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Species invasion and range expansion are currently under scrutiny due to increasing anthropogenic impact on the natural environment. This is also true for harmful algal blooms, which have been reported to have increased in frequency. However, this research is challenging due to the ephemeral nature, small size and mostly low concentrations of microalgae in the environment. One such species is the nuisance microalga Gonyostomum semen (Raphidophyceae), which has increased in occurrence in northern Europe in recent decades. The question of whether the species has expanded its habitat range or if it was already present in the lakes but was too rare to be detected remains unanswered. The aim of the present study was to determine the genetic structure and dispersal pathways of G. semen using RAD (restriction-site-associated DNA) tag sequencing. For G. semen, which has a huge genome (32 Gbp), we faced particular challenges, but were nevertheless able to recover over 1000 single nucleotide polymorphisms at high coverage. Our data revealed a distinct population genetic structure, demonstrating a divide of western and eastern populations that probably represent different lineages. Despite significant genetic differentiation among lakes, we found only limited isolation-by-distance. While we had expected a pattern of recent expansion northwards, the data demonstrated gene flow from the northeast/east towards the southwest/west. This genetic signature suggests that the observed gene flow may be due to dispersal by autumn migratory birds, which act as dispersal vectors of resistant resting propagules that form at the end of the G. semen blooms.

Increases in water colour (brownification) have been observed in aquatic systems in the Northern Hemisphere, partly caused by increased loading of organic carbon from terrestrial origins. We investigated the effect of increase in water colour on the composition, structure and function of lake plankton communities (bacteria, phytoplankton and zooplankton) conducting a mesocosm experiment in 3 medium-coloured lakes (average absorbance at 420 nm: 0.034 cm(-1)), with different nutrient concentrations and phytoplankton community composition. To simulate an increase in water colour, we added humic substances (HuminFeed) at 3 different concentrations. The additions significantly affected the water colour of the mesocosms, but had no measurable effect on total organic carbon concentration, thus change in light conditions was the main effect of our treatment on the plankton communities. The increase in water colour did not significantly affect the measured functions (productivity, respiration) and biomass of the plankton communities (bacteria, phytoplankton and zooplankton), but led to changes in the relative abundance of some phytoplankton taxa and, to a lesser extent, the bacterial community (differences in relative abundance). The treatments had no significant effect on zooplankton biomass or composition. Our study suggests that increases in water colour favour low-light-adapted phytoplankton species, which in turn also can affect bacterial composition, whereas the change in light climate had no clear impact on the functioning of plankton communities in weakly humic lakes.

Ecological stability is the central framework to understand an ecosystem’s ability to absorb or recover from environmental change. Recent modelling and conceptual work suggests that stability is a multidimensional construct comprising different response aspects. Using two freshwater mesocosm experiments as case studies, we show how the response to single perturbations can be decomposed in different stability aspects (resistance, resilience, recovery, temporal stability) for both ecosystem functions and community composition. We find that extended community recovery is tightly connected to a nearly complete recovery of the function (biomass production), whereas systems with incomplete recovery of the species composition ranged widely in their biomass compared to controls. Moreover, recovery was most complete when either resistance or resilience was high, the latter associated with low temporal stability around the recovery trend. In summary, no single aspect of stability was sufficient to reflect the overall stability of the system.

The nuisance alga Gonyostomum semen (Raphidophyceae) has expanded in the Nordic countries during the last decades and can dominate lake phytoplankton communities almost completely. A possible explanation to its dominance could be limited grazing by zooplankton. We investigated the potential grazing pressure on G. semen using an experimental approach supported by field data. We determined the grazing rate by cladocerans, calanoid copepods, and Chaoborus larvae to determine which were able to feed on G. semen. Only the large cladoceran Daphnia magna was able to feed successfully on G. semen. The large cell size of G. semen was likely a limiting factor for the filtering apparatus of smaller cladocerans. The copepod Eudiaptomus gracilis did not graze on G. semen, although the mechanism behind this selective feeding is still unknown. In addition to the experimental study, we quantified the zooplankton and phytoplankton communities in 40 lakes to determine the composition and abundance of the zooplankton communities co-occurring with G. semen, suggesting that large cladoceran species were not present in lakes where G. semen occurred. Hence, the growth of G. semen is not significantly controlled by grazing in natural systems, which likely facilitates bloom formation and invasion success of G. semen.

Receding glaciers are among the most obvious changes caused by global warming, and glacial meltwater entering lakes generally forms plumes of particles. By taking vertical samples along a horizontal gradient from such a particle source, we found that photosynthetically active radiation (PAR) and ultraviolet radiation (UVR) attenuated 20-25% faster close to the inflow of suspended particles compared with the more transparent part of the gradient. All sampled stations had a deep chlorophyll a (Chl a) maximum at 15-20 m which was more distinct in the transparent part of the horizontal gradient. Picocyanobacteria increased in abundance in more transparent water and their numbers were tightly correlated with the intensity of the deep Chl a maxima. Motile species of phytoplankton had a deeper depth distribution in transparent versus less transparent water. Yet other species, like Chrysochromulina parva, that can withstand high PAR intensities and low nutrient concentrations, increased in abundance as the water became more transparent. Also copepods increased in abundance, indicating that they are more successful in transparent water. We conclude that sediment input into lakes creates horizontal gradients in PAR and UVR attenuation which strongly affect both distribution and behavior of phyto-and zooplankton. The input of glacial flour creates a sub-habitat that can function as a refuge for species that are sensitive to high PAR and UVR exposure. When the glacier has vanished, this habitat may disappear. During the melting period, with heavy sediment input, we predict that competitive species in transparent waters, like Chrysocromulina, picocyanobacteria and copepods, will become less common. The deep Chl a maxima is also likely to become less developed. Hence, glacier melting will probably have profound effects on both species composition and behavior of several planktonic taxa with potential effects on the food web.

Climate change is altering temperatures and precipitation patterns all over the world. In Patagonia, Argentina, predicted increase in precipitation together with rapidly melting glaciers increase the surface runoff, and thereby the transport of suspended solids to recipient lakes. Suspended solids affect the visual conditions in the water which in turn restricts visual foraging. The native fish Aplochiton zebra Jenyns, and its filter-feeding cladoceran prey, Daphnia commutata Ekman, were subjected to foraging experiments at three turbidity levels. A. zebra foraging rate was substantially reduced at naturally occurring turbidity levels and the filtering rate of D. commutata was reduced at the highest turbidity level. This indicates that Daphnia may be partly released from predation from A. zebra at the same time as it can maintain relatively high feeding rates as turbidity increases. Lower foraging rates at the same time as the metabolic demand increases, through increased temperatures, may result in larger effects on A. zebra than could be expected from increases in turbidity or temperature alone. Turbidity may, as an indirect effect of climate change, decrease planktivore foraging rates and thereby alter the interaction strength between trophic levels.

Gonyostmum semen is a freshwater raphidophyte that has increased in occurrence and abundance in several countries in northern Europe since the 1980s. More recently, the species has expanded rapidly also in north-eastern Europe, and it is frequently referred to as invasive. To better understand the species history, we have explored the phylogeography of G. semen using strains from northern Europe, United States, and Japan. Three regions of the ribosomal RNA gene (small subunit [SSU], internal transcribed spacer [ITS] and large subunit [LSU]) and one mitochondrial DNA marker (cox1) were analyzed. The SSU and partial LSU sequences were identical in all strains, confirming that they belong to the same species. The ITS region differentiated the American from the other strains, but showed high intra-strain variability. In contrast, the mitochondrial marker cox1 showed distinct differences between the European, American, and Japanese strains. Interestingly, only one cox1 haplotype was detected in European strains. The overall low diversity and weak geographic structure within northern European strains supported the hypothesis of a recent invasion of new lakes by G. semen. Our data also show that the invasive northern European lineage is genetically distinct from the lineages from the other continents. Finally, we concluded that the mitochondrial cox1 was the most useful marker in determining large-scale biogeographic patterns in this species.

Biological invasions often cause major perturbations in the environment and are well studied among macroorganisms. Less is known about invasion by free-living microbes. Gonyostomum semen (Raphidophyceae) is a freshwater phytoplankton species that has increased in abundance in Northern Europe since the 1980's and has expanded its habitat range. In this study, we aimed to determine the genetic population structure of G. semen in Northern Europe and to what extent it reflects the species' recent expansion. We sampled lakes from 12 locations (11 lakes) in Norway, Sweden and Finland. Multiple strains from each location were genotyped using Amplified Fragment Length Polymorphism (AFLP). We found low differentiation between locations, and low gene diversity within each location. Moreover, there was an absence of genetic isolation with distance (Mantel test, p = 0.50). According to a Bayesian clustering method all the isolates belonged to the same genetic population. Together our data suggest the presence of one metapopulation and an overall low diversity, which is coherent with a recent expansion of G. semen.

This study evaluates the treatment efficiency of a chemical-free water treatment for treating the secondary effluent of a municipal wastewater treatment plant with the aim of reusing the water for agriculture. Urban wastewater was treated by three units run in series: a full-scale horizontal sub-surface flow constructed wetland, a small pond with an ultrasound (US) system and a UV device. The treatment efficiency was evaluated in terms of the Italian wastewater limits for irrigation reuse, water quality improvement (removal percentage) and algae bloom control. The tolerable infection risk, associated with the use of wastewaters for irrigating crops, was also assessed by applying the microbial risk analyses proposed in the WHO guidelines for wastewater reuse. The constructed wetland was efficient in reducing physical-chemical and microbiological concentrations, and its efficiency was very steady over the investigation period. The UV system significantly improved water quality (p

Sterols are a class of membrane-reinforcing, ringed lipids which have a long history of examination in algae as a means of deriving chemotaxonomic relationships and as potential lipidic biomarkers. The Raphidophyceae represent a class of harmful, bloom-forming, marine and freshwater algae. To date, there have been four published examinations of their sterol composition, focusing primarily on brown-pigmented, marine species within the genera, Chattonella, Fibrocapsa, and Heterosigma. Lacking in these examinations has been the species Gonyostomum semen Ehrenb., which is a green-pigmented, freshwater raphidophyte with a worldwide distribution. The goal of this study was to examine the sterol composition of this nuisance alga, determine the potential of using its sterol profile as a biomarker, and finally to determine if there is any intraspecific variability between isolates. We have examined 21 isolates of G. semen from a number of Scandinavian lakes, and all were found to produce two major sterols, 24-ethylcholesta-5,22E-dien-3β-ol and 24-ethylcholest-5-en-3β-ol, and 24-methylcholest-5-en-3β-ol as a minor sterol; the presence of 24-ethylcholesta-5,22E-dien-3β-ol differentiates G. semen from brown-pigmented, marine raphidophytes which generally lack it. The results of this study indicate that isolates of G. semen from geographically separate lakes across Finland and Scandinavia have the same sterol biosynthetic pathway, and that there is no evolutionary divergence between the isolates with regard to sterol composition. The sterols of G. semen are not considered to be useful biomarkers for this particular organism because they are commonly found in other algae and plants.

The field of genetic diversity in protists, particularly phytoplankton, is under expansion. However, little is known regarding variation in genetic diversity within populations over time. The aim of our study was to investigate intrapopulation genetic diversity and genetic differentiation in the freshwater bloom-forming microalga Gonyostomum semen (Raphidophyceae). The study covered a 2-year period including all phases of the bloom. Amplified fragment length polymorphism (AFLP) was used to determine the genetic structure and diversity of the population. Our results showed a significant differentiation between samples collected during the two blooms from consecutive years. Also, an increase of gene diversity and a loss of differentiation among sampling dates were observed over time within a single bloom. The latter observations may reflect the continuous germination of cysts from the sediment. The life cycle characteristics of G. semen, particularly reproduction and recruitment, most likely explain a high proportion of the observed variation. This study highlights the importance of the life cycle for the intraspecific genetic diversity of microbial species, which alternates between sexual and asexual reproduction., SI

Human impacts on coastal waters, through eutrophication and overfishing, associated with the global change phenomenon are leading to a proliferation of both micro- and macroalgae. Algae can settle and develop on a wide range of surfaces including both natural and man-made structures. They are fast colonisers and can out-compete numerous species. The fixation of marine organisms on immersed manmade structures is responsible for major economic costs and the recent recrudescence of algae in the environment might increase the fouling phenomenon observed: the resistance penalty for ships is known to be increased by 11% in presence of a light microalgal slime to 34% in case of colonisation by macroalgae (Schultz, 2007). The paints used nowadays often contain biocides, are non selective against target organisms and are not environmentally friendly. The development of new antifouling paints requires a better understanding of both the mechanisms of fixation and colonisation of the fouler organisms. This chapter focuses on the fixation of algae (both micro and macro), the damages they cause and the method of prevention currently used.