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The physiological performance of organisms depends on their environmental context, resulting in performance-response curves along environmental gradients. Parasite performance-response curves are generally expected to be broader than those of their hosts due to shorter generation times and hence faster adaptation. However, certain environmental conditions may limit parasite performance more than that of the host, thereby providing an environmental refuge from disease. Thermal disease refuges have been extensively studied in response to climate warming, but other environmental factors may also provide environmental disease refuges which, in turn, respond to global change. Here, we 1) showcase laboratory and natural examples of refuges from parasites along various environmental gradients, and 2) provide hypotheses on how global environmental change may affect these refuges. We strive to synthesise knowledge on potential environmental disease refuges along different environmental gradients including salinity and nutrients, in both natural and food-production systems. Although scaling up from single host-parasite relationships along one environmental gradient to their interaction outcome in the full complexity of natural environments remains difficult, integrating host and parasite performance-response can serve to formulate testable hypotheses about the variability in parasitism outcomes and the occurrence of environmental disease refuges under current and future environmental conditions. This article is protected by copyright. All rights reserved.

Zoosporic parasites (i.e. fungi and fungi-like aquatic microorganisms) constitute important drivers of natural populations, causing severe host mortality. Economic impacts of parasitic diseases are notable in the microalgae biotech industry, affecting production of food ingredients, biofuels, pharma- and nutraceuticals. While scientific research on this topic is gaining traction by increasing studies elucidating the functional role of zoosporic parasites in natural ecosystems, we are currently lacking integrated and interdisciplinary efforts for effectively detecting and controlling parasites in the microalgae industry. To fill this gap we propose to establish an innovative, dynamic European network connecting scientists, industries and stakeholders to optimize information exchange, equalize access to resources and to develop a joint research agenda. ParAqua aims at compiling and making available all information on the occurrence of zoosporic parasites and their relationship with hosts, elucidate drivers and evaluate impacts of parasitism in natural and man-made aquatic environments. We aim to implement new tools for monitoring and prevention of infections, and to create protocols and a Decision Support Tool for detecting and controlling parasites in the microalgae biotech production. Applied knowledge on zoosporic parasites can feed back from industry to ecology, and we therefore will explore whether the developed tools can be applied for monitoring lakes and reservoirs. Short-Term Scientific Missions and Training Schools will be organised specifically for early stage scientists and managers – with a specific focus on ITC – with the aim to share and integrate both scientific and applied expertise and increase exchange between basic and applied researchers and stakeholders.

Datasets and R scripts for the COVID anthropause case study sites for the publication entitled "Lessons learned from the COVID-19 anthropause in the Netherlands on urban aquatic ecosystem services provisioning and management" When using the .csv files for Figure 2B (bathing waters, file: Zwemmen buiten) and Figure 2C (national parks; files: De Alde Feanen, De Biesbosch, De Groote Peel, Dwingelderveld, Lauwersmeer and Weerribben-Wieden), make sure the dates are in the following format: dd-mm-yyyy When running the Figure 2C codes (in R script "COVID_GoogleTrends_Script"), set the working directory to a folder containing only the national parks .csv files.

1. Parasites are generally considered the most commonly occurring type of consumers, yet their biomass and population dynamics are rarely quantified at community level. 2. Here, we used 12 years of weekly or fortnightly monitoring data (518 time points) to determine the occurrence of chytrids, fungal parasites of phytoplankton, to assess their seasonality and long-term (seasonally-detrended) dynamics in the pelagic plankton community of a temperate, eutrophic, and polymictic lake. 3. Chytrid infections were observed in c. 75% of all samples with recurrent infections in multiple host taxa. Infection prevalence was highest in spring, but infections occurred throughout the entire year with an average of 2.3 host taxa infected per time point (ranging from 0 to 10 host taxa) and an average infection prevalence of 2.78% (ranging from 0% to 47.35%). Infected host biomass equalled that of the carnivorous zooplankton and decreased over time, while infection prevalence remained unchanged. Seasonal infection prevalence increased with phytoplankton biomass, but decreased with increasing temperature and phosphorus concentrations, reflecting that peak prevalence occurred in spring when temperature and phosphorus concentrations were relatively low. In contrast, seasonally-detrended prevalence increased with temperature, but decreased with increasing phosphorus concentrations. 4. Chytrids are a common component of the pelagic plankton community with sizeable biomass and removing an—at times—substantial proportion of the primary production, challenging the long-standing underrepresentation of parasites in ecological studies. 5. Chytrids responded differentially to seasonal variation and long-term trends in host density, water temperature and nutrient availability, highlighting the need to disentangle seasonal signals from long-term changes.

Datasets and R scripts for the COVID anthropause case study sites for the publication entitled "Lessons learned from the COVID-19 anthropause for water quality management"

SUMMARY: As human mobility decreased in 2020, the interaction between humans and nature changed significantly. On one hand, water clarity improved in the Amsterdam canals because boat traffic was reduced. On the other hand increased use of fishing water and national parks formed potential threats to the aquatic ecosystems. It is important to use these experiences to foster a more eco-centric mindset, building up to handling handling climate change and future pandemics. ABSTRACT: The anomalous past two years of the COVID-19 pandemic have been a test of human response to global crisis management as typical human activities were significantly altered. The COVID-instigated anthropause has illustrated the influence that humans and the biosphere have on each other, especially given the variety of national mobility interventions that have been implemented globally. These local COVID-19-era restrictions influenced human-ecosystem interactions through changes in accessibility of water systems and changes in ecosystem service demand. Four urban aquatic case studies in the Netherlands demonstrated shifts in human demand during the anthropause. For instance, reduced boat traffic in Amsterdam canals led to improved water clarity. In comparison, ongoing service exploitation from increased recreational fishing, use of bathing waters and national parks visitation are heightening concerns about potential ecosystem degradation. We distilled management lessons from both the case studies as well as from recent literature pertaining to ecological intactness and social relevance. Equally important to the lessons themselves, however, is the pace at which informed management practices are established after the pandemic ends, particularly as many communities currently recognize the importance of aquatic ecosystems and are amenable to their protection.

Fungal diseases threaten natural and man‐made ecosystems. Chytridiomycota (chytrids) infect a wide host range, including phytoplankton species that form the basis of aquatic food webs and produce roughly half of Earth's oxygen. However, blooms of large or toxic phytoplankton form trophic bottlenecks, as they are inedible to zooplankton. Chytrids infecting inedible phytoplankton provide a trophic link to zooplankton by producing edible zoospores of high nutritional quality. By grazing chytrid zoospores, zooplankton may induce a trophic cascade, as a decreased zoospore density will reduce new infections. Conversely, fewer infections will not produce enough zoospores to sustain long‐term zooplankton growth and reproduction. This intricate balance between zoospore density necessary for zooplankton energetic demands (growth/survival), and the loss in new infections (and thus new zoospores) because of grazing was tested empirically. To this end, we exposed a cyanobacterial host (Planktothrix rubescens) infected by a chytrid (Rizophydium megarrhizum) to a grazer density gradient (the rotifer Keratella cf. cochlearis). Rotifers survived and reproduced on a zoospore diet, but the Keratella population growth was limited by the amount of zoospores provided by chytrid infections, resulting in a situation where zooplankton survived but were restricted in their ability to control disease in the cyanobacterial host. We subsequently developed and parameterized a dynamical food‐chain model using an allometric relationship for clearance rate to assess theoretically the potential of different‐sized zooplankton groups to restrict disease in phytoplankton hosts. Our model suggests that smaller‐sized zooplankton may have a high potential to reduce chytrid infections on inedible phytoplankton. Together, our results point out the complexity of three‐way interactions between hosts, parasites, and grazers and highlight that trophic cascades are not always sustainable and may depend on the grazer's energetic demand.

Eukaryotic phytoplankton form the basis of aquatic food webs and play a key role in the global carbon cycle. Many of these evolutionarily diverse microalgae are also capable of feeding on other microbes, and hence simultaneously act both as primary producers and consumers. The net ecosystem impact of such mixotrophs depends on their nutritional strategy which is likely to alter with environmental change. Many temperate lakes are currently warming at unprecedented rates and are simultaneously increasing in water color (browning) due to increased run-off of humic substances. We hypothesized that the resulting reduction in light intensity and increased bacterial abundances would favor mixotrophic phytoplankton over obligate autotrophs, while higher temperatures might boost their rates of bacterivory. We tested these hypotheses in a mesocosm experiment simulating a gradient of increasing temperature and water color in temperate shallow lakes as expected to occur over the coming century. Mixotrophs showed a faster increase in abundance under the climate change scenario during spring, when they dominated the phytoplankton community. Furthermore, both bacterial abundances and rates of phytoplankton bacterivory increased under future climate conditions. Bacterivory contributed significantly to phytoplankton resource acquisition under future climate conditions, while remaining negligible throughout most of the season in treatments resembling today's conditions. Hence, to our knowledge, we here provide the first evidence for an increasing importance of bacterivory by phytoplankton in future temperate shallow lakes. Such a change in phytoplankton nutritional strategies will likely impact biogeochemical cycles and highlights the need to conceptually integrate mixotrophy into current ecosystem models.

Methane (CH4) strongly contributes to observed global warming. As natural CH4 emissions mainly originate from wet ecosystems, it is important to unravel how climate change may affect these emissions. This is especially true for ebullition (bubble flux from sediments), a pathway that has long been underestimated but generally dominates emissions. Here we show a remarkably strong relationship between CH4 ebullition and temperature across a wide range of freshwater ecosystems on different continents using multi-seasonal CH4 ebullition data from the literature. As these temperature–ebullition relationships may have been affected by seasonal variation in organic matter availability, we also conducted a controlled year-round mesocosm experiment. Here 4 °C warming led to 51% higher total annual CH4 ebullition, while diffusion was not affected. Our combined findings suggest that global warming will strongly enhance freshwater CH4 emissions through a disproportional increase in ebullition (6–20% per 1 °C increase), contributing to global warming., The impacts of climate change on natural methane (CH4) emissions via ebullition are unclear. Here, using published and experimental multi-seasonal CH4 ebullition data, the authors find a strong relationship between CH4 ebullition and temperature across a wide range of freshwater ecosystems globally.

Contains fulltext : 195210.pdf (Publisher’s version ) (Open Access) Abstract Temperatures have been rising throughout recent decades and are predicted to rise further in the coming century. Global warming affects carbon cycling in freshwater ecosystems, which both emit and bury substantial amounts of carbon on a global scale. Currently, most studies focus on the effect of warming on overall carbon emissions from freshwater ecosystems, while net effects on carbon budgets may strongly depend on burial in sediments. Here, we tested whether year-round warming increases the production, sedimentation, or decomposition of particulate organic carbon and eventually alters the carbon burial in a typical shallow freshwater system. We performed an indoor experiment in eight mesocosms dominated by the common submerged aquatic plant Myriophyllum spicatum testing two temperature treatments: a temperate seasonal temperature control and a warmed (+4°C) treatment (n = 4). During a full experimental year, the carbon stock in plant biomass, dissolved organic carbon in the water column, sedimented organic matter, and decomposition of plant detritus were measured. Our results showed that year-round warming nearly doubled the final carbon stock in plant biomass from 6.9 ± 1.1 g C in the control treatment to 12.8 ± 0.6 g C (mean ± SE), mainly due to a prolonged growing season in autumn. DOC concentrations did not differ between the treatments, but organic carbon sedimentation increased by 60% from 96 ± 9.6 to 152 ± 16 g C m?2 yaer?1 (mean ± SE) from control to warm treatments. Enhanced decomposition of plant detritus in the warm treatment, however, compensated for the increased sedimentation. As a result, net carbon burial was 40 ± 5.7 g C m?2 year?1 in both temperature treatments when fluxes were combined into a carbon budget model. These results indicate that warming can increase the turnover of organic carbon in shallow macrophyte-dominated systems, while not necessarily affecting net carbon burial on a system scale.

During the end of spring and throughout summer, large‐sized phytoplankton taxa often proliferate and form dense blooms in freshwater ecosystems. In many cases, they are inedible to zooplankton and prevent efficient transfer of energy and elements to higher trophic levels. Such a constraint may be alleviated by fungal parasite infections on large‐sized phytoplankton taxa like diatoms and filamentous cyanobacteria, as infections may provide zooplankton with a complementary food source in the form of fungal zoospores. Zoospores have been shown to support somatic growth of large filter feeding zooplankton species. Here, we tested if selectively feeding zooplankton, more specifically rotifers, also can use fungal zoospores as a food source. Our results show that chytrid fungal parasites can indeed support population growth of rotifers (Keratella sp.). Specifically, in cultures of an inedible filamentous cyanobacterium (Planktothrix rubescens), Keratella populations rapidly declined, while in Planktothrix cultures infected with chytrids, Keratella population growth rate equaled the growth observed for populations fed with a more suitable green algal diet (Chlorella sorokiniana). Feeding of Keratella on zoospores was furthermore indicated by a reduced number of zoospores during the last sampling day. These findings not only imply that rotifers may survive on zoospores, but also that the zoospores can support high rotifer population growth rates. We thus show that fungal parasites of inedible cyanobacteria can facilitate grazers by providing them alternative food sources. Together, these results highlight the important role that parasites may play in the aquatic plankton food web.

Internal phosphorus loading has become a major problem in many shallow freshwater lakes over the past decades due to the build-up of phosphorus stocks in the sediment. Iron is a natural capping agent which can enhance sediment P binding capacity, thus reducing P availability and shifting a lake from an algal to a macrophyte dominated state. Iron could, however, also impose toxic effects on the biota. We therefore provide a synopsis of iron toxicity studies and lake restoration measures using iron addition. Iron toxicity studies revealed that, even though iron is an essential nutrient for growth, when added in excess, it can negatively affect aquatic organisms. We found 13 studies testing the effect of iron addition as a restoration measure in the field (10) or using sediment from lakes and reservoirs in the laboratory (3). Twelve of the studies reported increased P retention after iron addition, which depended on the iron salts used and the concentrations added in two studies, whereas one study found no effect on P retention. Eight out of the nine field studies that reported biotic responses found reduced chlorophyll concentrations in the water column, whereas toxic effects of iron on organisms remained absent. Iron addition was most successful when external P loading, and concentrations of organic matter and sulphate were low as well as densities of sediment disturbing fish and crayfish. We conclude that iron addition can be a successful restoration method when these conditions are met.

Climate change is expected to favour infectious diseases across ecosystems worldwide. In freshwater and marine environments, parasites play a crucial role in controlling plankton population dynamics. Infection of phytoplankton populations will cause a transfer of carbon and nutrients into parasites, which may change the type of food available for higher trophic levels. Some phytoplankton species are inedible to zooplankton, and the termination of their population by parasites may liberate otherwise unavailable carbon and nutrients. Phytoplankton spring blooms often consist of large diatoms inedible for zooplankton, but the zoospores of their fungal parasites may serve as a food source for this higher trophic level. Here, we investigated the impact of warming on the fungal infection of a natural phytoplankton spring bloom and followed the response of a zooplankton community. Experiments were performed in ca. 1000 L indoor mesocosms exposed to a controlled seasonal temperature cycle and a warm (+4 °C) treatment in the period from March to June 2014. The spring bloom was dominated by the diatom Synedra. At the peak of infection over 40% of the Synedra population was infected by a fungal parasite (i.e. a chytrid) in both treatments. Warming did not affect the onset of the Synedra bloom, but accelerated its termination. Peak population density of Synedra tended to be lower in the warm treatments. Furthermore, Synedra carbon: phosphorus stoichiometry increased during the bloom, particularly in the control treatments. This indicates enhanced phosphorus limitation in the control treatments, which may have constrained chytrid development. Timing of the rotifer Keratella advanced in the warm treatments and closely followed chytrid infections. The chytrids' zoospores may thus have served as an alternative food source to Keratella. Our study thus emphasizes the importance of incorporating not only nutrient limitation and grazing, but also parasitism in understanding the response of plankton communities towards global warming.

Theory predicts that intraguild predation leads to different community dynamics than the trophic cascades of a linear food chain. However, experimental comparisons of these two food-web modules are rare. Mixotrophic plankton species combine photoautotrophic and heterotrophic nutrition by grazing upon other phytoplankton species. We found that the mixotrophic chrysophyte Ochromonas can grow autotrophically on ammonium, but not on nitrate. This offered a unique opportunity to compare predator–prey interactions in the presence and absence of intraguild predation, without changing the species composition of the community. With ammonium as nitrogen source, Ochromonas can compete with its autotrophic prey for nitrogen and therefore acts as intraguild predator. With nitrate, Ochromonas acts solely as predator, and is not in competition with its prey for nitrogen. We parameterized a simple intraguild predation model based on chemostat experiments with monocultures of Ochromonas and the toxic cyanobacterium Microcystis. Subsequently, we tested the model predictions by inoculating Ochromonas into the Microcystis monocultures, and vice versa. The results showed that Microcystis was a better competitor for ammonium than Ochromonas. In agreement with theoretical predictions, Microcystis was much more strongly suppressed by intraguild predation on ammonium than by top–down predation on nitrate. Yet, Microcystis persisted at very low population densities, because the type III functional response of Ochromonas implied that the grazing pressure upon Microcystis became low when Microcystis was rare. Our results provide experimental support for intraguild predation theory, and indicate that intraguild predation may enable biological control of microbial pest species.

Ecosystems can show sudden and persistent changes in state despite only incremental changes in drivers. Such critical transitions are difficult to predict, because the state of the system often shows little change before the transition. Early-warning indicators (EWIs) are hypothesized to signal the loss of system resilience and have been shown to precede critical transitions in theoretical models, paleo-climate time series, and in laboratory as well as whole lake experiments. The generalizability of EWIs for detecting critical transitions in empirical time series of natural aquatic ecosystems remains largely untested, however. Here we assessed four commonly used EWIs on long-term datasets of five freshwater ecosystems that have experienced sudden, persistent transitions and for which the relevant ecological mechanisms and drivers are well understood. These case studies were categorized by three mechanisms that can generate critical transitions between alternative states: competition, trophic cascade, and intraguild predation. Although EWIs could be detected in most of the case studies, agreement among the four indicators was low. In some cases, EWIs were detected considerably ahead of the transition. Nonetheless, our results show that at present, EWIs do not provide reliable and consistent signals of impending critical transitions despite using some of the best routinely monitored freshwater ecosystems. Our analysis strongly suggests that a priori knowledge of the underlying mechanisms driving ecosystem transitions is necessary to identify relevant state variables for successfully monitoring EWIs.

Until 1992, zebra mussels (Dreissena polymorpha) were an important food source for diving ducks in Lake Markermeer (The Netherlands). After 1993, the mussel biomass sharply declined, and the current population is in poor condition (maximum shell length 0.5 g L −1 . The NS concentrations used in this study are high, but analyses of the near-bottom TSM concentrations show that these values occur frequently and may be partly responsible for the poor mussel status in Lake Markermeer.

Diatom-based assays have been successfully associated worldwide with the tropic status of lakes. Several studies have demonstrated a correlation between epiphytic diatoms and nutrient load in shallow lakes and wetlands. We examine the relative importance of environmental factors in explaining the structure of epiphytic diatom communities in a set of Western European shallow lakes. The effects of lake physical/chemical, morphometric, and geographical variables on diatom assemblages were tested using distance-based, canonical correspondence and regression analyses. Our results show that epiphytic diatom communities respond, mainly to physical/chemical variables, overriding the effects of lake depth, size, and location. The clustering of studied systems based along a total phosphorus concentration gradient concurs with previous classification of diatom taxa and nutrient state. Assessment of epiphytic diatoms provided a potential method for shallow lake classification.

UV/H2O2 treatment can be part of the process converting surface water to drinking water, but would pose a potential problem when resulting in genotoxicity. This study investigates the genotoxicity of samples collected from the water treatment plant Andijk, applying UV/H2O2 treatment with an electrical energy dose of 0.54 kWh/m(3) and a H2O2 dose of 6 mg/l. Genotoxicity was tested in vitro using the Ames and Comet assay. All samples showed negative results. Samples were also tested in in vivo genotoxicity tests in Eastern mudminnow fish (Umbra pygmaea) by a sister chromatid exchange (SCE) and a Comet assay. No significant increases in SCEs were observed, but gill cells isolated from fish exposed to water obtained immediately after UV/H2O2 treatment and to Lake IJsselmeer water showed significantly increased DNA damage in the Comet assay. All other samples tested negative in this Comet assay. This indicates that DNA damaging compounds may result from the UV/H2O2 treatment, but also that these can be efficiently eliminated upon granular activated carbon (GAC) treatment of the water. It is concluded that when combined with this subsequent GAC treatment, UV/H2O2 treatment for the production of drinking water from surface water is not of concern with respect to genotoxicity.

Numerous studies show that increasing species richness leads to higher ecosystem productivity. This effect is often attributed to more efficient portioning of multiple resources in communities with higher numbers of competing species, indicating the role of resource supply and stoichiometry for biodiversity–ecosystem functioning relationships. Here, we merged theory on ecological stoichiometry with a framework of biodiversity–ecosystem functioning to understand how resource use transfers into primary production. We applied a structural equation model to define patterns of diversity–productivity relationships with respect to available resources. Meta-analysis was used to summarize the findings across ecosystem types ranging from aquatic ecosystems to grasslands and forests. As hypothesized, resource supply increased realized productivity and richness, but we found significant differences between ecosystems and study types. Increased richness was associated with increased productivity, although this effect was not seen in experiments. More even communities had lower productivity, indicating that biomass production is often maintained by a few dominant species, and reduced dominance generally reduced ecosystem productivity. This synthesis, which integrates observational and experimental studies in a variety of ecosystems and geographical regions, exposes common patterns and differences in biodiversity–functioning relationships, and increases the mechanistic understanding of changes in ecosystems productivity.

1. Different components of the climate system have been shown to affect temporal dynamics in natural plankton communities on scales varying from days to years. The seasonal dynamics in temperate lake plankton communities, with emphasis on both physical and biological forcing factors, were captured in the 1980s in a conceptual framework, the Plankton Ecology Group (PEG) model. 2. Taking the PEG model as our starting point, we discuss anticipated changes in seasonal and long-term plankton dynamics and extend this model to other climate regions, particularly polar and tropical latitudes. Based on our improved post-PEG understanding of plankton dynamics, we also evaluate the role of microbial plankton, parasites and fish in governing plankton dynamics and distribution. 3. In polar lakes, there is usually just a single peak in plankton biomass in summer. Lengthening of the growing season under warmer conditions may lead to higher and more prolonged phytoplankton productivity. Climate-induced increases in nutrient loading in these oligotrophic waters may contribute to higher phytoplankton biomass and subsequent higher zooplankton and fish productivity. 4. In temperate lakes, a seasonal pattern with two plankton biomass peaks – in spring and summer – can shift to one with a single but longer and larger biomass peak as nutrient loading increases, with associated higher populations of zooplanktivorous fish. Climate change will exacerbate these trends by increasing nutrient loading through increased internal nutrient inputs (due to warming) and increased catchment inputs (in the case of more precipitation). 5. In tropical systems, temporal variability in precipitation can be an important driver of the seasonal development of plankton. Increases in precipitation intensity may reset the seasonal dynamics of plankton communities and favour species adapted to highly variable environments. The existing intense predation by fish on larger zooplankters may increase further, resulting in a perennially low zooplankton biomass. 6. Bacteria were not included in the original PEG model. Seasonally, bacteria vary less than the phytoplankton but often follow its patterns, particularly in colder lakes. In warmer lakes, and with future warming, a greater influx of allochthonous carbon may obscure this pattern. 7. Our analyses indicate that the consequences of climate change for plankton dynamics are, to a large extent, system specific, depending on characteristics such as food-web structure and nutrient loading. Indirect effects through nutrient loading may be more important than direct effects of temperature increase, especially for phytoplankton. However, with warming a general picture emerges of increases in bacterivory, greater cyanobacterial dominance and smaller-bodied zooplankton that are more heavily impacted by fish predation.

1. Despite the strong impact parasites can have, only few models of phytoplankton ecology or aquatic food webs have specifically included parasitism. 2. Here, we provide a susceptible-infected model for a diatom-chytrid host–parasite system that explicitly includes nutrients, infected and uninfected hosts, reproduction of the parasite on the hosts and free-living infective stages. 3. A distinguishing feature of the model is that parasite reproduction on host increases with nutrient availability to the infected host, as has been observed for many parasites and viruses. 4. It follows from this assumption that the parasite’s basic reproduction number, R0, increases with nutrient concentration, because at higher nutrient concentrations, infected hosts consume more nutrients that are used for the reproduction of the parasite. 5. Another important result is that there may be two alternative states to which population densities can converge: one with only the host and one with host and parasite co-existing. In the latter, the parasite can invade a host population only if it is introduced above a threshold density. 6. Furthermore, the model shows a strong tendency for host–parasite cycles, which may be chaotic. Nutrient enrichment leads to increasing amplitude of these cycles, which may cause host or parasite population extinction caused by stochastic fluctuations during periods of low population density, which is the Paradox of Enrichment. 7. Finally, if alternative states and cycles co-occur, increased population cycle amplitude may drive the parasite below its threshold density for successful invasion, causing parasite extinction in a ‘deterministic Paradox of Enrichment’. Published results confirm that host–parasite cycles and collapse of host–parasite systems may occur in real plankton communities. 8. Our results underline that ecological detail in host–parasite models may have consequences for disease dynamics that may be overlooked when ecological interactions between environment, host and parasite are not explicitly taken into account.

SUMMARY 1. Many host–parasite interaction dynamics show distinct seasonality. Parasite population growth and invasion success are generally explained by host density dependence, while the direct influence of environmental factors on parasite life history traits has been underreported. 2. In waterbodies, resource availability and environmental conditions change with season (temperature, irradiance and rainfall patterns) and with depth (light, temperature and chemical gradients). Hence, hosts and parasites live in a spatially and temporally variable environment. Such environmental variation leads to structured populations, which in turn have implications for host–parasite interaction dynamics. Nevertheless, time-series data on the vertical distribution of aquatic hosts and their parasites are rare. 3. We present a dataset spanning 1.5 years (2008–2010) of weekly sampling in Lake Maarsseveen (the Netherlands) focussing on the dynamics of the diatom Asterionella formosa and its parasite, the chytrid Zygorhizidium planktonicum, at four depths. Environmental variables measured included ice cover, temperature, global irradiance, light extinction, pH, soluble reactive silicate (SRSi), dissolved nitrate and ortho-phosphate. 4. We observed four host blooms, two in early spring and one each in summer and autumn. Each host bloom was followed by a time-lagged parasite epidemic. Blooms and epidemics started in the uppermost water layers and showed a time lag in onset date with increasing depth. 5. Host abundance was related to SRSi, global irradiance, Schmidt stability and parasite abundance in the upper 10 m, whereas at 15 m only a relationship with parasite abundance prevailed. Parasite abundance was related to host abundance, light extinction, temperature, SRSi, stability and global irradiance within the upper 10 m; again, at 15 m, parasite abundance correlated only with host abundance and disease prevalence. 6. Host vertical distribution was less aggregated during isothermal conditions than during thermal stratification, when host abundance was higher in the mixed, photic epilimnion and lower in the dark, colder hypolimnion. Parasite vertical distribution was patchy most of the year. Parasite epidemics seemed to reduce host vertical patchiness as they impacted higher density patches in the photic zone more strongly, a result of both higher host abundance and favourable environmental conditions for the parasite. 7. Seasonal variability and vertical gradients in biotic and abiotic factors expose host and parasite individuals to different environmental conditions even within a single population. Environmental variability affects parasite transmission rates through changes in host abundance and through changes in the strength and outcome of host–parasite interactions.

1. Rising atmospheric CO2 concentrations might affect the primary production and community composition of freshwater ecosystems. 2. We investigated these potential effects in laboratory mesocosms (Limnotrons), using monoculture experiments and competition experiments with the green alga Scenedesmus obliquus and the cyanobacterium Synechocystis sp. PCC6803. The Limnotrons were sparged with ambient air (controls, 380 parts per million volume (ppmv) CO2), moderately elevated CO2 levels (3000 ppmv CO2) or highly elevated CO2 levels (18 800 ppmv CO2). 3. Growth at ambient air led to the depletion of dissolved CO2 during algal bloom development and hence a high pH. In contrast, growth at elevated CO2 levels resulted in high concentrations of dissolved CO2 and dissolved inorganic carbon (DIC), lower pH and low concentrations of nitrate and soluble reactive phosphorus. Elevated CO2 levels did not have a significant effect on the biomass of the algal species, but shifted their elemental composition towards higher carbon-to-nutrient ratios. 4. Competition experiments at ambient air were driven mainly by competition for inorganic carbon. In this case, the cyanobacterium Synechocystis was displaced by the green alga Scenedesmus. Elevated CO2 alleviated the community from carbon limitation, which shifted the species interactions towards competition for nitrogen and phosphorus, and resulted in coexistence of the two species. Hence, our findings do not support the hypothesis that cyanobacteria are generally better competitors for inorganic carbon than green algae. 5. In conclusion, our results show that elevated CO2 levels may cause major changes in freshwater ecosystems, including a reduction in pH, shifts in resource limitation patterns, and changes in the ecological stoichiometry and species composition of phytoplankton communities.

Although many lake restoration projects have led to decreased nutrient loads and increased water transparency, the establishment or expansion of macrophytes does not immediately follow the improved abiotic conditions and it is often unclear whether vegetation with high macrophyte diversity will return. We provide an overview of the potential bottlenecks for restoration of submerged macrophyte vegetation with a high biodiversity and focus on the biotic factors, including the availability of propagules, herbivory, plant competition and the role of remnant populations. We found that the potential for restoration in many lakes is large when clear water conditions are met, even though the macrophyte community composition of the early 1900s, the start of human-induced large-scale eutrophication in Northwestern Europe, could not be restored. However, emerging charophytes and species rich vegetation are often lost due to competition with eutrophic species. Disturbances such as herbivory can limit dominance by eutrophic species and improve macrophyte diversity. We conclude that it is imperative to study the role of propagule availability more closely as well as the biotic interactions including herbivory and plant competition. After abiotic conditions are met, these will further determine macrophyte diversity and define what exactly can be restored and what not.

Macrophytes and nutrient loading are two factors that can strongly determine the diversity and composition of aquatic invertebrate communities. Both factors may also interact, because macrophyte species may be differentially affected by nutrients. Macrophyte community characteristics, such as species composition, morphotype and biomass have the potential to mediate the response of invertebrate communities to nutrient loading. In 36 newly constructed experimental ponds, we orthogonally combined three macrophyte community types (Chara-, Potamogeton- and Elodea-dominated) with two levels of nutrient additions (no addition and an addition of 0.5 mg P and 3 mg N/L per week) and studied community assembly in three functional groups of invertebrates (epiphytic macroinvertebrates, littoral and pelagic crustacean zooplankton). Macrophyte biomass was negatively affected by nutrient addition. General linear models indicated negative responses of species richness in the zooplankton functional groups to nutrient addition and phytoplankton chlorophyll-a, but demonstrated no effects of macrophyte community type. Conversely, macroinvertebrate taxon richness differed among macrophyte community types but showed no response to nutrient enrichment. Macrophyte biomass correlated positively with the richness of littoral zooplankton and macroinvertebrates and was a better predictor of these diversity variables than macrophyte community type. Overall, our results indicate that lake management practices that aim at obtaining a nutrient poor and macrophyte dominated clear water state contribute also to the maintenance of aquatic invertebrate diversity.

1. We describe the dynamics of host-parasite interactions over a period of more than 30 years between the freshwater diatom Asterionella formosa and two highly virulent chytrid parasites (Rhizophydium planktonicum and Zygorhizidium planktonicum) in Lake Maarsseveen, The Netherlands. This period is characterised by a significant warming trend which is strongest in spring. 2. The key spring event in lakes, the diatom bloom, was in many years dominated by Asterionella. We examine whether and how climate warming has affected the prevalence of infection in Asterionella by chytrids. 3. In years with cold winters/early springs, a dense Asterionella bloom is followed by epidemic development of disease as high Asterionella densities greatly facilitate transmission of chytrid zoospores. This sequence of events is absent in milder winters. 4. Earlier experimental studies have shown that the parasite is almost non-infective at water temperatures below 3 degrees C, offering a disease-free window of opportunity for growth of Asterionella. Climate warming has reduced periods in which water temperature remains < 3 degrees C, narrowing the window of opportunity for uninfected growth. Consequently, Asterionella continuously suffers from infection, albeit at low levels. 5. Population reduction as a result of low level infection allows other diatoms to take over as dominant species, possibly through priority effects. 6. In mild winters, chytrid infections no longer reach epidemic levels, but remain at low prevalence since transmission is impaired at low host densities. Climate warming thus affects both host and parasite in intricate ways, with the host denied a bloom and consequently the parasite denied an epidemic. 7. A shift from Asterionella to a mixed diatom community in years with mild winters may benefit the food web, because of the poor edibility of Asterionella, unless the numerous chytrid zoospores produced during epidemics significantly contribute to zooplankton nutrition. 8. Our study demonstrates the potential complexity of climate change impacts on disease. A reduction in the likelihood of epidemic development of a virulent parasite would seem to be of great benefit to the host, but this was not the case. Unexpected, sometimes paradoxical consequences of climate change can be expected and suggest that the view of a 'warmer hence sicker world' may not always apply.

Planktothrix agardhii is a widespread harmful cyanobacterium of eutrophic waters, and can produce the hepatotoxins [Asp3]microcystin-LR and [Asp3]microcystin-RR. These two microcystin variants differ in their first variable amino acid position, which is occupied by either leucine (L) or arginine (R). Although microcystins are extensively investigated, little is known about the mechanisms that determine the production of different microcystin variants. We hypothesize that enhanced nitrogen availability will increase the intracellular content of the nitrogen-rich amino acid arginine, and thereby promote the production of the variant [Asp3]microcystin-RR. To test this hypothesis, we transferred P. agardhii strain 126/3 from nitrogen-replete to nitrogen-deficient conditions, and after 2 weeks of growth under nitrogen deficiency, we added a nitrogen pulse. We found a rapid increase in the cellular nitrogen to carbon ratio and the amino acids aspartic acid and arginine, indicative of cyanophycin synthesis. This was followed by a more gradual increase of the total amino acid content connected to balanced growth. As expected, the [Asp3]microcystin-RR variant increased strongly after the nitrogen pulse, while the [Asp3]microcystin-LR increased to a much lesser extent. We conclude that sudden nitrogen enrichment affects the amino acid composition of harmful cyanobacteria, which, in turn, affects the production and composition of their microcystins.

Diatoms form large spring blooms in lakes and oceans, providing fuel for higher trophic levels at the start of the growing season. Some of the diatom blooms, however, are not grazed by filter-feeding zooplankton like Daphnia due to their large size. Several of these large diatoms are susceptible to chytrid infections. Zoospores of chytrids appeared to be excellent food for Daphnia, both in terms of size, shape, and quality (PUFAs and cholesterol). Thus, zoospores of chytrids can bridge the gap between inedible diatoms and Daphnia. In order to examine the effects of diatoms and chytrids on the survival of copepods, we performed one grazing and one survival experiment. The grazing experiment revealed that the diatom, Asterionella formosa, was not grazed by the copepod, Eudiaptomus gracilis, even after being infected by the chytrid Zygorhizidium planktonicum. However, carbon and nitrogen concentrations were significantly reduced by E. gracilis only when A. formosa was infected by Z. planktonicum, indicating that the chytrids might facilitate material transfer from inedible diatoms to the copepods. The survival experiment revealed that E. gracilis lived shorter with A. formosa than with the cryptophyta Cryptomonas pyrenoidifera. However, the survival of E. gracilis increased significantly in the treatment where A. formosa cells were infected by Z. planktonicum. Since E. gracilis could not graze A. formosa cells due to their large colonial forms, E. gracilis may acquire nutrients by grazing on the zoospores, and were so able to survive in the presence of the A. formosa. This provides new insights into the role of parasitic fungi in aquatic food webs, where chytrids may improve copepod survival during diatom blooms.

Advances in ecological stoichiometry, a rapidly expanding research field investigating the elemental composition of organisms and their environment, have shed new light on the impacts of climate change on freshwater and marine ecosystems. Current changes in the Earth's climate alter the availability of carbon and nutrients in lakes and oceans. In particular, atmospheric CO2 concentrations will rise to unprecedented levels by the end of this century, while global warming will enhance stratification of aquatic ecosystems and may thereby diminish the supply of nutrients into the surface layer. These processes enrich phytoplankton with carbon, but suppress nutrient availability. Phytoplankton with a high carbon-to-nutrient content provide poor-quality food for most zooplankton species, which may shift the species composition of zooplankton and higher trophic levels to less nutrient-demanding species. As a consequence, climate-driven changes in plankton stoichiometry may alter the structure and functioning of entire aquatic food webs.

The elemental composition of primary producers reflects the availability of light, carbon and nutrients in their environment. According to the carbon-nutrient balance hypothesis, this has implications for the production of secondary metabolites. To test this hypothesis, we investigated a family of toxins, known as microcystins, produced by harmful cyanobacteria. The strain Microcystis aeruginosa HUB 5-2-4, which produces several microcystin variants of different N:C stoichiometry, was cultured in chemostats supplied with various combinations of nitrate and CO(2). Excess supply of both nitrogen and carbon yielded high cellular N:C ratios accompanied by high cellular contents of total microcystin and the nitrogen-rich variant microcystin-RR. Comparable patterns were found in Microcystis-dominated lakes, where the relative microcystin-RR content increased with the seston N:C ratio. In total, our results are largely consistent with the carbon-nutrient balance hypothesis, and warn that a combination of rising CO(2) and nitrogen enrichment will affect the microcystin composition of harmful cyanobacteria.

While there is a general sense that lakes can act as sentinels of climate change, their efficacy has not been thoroughly analyzed. We identified the key response variables within a lake that act as indicators of the effects of climate change on both the lake and the catchment. These variables reflect a wide range of physical, chemical, and biological responses to climate. However, the efficacy of the different indicators is affected by regional response to climate change, characteristics of the catchment, and lake mixing regimes. Thus, particular indicators or combinations of indicators are more effective for different lake types and geographic regions. The extraction of climate signals can be further complicated by the influence of other environmental changes, such as eutrophication or acidification, and the equivalent reverse phenomena, in addition to other land-use influences. In many cases, however, confounding factors can be addressed through analytical tools such as detrending or filtering. Lakes are effective sentinels for climate change because they are sensitive to climate, respond rapidly to change, and integrate information about changes in the catchment.

The effects of inducible defenses and constitutive defenses on population dynamics were investigated in a freshwater plankton system with rotifers as predators and different algal strains as prey. We made predictions for these systems using a chemostat predator-prey model and focused on population stability and predator persistence as a function of flow-through rate. The model exhibits three major types of behavior at a high nutrient concentration: (1) at high dilution rates, only algae exist; (2) at intermediate dilution rates, algae and rotifers show stable coexistence; (3) at low dilution rates, large population fluctuations occur, with low minimum densities entailing a risk of stochastic rotifer extinctions. The size and location of the corresponding areas in parameter space critically depend on the type of algal defense strategy. In an 83-day high-nutrient chemostat experiment we changed the dilution rate every 3 weeks, from 0.7 to 0.5 to 0.3 to 0.1 per day. Within this range of dilution rates, rotifers and algae coexisted, and population fluctuations of algae clearly increased as dilution rates decreased. The CV of herbivore densities was highest at the end of the experiment, when the dilution rate was low. On day 80, herbivorous rotifers had become undetectable in all three chemostats with permanently defended algae (where rotifer densities had already been low) and in two out of three chemostats where rotifers had been feeding on algae with inducible defenses (that represented more edible food). We interpret our results in relation to the paradox of enrichment. © The Ecological Society of Japan 2009.

Replicated, factorial mesocosm experiments were conducted across Europe to study the effects of nutrient enrichment and fish density on macrophytes and on periphyton chlorophyll a (chl-a) with regard to latitude. Periphyton chl-a densities and plant decline were significantly related to nutrient loading in all countries. Fish effects were significant in a few sites only, mostly because of their contribution to the nutrient pool. A saturation-response type curve in periphyton chl-a with nutrients was found, and northern lakes achieved higher densities than southern lakes. Nutrient concentration and phytoplankton chl-a necessary for a 50% plant reduction followed a latitudinal gradient. Total phosphorus values for 50% plant disappearance were similar from Sweden (0.27 mg L−1) to northern Spain (0.35 mg L−1), but with a sharp increase in southern Spain (0.9 mg L−1). Planktonic chl-a values for 50% plant reduction increased monotonically from Sweden (30 μg L−1) to Valencia (150 μg L−1). Longer plant growing-season, higher light intensities and temperature, and strong water-level fluctuations characteristic of southern latitudes can lead to greater persistence of macrophyte biomass at higher turbidities and nutrient concentration than in northern lakes. Results support the evidence that latitudinal differences in the functioning of shallow lakes should be considered in lake management and conservation policies.

The structure of aquatic ecosystems is determined by complex interactions among individual organisms at different trophic levels. Although our basic understanding of how top-down and bottom-up processes interact to determine food-web dynamics has advanced, we still lack insights into how complex interactions and feedbacks affect the dynamics and structure of food webs. It is now becoming increasingly clear that, in addition to energy transfer from one trophic level to the other, there is exchange of information between these levels facilitated by the release of infochemicals by the organisms. There is evidence from recent studies that the exchange of chemical information in freshwater ecosystems is likely to play a decisive role in shaping structure and functioning of these systems. Chemical communication among freshwater organisms mediates many aspects of both predation and interspecific competition, which play key roles in determining community structure and ecosystem functioning. For example, consumer-induced defences in phytoplankton and zooplankton include modifications in the characteristics relating to life history, behaviour, morphology and biochemistry. These inducible defences affect trophic interactions by altering predator feeding rates through changes in attack rate or handling time, or both. Also host-specific fungal parasitism in phytoplankton is probably controlled by infochemicals. The motile fungi recognise their host by host-secreted compounds. Until now models describing the functioning of ecosystems mainly considered flows of biomass and energy. Integration of new knowledge about the role of chemical communication in these models may be one of the aims of ecological informatics. In this chapter I discuss how infochemicals may affect the dynamics and structure of planktonic food webs.

Submerged macrophytes are known to serve as refuge for zooplankton but also seem to suppress the zooplankton growth. Thus, there is a conflict between the positive and negative role of macrophytes for zooplankton. We tested the influence of physical structure using artificial macrophytes, chemicals released by a macrophyte (Myriophyllum verticillatum), as well as the combined effects of these two factors on the life history of Daphnia magna. Daphnids matured at a smaller size and produced fewer eggs but larger individual offspring in the presence of artificial and real plants. Thus, under conditions with plants, we observed a trade off between number of eggs produced and the individual size of the offspring. Daphnids grown in the presence of exudates without plants were larger at maturity and showed no reduction in clutch size as compared with the control. We suggest that the macrophytes (real and artificial) negatively affected the daphnids in two ways: (1) food particles were settling down faster on the plant structures and this reduced the available food for Daphnia, (2) the plant structures were obstacles for the daphnids causing them to spent more energy during swimming. Both effects resulted in a reduced somatic growth, whereas the increased individual offspring size was probably a response to the reduced food level. Daphnia was not negatively affected by Myriophyllum exudates.

We studied grazing on the green alga Scenedesmus obliquus and the cyanobacterium Planktothrix agardhii (a microcystin-producing and a microcystin- free strain) by three different filter-feeders to compare the efficiency with which these grazers remove Planktothrix from the water. The filterfeeders were two mussel species (the invasive species Dreissena polymorpha and the native Anodonta anatina) and one zooplankton species (Daphnia galeata). On a weight-specific basis, D. galeata had the highest observed clearance rates and A. anatina the lowest. However, D. galeata was only able to feed on Scenedesmus and not on Planktothrix, probably as a consequence of the morphology of Planktothrix (filaments), since neither microcystin-producing nor microcystin-free strains were grazed by Daphnia. Both mussel species were able to graze on Planktothrix, irrespective of microcystin content. Dreissena showed an especially high clearance rate. On an individual basis, however, Anodonta showed the highest clearance rates, which are a consequence of its large biomass. Densities of Anodonta in the field, however, are presently too low to exert a high grazing pressure on the phytoplankton. We suggest therefore that water managers, who may be interested in using filter-feeders as a biomanipulation tool, focus on the improvement of settlement conditions for native bivalves, such as Anodonta, instead of exotic species like Dreissena.

Pharmaceuticals consumption by humans and animals is increasing substantially, leading to unprecedented levels of these compounds in aquatic environments worldwide. Recent findings that concentrations reach levels that can directly have negative effects on organisms are important per se, but also sound an alarm for other potentially more pervasive effects that arise from the interconnected nature of ecological communities. Aquatic organisms use chemical cues to navigate numerous challenges, including the location of mates and food, and the avoidance of natural enemies. Low concentrations of pharmaceuticals can disrupt this "smellscape" of information leading to maladaptive responses. Furthermore, direct effects of pharmaceuticals on the traits and abundance of one species can cascade through a community, indirectly affecting other species. We review mechanisms by which pharmaceuticals in surface waters can disrupt natural chemical information flows and species interactions. Pharmaceuticals form a new class of chemical threats, which could have far-reaching implications for ecosystem functioning and conservation management.

Background In recent decades, many attempts have been made to restore eutrophic lakes through biomanipulation. Reducing the populations of planktivorous and benthivorous fish (either directly or through stocking of piscivorous fish) may induce ecosystem changes that increase water transparency and decrease the risk of algal blooms and fish kills, at least in the short term. However, the generality of biomanipulation effects on water quality across lake types and geographical regions is not known. Therefore, we have undertaken a systematic review of such effects in eutrophic lakes in temperate regions throughout the world. Methods Searches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Danish, Dutch and Swedish. Identified articles were screened for relevance using inclusion criteria set out in an a priori protocol. To reduce the risk of bias, we then critically appraised the combined evidence found on each biomanipulation. Data were extracted on outcomes such as Secchi depth and chlorophyll a concentration before, during and/or after manipulation, and on effect modifiers such as lake properties and amounts of fish removed or stocked. Results Our searches identified more than 14,500 articles. After screening for relevance, 233 of them remained. After exclusions based on critical appraisal, our evidence base included useful data on 128 biomanipulations in 123 lakes. Of these interventions, 85% had been made in Europe and 15% in North America. Meta-analysis showed that removal of planktivores and benthivores (with or without piscivore stocking) leads to increased Secchi depth and decreased chlorophyll a concentration during intervention and the first three years afterwards. Piscivore stocking alone has no significant effect. The response of chlorophyll a levels to biomanipulation is stronger in lakes where fish removal is intense, and in lakes which are small and/or have high pre-manipulation concentrations of total phosphorus. Conclusions Our review improves on previous reviews of biomanipulation in that we identified a large number of case studies from many parts of the world and used a consistent, repeatable process to screen them for relevance and susceptibility to bias. Our results indicate that removal of planktivorous and benthivorous fish is a useful means of improving water quality in eutrophic lakes. Biomanipulation tends to be particularly successful in relatively small lakes with short retention times and high phosphorus levels. More thorough fish removal increases the efficacy of biomanipulation. Nonetheless successes and failures have occurred across a wide range of conditions.