCBs), polybrominated diphenyl ethers (PBDEs) and hexachlorocyclohexanes (HCHs) in the northern watershed of Lake Victoria. Sediments and fish were collected from three highly polluted embayments (i.e. Murchison Bay, Napoleon Gulf and Thurston Bay) of the lake. The analysis for PCDD/Fs, PCBs and PBDEs was done using a high resolution mass spectrometer coupled to a gas chromatograph (GC), and a GC equipped with an electron capture detector was used for HCHs. Total (Σ) PCDD/Fs, PCBs and PBDEs in sediments ranged from 3.19 to 478, 313 to 4325 and 60.8 to 179 pg g-1 dry weight (dw), respectively. The highest concentrations of pollutants were found at sites close to industrial areas and wastewater discharge points. The maximum concentrations of PCDD/Fs, PCBs, PBDEs and HCHs in fish muscle homogenates were 49, 779, 495 and 45,900 pg g-1 wet weight (ww), respectively. The concentrations of the pollutants in Nile perch (Lates niloticus) were significantly greater than those in Nile tilapia (Oreochromis niloticus), possibly due to differences in trophic level and dietary feeding habits among fish species. World Health Organization-toxic equivalency quotient (WHO2005-TEQ) values in the sediments were up to 4.24 pg g-1 dw for PCDD/Fs and 0.55 pg TEQ g-1 dw for the 12 dioxin-like PCBs (dl-PCBs). 23.1% of the samples from the Napoleon Gulf were above the interim sediment quality guideline value of 0.85 pg WHO-TEQ g-1 dw set by the Canadian Council for Ministers of the Environment. The WHO2005-TEQs in fish were 0.001-0.16 pg g-1 for PCDD/Fs and 0.001-0.31 pg g-1 ww for dl- PCBs. The TEQ values were within a permissible level of 3.5 pg g−1 ww recommended by the European Commission. Based on the Commission set TEQs and minimum risk level criteria formulated by the Agency for Toxic Substances and Disease Registry, the consumption of fish from Lake Victoria gives no indication of health risks associated to PCDD/Fs and PCBs. Principal component analysis (PCA) indicated that anthropogenic activities such as agricultural straw open burning, medical waste incinerators and municipal solid waste combustors were the major sources of PCDD/Fs in the watershed of Lake Victoria. The ratios of α-/γ-HCH varied from 0.89 to 1.68 suggesting that the highest HCH residues mainly came from earlier usage and fresh γ-HCH (lindane). In the present study, the concentration of POPs in fish were not significantly related to those in sediments, and the biota sediment accumulation factor (BSAF) concept was found to be a poor predictor of the bioavailability and bioaccumulation of environmental pollutants.

Geochemistry and water quality of Lake Qarun, Egypt

Water geochemistry is a very important tool for studying the water quality in a given area. Geology and climate are the major natural factors controlling the chemistry of most natural waters. Anthropogenic impacts are the secondary sources of contamination in natural waters. This study presents the first integrative approach to the geochemistry and water quality of surface waters and Lake Qarun in the Fayoum catchment, Egypt. Moreover, geochemical modeling of Lake Qarun was firstly presented. The Nile River is the main source of water to the Fayoum watershed. To investigate the quality and geochemistry of this water, water samples from irrigation canals, drains and Lake Qarun were collected during the period 2010‒2013 from the whole Fayoum drainage basin to address the major processes and factors governing the evolution of water chemistry in the investigation area. About 34 physicochemical quality parameters, including major ions, oxygen isotopes, trace elements, nutrients and microbiological parameters were investigated in the water samples. Multivariable statistical analysis was used to interpret the interrelationship between the different studied parameters. Geochemical modeling of Lake Qarun was carried out using Hardie and Eugster’s evolutionary model and a model simulated by PHREEQC software. The crystallization sequence during evaporation of Lake Qarun brine was also studied using a Jänecke phase diagram involving the system Na‒K‒Mg‒ Cl‒SO4‒H2O. The results show that the chemistry of surface water in the Fayoum catchment evolves from Ca- Mg-HCO3 at the head waters to Ca‒Mg‒Cl‒SO4 and eventually to Na‒Cl downstream and at Lake Qarun. The main processes behind the high levels of Na, SO4 and Cl in downstream waters and in Lake Qarun are dissolution of evaporites from Fayoum soils followed by evapoconcentration. This was confirmed by binary plots between the different ions, Piper plot, Gibb’s plot and δ18O results. The modeled data proved that Lake Qarun brine evolves from drainage waters via an evaporation‒crystallization process. Through the precipitation of calcite and gypsum, the solution should reach the final composition "Na–Mg–SO4–Cl". As simulated by PHREEQC, further evaporation of lake brine can drive halite to precipitate in the final stages of evaporation. Significantly, the crystallization sequence during evaporation of the lake brine at the concentration ponds of the Egyptian Salts and Minerals Company (EMISAL) reflected the findings from both Hardie and Eugster’s evolutionary model and the PHREEQC simulated model. After crystallization of halite at the EMISAL ponds, the crystallization sequence during evaporation of the residual brine (bittern) was investigated using a Jänecke phase diagram at 35 °C. This diagram was more useful than PHREEQC for predicting the evaporation path especially in the case of this highly concentrated brine (bittern). The predicted crystallization path using a Jänecke phase diagram at 35 °C showed that halite, hexahydrite, kainite and kieserite should appear during bittern evaporation. Yet the actual crystallized mineral salts were only halite and hexahydrite. The absence of kainite was due to its metastability while the absence of kieserite was due to opposed relative humidity. The presence of a specific MgSO4.nH2O phase in ancient evaporite deposits can be used as a paleoclimatic indicator. Evaluation of surface water quality for agricultural purposes shows that some irrigation waters and all drainage waters have high salinities and therefore cannot be used for irrigation. Waters from irrigation canals used as a drinking water supply show higher concentrations of Al and suffer from high levels of total coliform (TC), fecal coliform (FC) and fecal streptococcus (FS). These waters cannot be used for drinking or agricultural purposes without treatment, because of their high health risk. Therefore it is crucial that environmental protection agencies and the media increase public awareness of this issue, especially in rural areas.

“Good night, dear Lake of Shining Waters”. Luonto L. M. Montgomeryn Anne of Green Gables –teoksessa

Kanadalaisen kirjailijan L. M. Montgomeryn tunnetun ja rakastetun Anna-sarjan ensimmäinen osa Anne of Green Gables -teos (1908) kertoo orvosta tytöstä, joka pääsee Prinssi Edwardin saaren Avonlean kylään asumaan naimattoman sisarusparin Marilla ja Matthew`n kasvattilapseksi. Luonto on keskeisellä sijalla teoksessa: Anna on luontoa rakastava ja luonnon keskellä viihtyvä tyttö ja myös teoksen kertoja kuvaa paljon luontoa. Tutkimukseni käsittelee luontoa Anne of Green Gables -teoksessa. Tutkin luontoteemaa teoksessa seuraavista näkökulmista: Analysoin, kuinka luonto kuvataan teoksessa ja millainen on teoksen luontokuva. Luontokuvaa käsittelen pastoraalin käsitteen kautta, ja kysyn teokselta, voidaanko se tulkita pastoraaliksi ja kuinka teos asettuu pastoraalin traditioon. Lisäksi erittelen, millainen on päähenkilön Annan luontosuhde. Millaisen arvon hän antaa luonnolle ja mitä merkityksiä hän liittää luontoon? Tutkimus asettuu ekokriittiseen viitekehykseen, ja pohdin myös sitä, minkälainen on teoksen representaatio luonnosta ekokriittisesti tarkastellen. Amerikkalaisen transsendentalistin Ralph Waldo Emersonin luontofilosofia kulkee myös taustajuonteena tutkielmassa ja erittelen, mitä yhtymäkohtia hänen luontofilosofiassaan on Anne of Green Gables -teokseen. Anne of Green Gables -teoksen luontokuvaa voidaan luonnehtia romantisoiduksi ja idealisoivaksi. Luonto kuvataan kauniiksi, ja sen kauneutta ylistää niin Anna kuin teoksen kaikkitietävä kertojakin. Teoksen luontokuva on pastoraalinen, sillä siinä esiintyy vahvasti pastoraaliin liittyvä diskurssi maaseudun ja kaupungin vastakkainasettelusta siten, että maaseutu assosioituu positiivisesti, kun taas kaupunki nähdään negatiivisessa valossa. Luonnonläheinen Avonlean kylä näyttäytyy teoksessa niin idealisoidussa valossa, että se muistuttaa jo jonkinlaista myyttistä paratiisia. Näin ollen teoksen pastoraalisuus saa pejoratiivisiakin piirteitä. Annan luontosuhdetta värittää voimakas kunnioitus ja arvostus luontoa kohtaan. Hänen luontoon liittämistä arvoista keskeiselle sijalle nousee luonnon kauneus. Luonnon kauneuden arvostaminen toimii kuitenkin porttina luonnon toiseuttamiseen, mistä syntyy pieni arvoristiriita. Toisaalta kuitenkin Anna samaistuu luontoon tavalla, joka hämärtää perinteistä subjekti–objekti-dikotomiaa. Anna näyttäytyy vihreän maailman arkkityyppinä, lapsena, joka sulautuu luontoon ja kokee sen kodikseen. Luonto edustaa Annalle vapautta ja herättää myös hänen mielikuvituksena ja taiteellisen inspiraation. Lisäksi luonto on Annalle paikka, jossa voi saavuttaa autenttisen yhteyden Jumalaan.