Droevendaalsesteeg 10
6708 PB Wageningen
The Netherlands
Ellen van Donk
Netwerk
Over
She has been interested in how ecological mechanisms and abiotic factors govern the dynamics and structure of food webs in aquatic ecosystems, acknowledging the demands from society for scientific input for nature conservation and ecosystem restoration
Biografie
She graduated from the University of Amsterdam in 1983 (PhD in Aquatic Ecology) and visited the University of Michigan (Ann Arbor, USA) for several months in 1981. From 1983 until 1990, she worked as the Head of the Research Department at the Water Board in the State of Utrecht, examining lakes and rivers and applying restoration measures to these waters. Thereafter, she was appointed as an Associate Professor at the Department of Aquatic Ecology and Water Quality Management of the University of Wageningen (1990 -1998). Here she continued her research, along with teaching and supervising the research of numerous graduate and undergraduate students. She joined the NIOO-KNAW (Netherlands Institute for Ecology) in 1998 as Head of the Department of Aquatic Ecology where she continues to examine her main research interests in plankton dynamics and ecology, phytoplankton succession, lake eutrophication and ecosystem stress, and food web studies. From 2000- 2009 she had a professorship in Limnology at the University of Nijmegen (The Netherlands) on a one-day-a-week basis, which was continued at the University of Utrecht (now as emeritus). From 2001-2011 she also had a visiting professorship in Aquatic Ecology at the University of Oslo (Norway). After her step down as Head Department Aquatic Ecology, due to retirement (Oct. 2019), she continued her work at NIOO as researcher and advisor within the Translational Science Unit AKWA.
CV
Nevenfuncties
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Professor in Aquatic Ecology
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Peer-reviewed publicaties
Serving many masters at once
https://doi.org/10.1080/20442041.2021.1944765Globally the number of relatively deep, isolated lakes is increasing because of sand, gravel, or clay excavation activities. The major excavation areas are located within the delta of rivers, and thus the deep freshwater ecosystems formed upon excavation, called quarry lakes, are unique to the landscape. They are embedded in a landscape comprised of shallow, naturally formed lakes. Given that quarry lakes are by definition novel ecosystems, water managers face difficulties in optimally managing them to deliver ecosystem services using existing frameworks designed for natural ecosystems. All lakes in delta areas are subject to similar pressures such as urbanization and eutrophication, leading to shifts in biodiversity and ecosystem functioning, and ultimately changing the ecosystem services the systems can provide. We propose a framework to enable water managers to assess the provision of ecosystem services by quarry lakes based on their ecological quality. For each ecosystem service we determined threshold values of ecological quality based on available scientific literature, an extensive field survey of 51 quarry lakes in the Netherlands, or expert knowledge. To illustrate the usefulness of our approach, we applied our framework to a lake before and after a rehabilitation focused on improving the nutrient status of the waterbody. Assessing ecosystem services under varying levels of ecological health is important to initiate action from legislators, managers, and communities.
Differential effects of elevated pCO2 and warming on marine phytoplankton stoichiometry
Phytoplankton stand at the base of the marine food-web, and play a major role in global carbon cycling. Rising CO2 levels and temperatures are expected to enhance growth and alter carbon:nutrient stoichiometry of marine phytoplankton, with possible consequences for the functioning of marine food-webs and the oceanic carbon pump. To date, however, the consistency of phytoplankton stoichiometric responses remains unclear. We therefore performed a meta-analysis on data from experimental studies on stoichiometric responses of marine phytoplankton to elevated pCO2 and 3–5° warming under nutrient replete and limited conditions. Our results demonstrate that elevated pCO2 increased overall phytoplankton C:N (by 4%) and C:P (by 9%) molar ratios under nutrient replete conditions, as well as phytoplankton growth rates (by 6%). Nutrient limitation amplified the CO2 effect on C:N and C:P ratios, with increases to 27% and 17%, respectively. In contrast to elevated pCO2, warming did not consistently alter phytoplankton elemental composition. This could be attributed to species- and study-specific increases and decreases in stoichiometry in response to warming. While our observed moderate CO2-driven changes in stoichiometry are not likely to drive marked changes in food web functioning, they are in the same order of magnitude as current and projected estimations of oceanic carbon export. Therefore, our results may indicate a stoichiometric compensation mechanism for reduced oceanic carbon export due to declining primary production in the near future.https://doi.org/10.1002/lno.12020Distribution patterns of epiphytic reed-associated macroinvertebrate communities across European shallow lakes
https://doi.org/10.1016/j.scitotenv.2020.144117So far, research on plant-associated macroinvertebrates, even if conducted on a large number of water bodies, has mostly focused on a relatively small area, permitting limited conclusions to be drawn regarding potentially broader geographic effects, including climate. Some recent studies have shown that the composition of epiphytic communities may differ considerably among climatic zones. To assess this phenomenon, we studied macroinvertebrates associated with the common reed Phragmites australis (Cav.) Trin. ex Steud in 46 shallow lakes using a common protocol. The lakes, located in nine countries, covered almost the entire European latitudinal range (from <48°N to 61°N) and captured much of the variability in lake size and nutrient content in the region. A Poisson Generalized Linear Mixed Model (GLMM) showed the number of macroinvertebrate epiphytic taxa to be negatively associated with water conductivity and positively associated with medium ice cover duration (approximately 1 month). A Gamma GLMM showed a positive effect of chlorophyll a on the density of macroinvertebrates, and a significantly greater density in lakes located at the lowest and highest latitudes. Individual taxa responded differently to lake environmental conditions across climate zones. Chironomidae dominated in all climate zones, but their contribution to total density decreased with increasing latitude, with progressively greater proportions of Naidinae, Asellidae, Ephemeroptera and Trichoptera. Our study demonstrates that epiphytic macroinvertebrate fauna, even when analyzed at low taxonomic resolution, exhibits clear differences in diversity, relative abundance of individual taxa and total density, shaped both by geographic and anthropogenic variables. The results were discussed in the context of climate change. To our best knowledge this is the first study to examine epiphytic fauna carried out on a European scale.
Direct and indirect effects of native plants and herbivores on biotic resistance to alien aquatic plant invasions
Biotic resistance to alien plant invasions is mainly determined by ecological interactions in two layers of the food web: competition with native plant species and herbivory by native herbivores. While the direct effect of native plants on alien plant performance via competition has been well documented across ecosystems, less is known about the direct and indirect effects of herbivores in providing biotic resistance. Our main aims were to determine whether temperate native aquatic plants and herbivores can provide biotic resistance to plant invasions, understand the underlying mechanisms and search for potential interactive effects of competition and herbivory on invader performance (i.e. growth).https://doi.org/10.1111/1365-2745.13380
We mimicked natural temperate mesotrophic and eutrophic freshwater lakes in mesoscosms, by growing three native submerged plant species in monocultures (Ceratophyllum demersum, Myriophyllum spicatum and Potamogeton perfoliatus) at three competition levels (no, low and high) without and with the native aquatic generalist snail Lymnaea stagnalis. We subsequently simulated an early stage of establishment of the South American highly invasive alien plant species Egeria densa.
We found that competition by native plant biomass significantly reduced invader performance but depended on native species identity. Herbivory had no direct negative effect on invader performance as the snails fed mainly on the available filamentous algae, which are commonly found in meso‐ and eutrophic systems, instead of on the plants. However, the consumption of filamentous algae by herbivores indirectly had positive effects on the invader total biomass, thus facilitating the invasion by E. densa. Nonetheless, these indirect effects worked through different pathways depending on the native plant identity.
Synthesis. We found evidence for biotic resistance through competition by native plant species. However, we show that herbivores can indirectly facilitate South American plant E. densa invasion promoting its growth through selective feeding on filamentous algae, but this effect depends on the native plant species involved. Our experiment illustrates the important role of indirect interactions to understand the potential of biotic resistance in natural ecosystems.Warming advances virus population dynamics in a temperate freshwater plankton community
Viruses are important drivers in the cycling of carbon and nutrients in aquatic ecosystems. Since viruses are obligate parasites, their production completely depends on growth and metabolism of hosts and therefore can be affected by climate change. Here, we investigated if warming (+4°C) can change the outcome of viral infections in a natural freshwater virus community over a 5‐month period in a mesocosm experiment. We monitored dynamics of viruses and potential hosts. Results show that warming significantly advanced the early summer peak of the virus community by 24 d, but neither affected viral peak abundances nor time‐integrated number of viruses present. Our results demonstrate that warming advances the timing of viruses in a natural community. Although warming may not necessarily result in a stronger viral control of bacterial and phytoplankton communities, our results suggest it can alter host population dynamics through advanced timing of infections, and thus timing of carbon and nutrient recycling.https://doi.org/10.1002/lol2.10160The potential of zooplankton in constraining chytrid epidemics in phytoplankton hosts
Abstract Fungal diseases threathen 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) due to 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 was restricted in their ability to control disease in the cyanobacterial host. We subesequently developed and parameterized a dynamical food-chain model using an allometric relationship for clearance rate to theoretically assess 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 tri-way interactions between hosts-parasites-grazers, and highlight that trophic cascades are not always sustainable and may depend on the grazer's energetic demand.https://doi.org/10.1002/ecy.2900Trophic position, elemental ratios and nitrogen transfer in a planktonic host–parasite–consumer food chain including a fungal parasite
Parasitism is arguably the most commonly occurring consumer strategy. However, only a few food web studies assess how well stable isotopes reflect the trophic position of parasitic consumers and results are variable. Even fewer studies have measured the nutrient transfer by parasitic consumers, hindering an assessment of their role in nutrient transfer through food webs. Here we used a food chain consisting of a diatom as host, a chytrid as its parasitic consumer and a rotifer as the predatory consumer of the chytrid, to assess the trophic position of all three food-chain components using their natural 13C and 15N isotope signatures, and to measure the nitrogen transfer from the host via the chytrid to the rotifer by tracing 15N of a labelled host up the food chain. Additionally, we measured the carbon to nitrogen (C:N) ratios of all food-chain components. Natural isotope abundance results showed no clear 15N enrichment in the chytrid or rotifer relative to the primary producer. However, estimates of nitrogen transfer indicated that about 14% of host nitrogen was transferred per day from host to chytrid during infection epidemics and that some of this nitrogen was also transferred onward to the rotifer. Moreover, C:N ratios decreased with trophic level, suggesting that the chytrid provided a high-quality food source to the rotifer. In conclusion, our results support the “mycoloop”. The mycooloop proposes that chytrid infections allow the transfer of nutrients bound in large, inedible phytoplankton to zooplankton through the production of edible transmission spores, thereby rerouting nutrients back into the food web.https://doi.org/10.1007/s00442-020-04721-wImpacts of warming on top-down and bottom-up controls of periphyton production
Global warming profoundly impacts the functioning of aquatic ecosystems. Nonetheless, the effect of warming on primary producers is poorly understood, especially periphyton production, which is affected both directly and indirectly by temperature-sensitive top-down and bottom-up controls. Here, we study the impact of warming on gross primary production in experimental ecosystems with near-realistic foodwebs during spring and early summer. We used indoor mesocosms following a temperate temperature regime (control) and a warmed (+4 °C) treatment to measure biomass and production of phytoplankton and periphyton. The mesocosms’ primary production was dominated by periphyton (>82%) during the studied period (April-June). Until May, periphyton production and biomass were significantly higher in the warm treatment (up to 98% greater biomass compared to the control) due to direct temperature effects on growth and indirect effects resulting from higher sediment phosphorus release. Subsequently, enhanced grazer abundances seem to have counteracted the positive temperature effect causing a decline in periphyton biomass and production in June. We thus show, within our studied period, seasonally distinct effects of warming on periphyton, which can significantly affect overall ecosystem primary production and functioning.https://doi.org/10.1038/s41598-018-26348-xPrimary producers or consumers? Increasing phytoplankton bacterivory along a gradient of lake warming and browning
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.https://doi.org/10.1002/lno.10728Fungal parasites of a toxic inedible cyanobacterium provide food to zooplankton
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.https://doi.org/10.1002/lno.10945Warming enhances sedimentation and decomposition of organic carbon in shallow macrophyte-dominated systems with zero net effect on carbon burial
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.https://doi.org/10.1111/gcb.14387Response of Submerged Macrophyte Communities to External and Internal Restoration Measures in North Temperate Shallow Lakes
Submerged macrophytes play a key role in north temperate shallow lakes by stabilising clear-water conditions. Eutrophication has resulted in macrophyte loss and shifts to turbid conditions in many lakes. Considerable efforts have been devoted to shallow lake restoration in many countries, but long-term success depends on a stable recovery of submerged macrophytes. However, recovery patterns vary widely and remain to be fully understood. We hypothesize that reduced external nutrient loading leads to an intermediate recovery state with clear spring and turbid summer conditions similar to the pattern described for eutrophication. In contrast, lake internal restoration measures can result in transient clear-water conditions both in spring and summer and reversals to turbid conditions. Furthermore, we hypothesize that these contrasting restoration measures result in different macrophyte species composition, with added implications for seasonal dynamics due to differences in plant traits. To test these hypotheses, we analysed data on water quality and submerged macrophytes from 49 north temperate shallow lakes that were in a turbid state and subjected to restoration measures. To study the dynamics of macrophytes during nutrient load reduction, we adapted the ecosystem model PCLake. Our survey and model simulations revealed the existence of an intermediate recovery state upon reduced external nutrient loading, characterised by spring clear-water phases and turbid summers, whereas internal lake restoration measures often resulted in clear-water conditions in spring and summer with returns to turbid conditions after some years. External and internal lake restoration measures resulted in different macrophyte communities. The intermediate recovery state following reduced nutrient loading is characterised by a few macrophyte species (mainly pondweeds) that can resist wave action allowing survival in shallow areas, germinate early in spring, have energy-rich vegetative propagules facilitating rapid initial growth and that can complete their life cycle by early summer. Later in the growing season these plants are, according to our simulations, outcompeted by periphyton, leading to late-summer phytoplankton blooms. Internal lake restoration measures often coincide with a rapid but transient colonisation by hornworts, waterweeds or charophytes. Stable clear-water conditions and a diverse macrophyte flora only occurred decades after external nutrient load reduction or when measures were combined.https://doi.org/10.3389/fpls.2018.00194Environmental risk assessment for invasive alien species: A case study of apple snails affecting ecosystem services in Europe
Abstract The assessment of the risk posed by invasive alien species (IAS) to the environment is a component of increasing importance for Pest Risk Analysis. Standardized and comprehensive procedures to assess their impacts on ecosystem services have been developed only recently. The invasive apple snails (Pomacea canaliculata and P. maculata) are used as a case study to demonstrate the application of an innovative procedure assessing the potential impact of these species on shallow freshwater ecosystems with aquatic macrophytes in Europe. The apple snail, Pomacea maculata, recently established in the Ebro delta in Spain resulting in a serious threat to rice production and wetlands, having also a high risk to spread to other European wetlands. Here, the population abundance of apple snails is regarded as the main driver of ecosystem change. The effects of ecosystem resistance, resilience and pest management on snail population abundance are estimated for the short (5 years) and the long (30 years) term. Expert judgment was used to evaluate the impacts on selected ecosystem services in a worst-case scenario. Our study shows that the combined effects of apple snails are estimated to have profound effects on the ecosystem services provided by shallow, macrophyte-dominated ecosystems in Europe. This case study illustrates that quantitative estimates of environmental impacts from different IAS are feasible and useful for decision-makers and invasive species managers that have to balance costs of control efforts against environmental and economic impacts of invasive species.https://doi.org/10.1016/j.eiar.2017.03.008Combined physical, chemical and biological factors shape Alexandrium ostenfeldii blooms in the Netherlands
Abstract Harmful algal blooms (HABs) are globally expanding, compromising water quality worldwide. HAB dynamics are determined by a complex interplay of abiotic and biotic factors, and their emergence has often been linked to eutrophication, and more recently to climate change. The dinoflagellate Alexandrium is one of the most widespread HAB genera and its success is based on key functional traits like allelopathy, mixotrophy, cyst formation and nutrient retrieval migrations. Since 2012, dense Alexandrium ostenfeldii blooms (up to 4500 cells mL−1) have recurred annually in a creek located in the southwest of the Netherlands, an area characterized by intense agriculture and aquaculture. We investigated how physical, chemical and biological factors influenced A. ostenfeldii bloom dynamics over three consecutive years (2013–2015). Overall, we found a decrease in the magnitude of the bloom over the years that could largely be linked to changing weather conditions during summer. More specifically, low salinities due to excessive rainfall and increased wind speed corresponded to a delayed A. ostenfeldii bloom with reduced population densities in 2015. Within each year, highest population densities generally corresponded to high temperatures, low DIN:DIP ratios and low grazer densities. Together, our results demonstrate an important role of nutrient availability, absence of grazing, and particularly of the physical environment on the magnitude and duration of A. ostenfeldii blooms. Our results suggest that predicted changes in the physical environment may enhance bloom development in future coastal waters and embayments.https://doi.org/10.1016/j.hal.2017.02.004Warming advances top-down control and reduces producer biomass in a freshwater plankton community
Global warming has been shown to affect ecosystems worldwide. Warming may, for instance, disrupt plant herbivore synchrony and bird phenology in terrestrial systems, reduce primary production in oceans, and promote toxic cyanobacterial blooms in freshwater lakes. Responses of communities will not only depend on direct species-specific temperature effects, but also on indirect effects related to bottom-up and top-down processes. Here, we investigated the impact of warming on freshwater phytoplankton community dynamics, and assessed the relative contribution of nutrient availability, fungal parasitism, and grazing therein. For this purpose, we performed an indoor mesocosm experiment following seasonal temperature dynamics of temperate lakes and a warmed (+4°C) scenario from early spring to late summer. We assessed phytoplankton biomass, C:N:P stoichiometry and community composition, dissolved nutrient availabilities, fungal parasite (i.e., chytrid) prevalence, and zooplankton abundance. Warming led to an overall reduction in phytoplankton biomass as well as lower C:P and N:P ratios, while phytoplankton community composition remained largely unaltered. Warming resulted in an earlier termination of the diatom spring bloom, and an epidemic of its fungal parasite ended earlier as well. Furthermore, warming advanced zooplankton phenology, leading to an earlier top-down control on phytoplankton in the period after the spring bloom. Linear model analysis showed that most of the observed variance in phytoplankton biomass was related to seasonal temperature dynamics in combination with zooplankton abundance. Our findings showed that warming advanced grazer phenology and reduced phytoplankton biomass, thereby demonstrating how bottom-up and top-down related processes may shape future phytoplankton dynamics.https://doi.org/10.1002/ecs2.1651Species sorting and stoichiometric plasticity control community C:P ratio of first-order aquatic consumers
Ecological stoichiometry has proven to be invaluable for understanding consumer response to changes in resource quality. Although interactions between trophic levels occur at the community level, most studies focus on single consumer species. In contrast to individual species, communities may deal with trophic mismatch not only through elemental plasticity but also through changes in species composition. Here, we show that a community of first-order consumers (e.g. zooplankton) is able to adjust its stoichiometry (C:P) in response to experimentally induced changes in resource quality, but only to a limited extent. Furthermore, using the Price equation framework we show the importance of both elemental plasticity and species sorting. These results illustrate the need for a community perspective in ecological stoichiometry, requiring consideration of species-specific elemental composition, intraspecific elemental plasticity and species turnover.https://doi.org/10.1111/ele.12773Integrating chytrid fungal parasites into plankton ecology: research gaps and needs.
Chytridiomycota, often referred to as chytrids, can be virulent parasites with the potential to inflict mass mortalities on hosts, causing e.g. changes in phytoplankton size distributions and succession, and the delay or suppression of bloom events. Molecular environmental surveys have revealed an unexpectedly large diversity of chytrids across a wide range of aquatic ecosystems worldwide. As a result, scientific interest towards fungal parasites of phytoplankton has been gaining momentum in the past few years. Yet, we still know little about the ecology of chytrids, their life cycles, phylogeny, host specificity and range. Information on the contribution of chytrids to trophic interactions, as well as co-evolutionary feedbacks of fungal parasitism on host populations is also limited. This paper synthesizes ideas stressing the multifaceted biological relevance of phytoplankton chytridiomycosis, resulting from discussions among an international team of chytrid researchers. It presents our view on the most pressing research needs for promoting the integration of chytrid fungi into aquatic ecology. This article is protected by copyright. All rights reserved.https://doi.org/10.1111/1462-2920.13827Changes in N:P Supply Ratios affect the Ecological Stoichiometry of a Toxic Cyanobacterium and its Fungal Parasite
Human activities have dramatically altered nutrient fluxes from the landscape into receiving waters. As a result, not only the concentration of nutrients in surface waters has increased, but also their elemental ratios have changed. Such shifts in resource supply ratios will alter autotroph stoichiometry, which may in turn have consequences for higher trophic levels, including parasites. Here, we hypothesize that parasite elemental composition will follow changes in the stoichiometry of its host, and that its reproductive success will decrease with host nutrient limitation. We tested this hypothesis by following the response of a host-parasite system to changes in nitrogen (N) and phosphorus (P) supply in a controlled laboratory experiment. To this end, we exposed a fungal parasite (the chytrid Rhizophydium megarrhizum) to its host (the freshwater cyanobacterium Planktothrix rubescens) under control, low N:P and high N:P conditions. Host N:P followed treatment conditions, with a decreased N:P ratio under low N:P supply, and an increased N:P ratio under high N:P supply, as compared to the control. Shifts in host N:P stoichiometry were reflected in the parasite stoichiometry. Furthermore, at low N:P supply, host intracellular microcystin concentration was lowered as compared to high N:P supply. In contrast to our hypothesis, zoospore production decreased at low N:P and increased at high N:P ratio as compared to the control. These findings suggest that fungal parasites have a relatively high N, but low P requirement. Furthermore, zoospore elemental content, and thereby presumably their size, decreased at high N:P ratios. From these results we hypothesize that fungal parasites may exhibit a trade-off between zoospore size and production. Since zooplankton can graze on chytrid zoospores, changes in parasite production, stoichiometry and cell size may have implications for aquatic food web dynamics.https://doi.org/10.3389/fmicb.2017.01015Mass development of monospecific submerged macrophyte vegetation after the restoration of shallow lakes: roles of light, sediment nutrient levels, and propagule density
Abstract After restoration, eutrophicated shallow freshwaters may show mass development of only one or two submerged macrophyte species, lowering biodiversity and hampering recreation. It is unclear which environmental factors govern this high percentage of the volume inhabited (PVI11 PVI: The percent of the water volume inhabited by submerged macrophytes. ) by submerged macrophytes, and whether the development of a more diverse, low canopy vegetation is likely to occur if dominant species decline in abundance. We hypothesized that (1) adequate light and high sediment nutrient availability leads to massive development of submerged macrophytes, and (2) that macrophyte species richness is low at high PVI, but that this is not caused by a lack of viable propagules of non-dominant species (especially charophytes). To test these hypotheses, fifteen shallow waters in the Netherlands were studied with respect to submerged vegetation (including propagules), water, and sediment characteristics. The probability of high submerged macrophyte PVI is highest in shallow waters where light availability in the water layer and phosphorus availability in the sediment are abundant. These conditions typically occur upon restoration of eutrophic waterbodies by reducing water nutrient loading or applying biomanipulation. Other factors, as top-down control, can additionally influence realised PVI. Viable propagules of species other than the dominant ones, including charophytes, were found in most of the sediments, indicating that once the dominant species declines, there is local potential for a diverse submerged vegetation to develop. Results can be used to predict when mass development occurs and to tackle the factors causing mass development.https://doi.org/10.1016/j.aquabot.2017.04.004Periphyton density is similar on native and non-native plant species
Non-native plants increasingly dominate the vegetation in aquatic ecosystems and thrive in eutrophic conditions. In eutrophic conditions, submerged plants risk being overgrown by epiphytic algae; however, if non-native plants are less susceptible to periphyton than natives, this would contribute to their dominance. Non-native plants may differ from natives in their susceptibility to periphyton growth due to differences in nutrient release, allelopathy and architecture. Yet, there is mixed evidence for whether plants interact with periphyton growth through nutrient release and allelopathy, or whether plants are neutral so that only their architecture matters for periphyton growth.https://doi.org/10.1111/fwb.12911
We hypothesised that (1) non-native submerged vascular plants support lower periphyton density than native species, (2) native and non-native species are not neutral substrate for periphyton and interact with periphyton and (3) periphyton density increases with the plant structural complexity of plant species.
We conducted an experiment in a controlled climate chamber where we grew 11 aquatic plant species and an artificial plant analogue in monocultures in buckets. These buckets were inoculated with periphyton that was collected locally from plants and hard substrate. Of the 11 living species, seven are native to Europe and four are non-native. The periphyton density on these plants was quantified after five weeks.
We found that the periphyton density did not differ between non-native and native plants and was not related to plant complexity. Three living plant species supported lower periphyton densities than the artificial plant, one supported a higher periphyton density and the other plants supported similar densities. However, there was a strong negative correlation between plant growth and periphyton density.
We conclude that the periphyton density varies greatly among plant species, even when these were grown under similar conditions, but there was no indication that the interaction with periphyton differs between native and non-native plant species. Hence, non-native plants do not seem to benefit from reduced periphyton colonisation compared to native species. Instead, certain native and non-native species tolerate eutrophic conditions well and as a consequence, they seem to host less periphyton than less tolerant species.Impact of temperature and nutrients on carbon:nutrient tissue stoichiometry of submerged aquatic plants: an experiment and meta-analysis
Human activity is currently changing our environment rapidly, with predicted temperature increases of 1–5°C over the coming century and increased nitrogen and phosphorus inputs in aquatic ecosystems. In the shallow parts of these ecosystems, submerged aquatic plants enhance water clarity by resource competition with phytoplankton, provide habitat, and serve as a food source for other organisms. The carbon:nutrient stoichiometry of submerged aquatic plants can be affected by changes in both temperature and nutrient availability. We hypothesized that elevated temperature leads to higher carbon:nutrient ratios through enhanced nutrient-use efficiency, while nutrient addition leads to lower carbon:nutrient ratios by the luxurious uptake of nutrients. We addressed these hypotheses with an experimental and a meta-analytical approach. We performed a full-factorial microcosm experiment with the freshwater plant Elodea nuttallii grown at 10, 15, 20, and 25°C on sediment consisting of pond soil/sand mixtures with 100, 50, 25, and 12.5% pond soil. To address the effect of climatic warming and nutrient addition on the carbon:nutrient stoichiometry of submerged freshwater and marine plants we performed a meta-analysis on experimental studies that elevated temperature and/or added nutrients (nitrogen and phosphorus). In the microcosm experiment, C:N ratios of Elodea nuttallii decreased with increasing temperature, and this effect was most pronounced at intermediate nutrient availability. Furthermore, higher nutrient availability led to decreased aboveground C:P ratios. In the meta-analysis, nutrient addition led to a 25, 22, and 16% reduction in aboveground C:N and C:P ratios and belowground C:N ratios, accompanied with increased N content. No consistent effect of elevated temperature on plant stoichiometry could be observed, as very few studies were found on this topic and contrasting results were reported. We conclude that while nutrient addition consistently leads to decreased carbon:nutrient ratios, elevated temperature does not change submerged aquatic plant carbon:nutrient stoichiometry in a consistent manner. This effect is rather dependent on nutrient availability and may be species-specific. As changes in the carbon:nutrient stoichiometry of submerged aquatic plants can impact the transfer of energy to higher trophic levels, these results suggest that eutrophication may enhance plant consumption and decomposition, which could in turn have consequences for carbon sequestration.https://doi.org/10.3389/fpls.2017.00655Cross continental increase in methane ebullition under climate change
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.https://doi.org/10.1038/s41467-017-01535-yThe influence of balanced and imbalanced resource supply on biodiversity–functioning relationship across ecosystems
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.https://doi.org/10.1098/rstb.2015.0283Warming accelerates termination of a phytoplankton spring bloom by fungal parasites
Climate change is expected to favour infectious diseases across ecosystems worldwide. In freshwater and marinehttps://doi.org/10.1111/gcb.13095
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.Lake restoration by in-lake iron addition: A synopsis of iron impact on aquatic organisms and shallow lake ecosystems
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.https://doi.org/10.1007/s10452-015-9552-1Pharmaceuticals may disrupt the natural chemical information flows and species interactions in aquatic systems: ideas and perspectives on a hidden global change
Over the last decades, anthropogenic activities have discharged into the environment many manmade chemicals. There is a rising concern regarding pharmaceutical products and their spread into the environment (e.g. Kümmerer 2008). Due to the enormous quantities consumed, anti-inflammatories, antibiotics, anti-depressives, hormones and blood lipid regulators are found in almost all aquatic environments (Kolpin et al. 2002; Loos et al. 2009). Most pharmaceuticals tend to enter the aquatic environment continuously (but see Sacher et al. 2008 for seasonal exception) in contrast to other pollutants such as herbicides and insecticides which are applied only at specific times related to the life cycle of the target organism, or in response to observed pest outbreaks (Rosi-Marshall and Royer 2012). Pharmaceuticals are designed to be biologically active at very low concentrations and end up in surface waters either unchanged, or as active metabolites/polar conjugates, mostly via municipal wastewater and agricultural discharges (Boxall et al. 2012).https://doi.org/10.1007/398_2015_5002Evaluating early-warning indicators of critical transitions in natural aquatic ecosystems
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.https://doi.org/10.1073/pnas.1608242113What is the influence of a reduction of planktivorous and benthivorous fish on water quality in temperate eutrophic lakes? A systematic review
Background: In recent decades, many attempts have been made to restore eutrophic lakes through biomanipulation.https://doi.org/10.1186/s13750-015-0032-9
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.
Keywords: Biomanipulation, Planktivore, Benthivore, Piscivore stocking, Fish removal, Lake restoration, Eutrophication,
Water quality, PhytoplanktonFighting internal phosphorus loading: An evaluation of the large scale application of gradual Fe-addition to a shallow peat lake
Abatement of external phosphorus (P) loading and biomanipulation are measures that are often applied with the aim to restore a macrophyte dominated clearwater state in turbid, anthropogenically eutrophied lakes. The recovery of such lakes, however, is often hampered by ‘internal eutrophication’, as a result of the release of historically accumulated P from the sediment into the water column. One way to combat this internal P loading is by adding iron (Fe) into the lake, which naturally binds to phosphate. Although studied in the laboratory or mesocosms, the effects of iron addition on a whole-lake scale are largelyhttps://doi.org/10.1016/j.ecoleng.2015.05.034
unknown. In this study we therefore compiled lake monitoring data to evaluate the effect of a gradual dose of 33 g Fe m2 on the water quality and biotic communities (phytoplankton, zooplankton and macrophytes) of Lake Terra Nova. During and after the Fe-addition, we also carried out assays to evaluate the effect of the Fe-addition on sediment P-release rates. Lake Terra Nova is a eutrophied, shallow peaty lake that has been subjected to biomanipulation measures for 10 years.
The large scale addition of iron during 2010 and 2011 resulted in a substantial reduction of dissolved P, suspended matter (SM), phytoplankton biomass and relative Cyanobacterial biomass, whereas macrophytes reappeared. Fe-addition also resulted in strongly reduced sediment P-mobilisation rates.
Nevertheless, reductions of water column TP in response to the Fe-addition were short-lived. This discrepancy between the observed TP-levels in the water column and sediment P-release rates could indicate that sustained loading with external P and interactions of chemical compounds with Fe may jeopardize long term restoration success by depleting the water column reservoir of reactive Fe. Ecological processes can in addition enhance this process by shunting P from the sediment to the water column.Biological control of toxic cyanobacteria by mixotrophic predators: an experimental test of intraguild predation theory
Intraguild predators both feed on and compete with their intraguild prey. In theory, intraguild predators can therefore be very effective as biological control agents of intraguild prey species, especially in productive environments. We investigated this hypothesis using the mixotrophic chrysophyte Ochromonas as intraguild predator and the harmful cyanobacterium Microcystis aeruginosa as its prey. Ochromonas can grow photoautotrophically, but can also graze efficiently on Microcystis. Hence, it competes with its prey for inorganic resources. We developed a mathematical model and parameterized it for our experimental food web. The model predicts dominance of Microcystis at low nutrient loads, coexistence of both species at intermediate nutrient loads, and dominance of Ochromonas but a strong decrease of Microcystis at high nutrient loads. We tested these theoretical predictions in chemostat experiments supplied with three different nitrogen concentrations. Ochromonas initially suppressed the Microcystis abundance by >97% compared to the Microcystis monocultures. Thereafter, however, Microcystis gradually recovered to ∼20% of its monoculture abundance at low nitrogen loads, but to 50–60% at high nitrogen loads. Hence, Ochromonas largely lost control over the Microcystis population at high nitrogen loads. We explored several mechanisms that might explain this deviation from theoretical predictions, and found that intraspecific interference at high Ochromonas densities reduced their grazing rates on Microcystis. These results illustrate the potential of intraguild predation to control pest species, but also show that the effectiveness of their biological control can be reduced in productive environments. Read More: http://www.esajournals.org/doi/abs/10.1890/13-0218.1https://doi.org/10.1890/13-0218.1Comparison of predator-prey interactions with and without intraguild predation by manipulation of the nitrogen source
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.https://doi.org/10.1111/j.1600-0706.2013.00736.xIron addition as a measure to restore water quality: Implications for macrophyte growth
Eutrophication of shallow lakes in North-West Europe has resulted in cyanobacterial blooms, turbid water, and a decline in submerged macrophytes. Even though external inputs of phosphorus (P) are declining, internal loading of P from the sediment may delay the recovery of these aquatic ecosystems. Iron can be a useful chemical binding agent to combat internal P loading in shallow lakes, but may potentially be harmful for macrophyte growth. We tested whether iron addition as a restoration measure harms the growth of submerged macrophytes. We hypothesized that this depends on the iron dosage and the rooting strategy of the macrophytes. We experimentally tested the effects of Fe (FeCl3) on the submerged macrophytes Potamogeton pectinatus L. and Elodea nuttallii (Planch.) H. St. John. Iron was dosed at a concentration of 20 g Fe m−2 and 40 g Fe m−2 to the surface water or to both the surface water and sediment. E. nuttallii growth was not affected by iron addition, whereas P. pectinatus growth significantly decreased with increasing iron concentrations. Nonetheless, biomass of both species increased in all treatments relative to starting conditions. During the experiment, propagules sprouted from a propagule bank in the sediment including species with a high conservation value and this spontaneous emergence was not influenced by increasing iron concentrations. We conclude that adding iron(III)chloride in dosages of 20–40 g m−2 may reduce growth of some macrophyte species, but does not prevent overall macrophyte recovery. It may however affect macrophyte community composition due to differential responses of macrophyte species to iron addition.https://doi.org/10.1016/j.aquabot.2014.01.007Community stoichiometry in a changing world: combined effects of warming and eutrophication on phytoplankton dynamics
The current changes in our climate will likely have far reaching consequences for aquatic ecosystems. These changes in the climate, however, do not act alone and are often accompanied by additional stressors such as eutrophication. Both global warming and eutrophication have been shown to affect the timing and magnitude of phytoplankton blooms. Little is known about the combined effects of rising temperatures and eutrophication on the stoichiometry of entire phytoplankton communities. We exposed a natural phytoplankton spring community to different warming and phosphorus loading scenarios using a full-factorial design. Our results demonstrate that rising temperatures promote the growth rate of an entire phytoplankton community. Furthermore, both rising temperatures and phosphorus loading stimulated the maximum biomass built up by the phytoplankton community. Rising temperatures led to higher carbon:nutrient stoichiometry of the phytoplankton community under phosphorus limited conditions. Such a shift towards higher carbon:nutrient ratios, in combination with a higher biomass build-up, suggests a temperature-driven increase in nutrient use efficiency of the phytoplankton community. Importantly, higher carbon:nutrient stoichiometry of phytoplankton is generally of poorer nutritional value for zooplankton. Thus, although warming may result in higher phytoplankton biomass, this may be accompanied by a stoichiometric mismatch between phytoplankton and their grazers, with possible consequences for the entire aquatic food web. Read More: http://www.esajournals.org/doi/abs/10.1890/13-1251.1https://doi.org/10.1890/13-1251.1Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf, southern Lake Victoria (Tanzania)
Summary 1.We investigated how dynamics in phytoplankton biomass are driven through light and nutrient limitations in the Mwanza Gulf, Lake Victoria (Tanzania). 2.We measured light attenuation, chlorophyll a and water quality parameters along a phytoplankton biomass gradient in the Mwanza Gulf at six sampling stations in three different seasons from August 2009 to March 2011. 3.We found a higher rate of attenuation of photosynthetic available radiation than in other gulfs of Lake Victoria which could only partially be explained by phytoplankton biomass, measured as chlorophyll a. Suspended particulate detritus strongly affected light attenuation, particularly in shallow water. 4.In shallow water in the southern part of the Mwanza Gulf, nutrients were in excess and phytoplankton biomass was limited by light. In deeper water near the entrance of the gulf, light was not limiting when nitrogen probably was, as indicated by the N:P ratio. 5.Low abundance of N-fixing phytoplankton species suggests that N-fixation was low in shallow water. Phytoplankton biomass was lower in Mwanza Gulf than in northern gulfs of Lake Victoria, but might increase in the future as a result of nutrient enrichment caused by an increase in intensive agriculture and rapid growth of the city of Mwanza.https://doi.org/10.1111/fwb.12253Rising CO2 levels will intensify phytoplankton blooms in eutrophic and hypertrophic lakes
Harmful algal blooms threaten the water quality of many eutrophic and hypertrophic lakes and cause severe ecological and economic damage worldwide. Dense blooms often deplete the dissolved CO2 concentration and raise pH. Yet, quantitative prediction of the feedbacks between phytoplankton growth, CO2 drawdown and the inorganic carbon chemistry of aquatic ecosystems has received surprisingly little attention. Here, we develop a mathematical model to predict dynamic changes in dissolved inorganic carbon (DIC), pH and alkalinity during phytoplankton bloom development. We tested the model in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa at different CO2 levels. The experiments showed that dense blooms sequestered large amounts of atmospheric CO2, not only by their own biomass production but also by inducing a high pH and alkalinity that enhanced the capacity for DIC storage in the system. We used the model to explore how phytoplankton blooms of eutrophic waters will respond to rising CO2 levels. The model predicts that (1) dense phytoplankton blooms in low- and moderately alkaline waters can deplete the dissolved CO2 concentration to limiting levels and raise the pH over a relatively wide range of atmospheric CO2 conditions, (2) rising atmospheric CO2 levels will enhance phytoplankton blooms in low- and moderately alkaline waters with high nutrient loads, and (3) above some threshold, rising atmospheric CO2 will alleviate phytoplankton blooms from carbon limitation, resulting in less intense CO2 depletion and a lesser increase in pH. Sensitivity analysis indicated that the model predictions were qualitatively robust. Quantitatively, the predictions were sensitive to variation in lake depth, DIC input and CO2 gas transfer across the air-water interface, but relatively robust to variation in the carbon uptake mechanisms of phytoplankton. In total, these findings warn that rising CO2 levels may result in a marked intensification of phytoplankton blooms in eutrophic and hypertrophic waters.https://doi.org/10.1371/journal.pone.0104325Coupled human and natural system dynamics as key to the sustainability of Lake Victoria’s ecosystem services
East Africa’s Lake Victoria provides resources and services to millions of people on the lake’s shores and abroad. In particular, the lake’s fisheries are an important source of protein, employment, and international economic connections for the whole region. Nonetheless, stock dynamics are poorly understood and currently unpredictable. Furthermore, fishery dynamics are intricately connected to other supporting services of the lake as well as to lakeshore societies and economies. Much research has been carried out piecemeal on different aspects of Lake Victoria’s system; e.g., societies, biodiversity, fisheries, and eutrophication. However, to disentangle drivers and dynamics of change in this complex system, we need to put these pieces together and analyze the system as a whole. We did so by first building a qualitative model of the lake’s social-ecological system. We then investigated the model system through a qualitative loop analysis, and finally examined effects of changes on the system state and structure. The model and its contextual analysis allowed us to investigate system-wide chain reactions resulting from disturbances. Importantly, we built a tool that can be used to analyze the cascading effects of management options and establish the requirements for their success. We found that high connectedness of the system at the exploitation level, through fisheries having multiple target stocks, can increase the stocks’ vulnerability to exploitation but reduce society’s vulnerability to variability in individual stocks. We describe how there are multiple pathways to any change in the system, which makes it difficult to identify the root cause of changes but also broadens the management toolkit. Also, we illustrate how nutrient enrichment is not a self-regulating process, and that explicit management is necessary to halt or reverse eutrophication. This model is simple and usable to assess system-wide effects of management policies, and can serve as a paving stone for future quantitative analyses of system dynamics at local scales.https://doi.org/10.5751/ES-06965-190431Food quality dominates the impact of food quantity on Daphnia life history: possible implications for re-oligotrophication
The elemental composition of phytoplankton is highly variable compared to the relatively narrow stoichiometry of zooplankton grazers. Using a full factorial design, we tested the effects of alterations in algal elemental composition (i.e., food quality) combined with food quantity on the life history of a Daphnia galeata clone from Lake IJsselmeer. Lower food quality reduced survival, growth, and reproduction. Food quantity became important at high food quality only. The strong effect of food quality indicates the potential for a stoichiometric bottleneck in Lake IJsselmeer, resulting in less high quality food for higher trophic levels as a result of re-oligotrophication.https://doi.org/10.5268/IW-4.4.701Epiphytic diatoms along environmental gradients in Western European shallow lakes
European shallow lakes; epiphyton; Bacillariophyta; bioindication; eutrophication; total phosphorus Diatom-based assays have been successfully associated worldwide with the trophic 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.https://doi.org/10.1002/clen.201200630Mixotrophic organisms become more heterotrophic with rising temperature
The metabolic theory of ecology predicts that temperature affects heterotrophic processes more strongly than autotrophic processes. We hypothesized that this differential temperature response may shift mixotrophic organisms towards more heterotrophic nutrition with rising temperature. The hypothesis was tested in experiments with the mixotrophic chrysophyte Ochromonas sp., grown under autotrophic, mixotrophic and heterotrophic conditions. Our results show that (1) grazing rates on bacterial prey increased more strongly with temperature than photosynthetic electron transport rates, (2) heterotrophic growth rates increased exponentially with temperature over the entire range from 13 to 33 °C, while autotrophic growth rates reached a maximum at intermediate temperatures and (3) chlorophyll contents during mixotrophic growth decreased at high temperature. Hence, the contribution of photosynthesis to mixotrophic growth strongly decreased with temperature. These findings support the hypothesis that mixotrophs become more heterotrophic with rising temperature, which alters their functional role in food webs and the carbon cycle.https://doi.org/10.1111/ele.12033Soil and Freshwater and Marine Sediment Food Webs: Their Structure and Function
The food webs of terrestrial soils and of freshwater and marine sediments depend on adjacent aboveground or pelagic ecosystems for organic matter input that provides nutrients and energy. There are important similarities in the flow of organic matter through these food webs and how this flow feeds back to primary production. In both soils and sediments, trophic interactions occur in a cycle in which consumers stimulate nutrient cycling such that mineralized resources are made available to the primary producers. However, aquatic sediments and terrestrial soils differ greatly in the connectivity between the production and the consumption of organic matter. Terrestrial soils and shallow aquatic sediments can receive organic matter within hours of photosynthesis when roots leak carbon, whereas deep oceanic sediments receive organic matter possibly months after carbon assimilation by phytoplankton. This comparison has implications for the capacity of soils and sediments to affect the global carbon balance.https://doi.org/10.1525/bio.2013.63.1.8Plankton dynamics under different climatic conditions in space and time
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.https://doi.org/10.1111/fwb.12053Temperature alters host genotype-specific susceptibility to chytrid infection
The cost of parasitism often depends on environmental conditions and host identity. Therefore, variation in the biotic and abiotic environment can have repercussions on both, species-level host-parasite interaction patterns but also on host genotype-specific susceptibility to disease. We exposed seven genetically different but concurrent strains of the diatom Asterionella formosa to one genotype of its naturally co-occurring chytrid parasite Zygorhizidium planktonicum across five environmentally relevant temperatures. We found that the thermal tolerance range of the tested parasite genotype was narrower than that of its host, providing the host with a “cold” and “hot” thermal refuge of very low or no infection. Susceptibility to disease was host genotype-specific and varied with temperature level so that no genotype was most or least resistant across all temperatures. This suggests a role of thermal variation in the maintenance of diversity in disease related traits in this phytoplankton host. The duration and intensity of chytrid parasite pressure on host populations is likely to be affected by the projected changes in temperature patterns due to climate warming both through altering temperature dependent disease susceptibility of the host and, potentially, through en- or disabling thermal host refugia. This, in turn may affect the selective strength of the parasite on the genetic architecture of the host population.https://doi.org/10.1371/journal.pone.0071737Alternative states and population crashes in a resource-susceptible-infected model for planktonic parasites and hosts
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.https://doi.org/10.1111/fwb.12010Elevated CO2 concentrations affect the elemental stoichiometry and species composition of an experimental phytoplankton community
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.https://doi.org/10.1111/j.1365-2427.2012.02833.xIron addition as a shallow lake restoration measure: impacts on charophyte growth
Eutrophication has caused a decline of charophyte species in many shallow lakes in Europe. Even though external inputs of phosphorus are declining, internal loading of P from the sediment seems to delay the recovery of these systems. Iron is a useful chemical binding agent to combat internal phosphorus loading. However, the effects of iron addition on charophytes are not yet known. In this study we experimentally tested the potential toxicity of iron(III)chloride (FeCl3) on two different charophytes, Chara virgata Kützing and Chara globularis Thuiller added at the concentration of 20 g Fe m−2 and 40 g Fe m−2 to the surface water. C. virgata growth was not significantly affected, whereas C. globularis growth significantly decreased with increasing iron concentrations. Nonetheless, biomass of both species increased in all treatments relative to starting conditions. The decrease of C. globularis biomass with high iron additions may have been caused by a drop in pH and alkalinity in combination with iron induced light limitation. Iron addition over a longer time scale, however, will not cause this rapid drop in pH. Therefore, we conclude that adding iron(III)chloride in these amounts to the surface water of a lake can potentially be a useful restoration method.https://doi.org/10.1007/s10750-011-0995-7Plankton dynamics under different climate conditions in tropical freshwater systems (a reply to the comment by Sarmento et al. 2013)
1.In our recent contribution to the special issue on plankton dynamics in a fast-changing world, we outlined some general predictions of plankton dynamics in different climate regions now and in future, building on the Plankton Ecology Group (PEG) model (de Senerpont Domis et al., 2013). 2.We proposed a stylised version of plankton dynamics in Fig. 3 of our article and stated that these patterns need to be further elaborated. Our figure displays annual plankton dynamics now and in future in oligotrophic, mesotrophic and eutrophic lakes in arctic, temperate and tropical climate zones. 3.We fully agree with Sarmento, Amado & Descy (2013) that more data on tropical regions are needed, and we are looking forward to the emergence of published data from tropical regions to extend our still-limited understanding of plankton dynamics in these regions. 4.Sarmento et al. (2013) did not agree with our predictions on plankton dynamics for hydrology-driven water systems in the tropics. Unfortunately, however, Sarmento et al. (2013) did not substantiate their statements with the much-needed data on plankton dynamics in the tropics. Moreover, they merely provide an overview of precipitation patterns in the tropics, not an alternative hypothesis for our predictions.https://doi.org/10.1111/fwb.12203Restoring macrophyte diversity in shallow temperate lakes: biotic versus abiotic constraints
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 humaninduced 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.https://doi.org/10.1007/s10750-012-1142-9Warmer climates boost cyanobacterial dominance in shallow lakes
Dominance by cyanobacteria hampers human use of lakes and reservoirs worldwide. Previous studies indicate that excessive nutrient loading and warmer conditions promote dominance by cyanobacteria, but evidence from global scale field data has so far been scarce. Our analysis, based on a study of 143 lakes along a latitudinal transect ranging from subarctic Europe to southern South America, shows that although warmer climates do not result in higher overall phytoplankton biomass, the percentage of the total phytoplankton biovolume attributable to cyanobacteria increases steeply with temperature. Our results also reveal that the percent cyanobacteria is greater in lakes with high rates of light absorption. This points to a positive feedback because restriction of light availability is often a consequence of high phytoplankton biovolume, which in turn may be driven by nutrient loading. Our results indicate a synergistic effect of nutrients and climate. The implications are that in a future warmer climate, nutrient concentrations may have to be reduced substantially from present values in many lakes if cyanobacterial dominance is to be controlled.https://doi.org/10.1111/j.1365-2486.2011.02488.xGenotype-by-temperature interactions may help to maintain clonal diversity in Asterionella formosa (Bacillariophyceae)
Marine and freshwater phytoplankton populations often show large clonal diversity, which is in disagreement with clonal selection of the most vigorous genotype(s). Temporal fluctuation in selection pressures in variable environments is a leading explanation for maintenance of such genetic diversity. To test the influence of temperature as a selection force in continually (seasonally) changing aquatic systems we carried out reaction norms experiments on co-occurring clonal genotypes of a ubiquitous diatom species, Asterionella formosa Hassall, across an environmentally relevant range of temperatures. We report within population genetic diversity and extensive diversity in genotype-specific reaction norms in growth rates and cell size traits. Our results showed genotype by environment interactions, indicating that no genotype could outgrow all others across all temperature environments. Subsequently, we constructed a model to simulate the relative proportion of each genotype in a hypothetical population based on genotype and temperature-specific population growth rates. This model was run with different seasonal temperature patterns. Our modeling exercise showed a succession of two to several genotypes becoming numerically dominant depending on the underlying temperature pattern. The results suggest that (temperature) context dependent fitness may contribute to the maintenance of genetic diversity in isolated populations of clonally reproducing microorganisms in temporally variable environments.https://doi.org/10.1111/j.1529-8817.2012.01205.xBeyond the Plankton Ecology Group (PEG) Model: Mechanisms Driving Plankton Succession
The seasonal succession of plankton is an annually repeated process of community assembly during which all major external factors and internal interactions shaping communities can be studied. A quarter of a century ago, the state of this understanding was described by the verbal plankton ecology group (PEG) model. It emphasized the role of physical factors, grazing and nutrient limitation for phytoplankton, and the role of food limitation and fish predation for zooplankton. Although originally targeted at lake ecosystems, it became also adopted by marine plankton ecologists. Since then, a suite of ecological interactions previously underestimated in importance have become research foci: overwintering of key organisms, the microbial food web, parasitism, and food quality as a limiting factor and an extended role of higher order predators. A review of the impact of these novel interactions on plankton seasonal succession reveals limited effects on gross seasonal biomass patterns, but strong effects on species replacements.https://doi.org/10.1146/annurev-ecolsys-110411-160251Induced defences in marine and freshwater phytoplankton: a review.
Many organisms have developed defences to avoid predation by species at higher trophic levels. The capability of primary producers to defend themselves against herbivores affects their own survival, can modulate the strength of trophic cascades and changes rates of competitive exclusion in aquatic communities. Algal species are highly flexible in their morphology, growth form, biochemical composition and production of toxic and deterrent compounds. Several of these variable traits in phytoplankton have been interpreted as defence mechanisms against grazing. Zooplankton feed with differing success on various phytoplankton species, depending primarily on size, shape, cell wall structure and the production of toxins and deterrents. Chemical cues associated with (i) mechanical damage, (ii) herbivore presence and (iii) grazing are the main factors triggering induced defences in both marine and freshwater phytoplankton, but most studies have failed to disentangle the exact mechanism( s) governing defence induction in any particular species. Induced defences in phytoplankton include changes in morphology (e.g. the formation of spines, colonies and thicker cell walls), biochemistry (such as production of toxins, repellents) and in life history characteristics (formation of cysts, reduced recruitment rate). Our categorization of inducible defences in terms of the responsible induction mechanism provides guidance for future work, as hardly any of the available studies on marine or freshwater plankton have performed all the treatments that are required to pinpoint the actual cue(s) for induction. We discuss the ecology of inducible defences in marine and freshwater phytoplankton with a special focus on the mechanisms of induction, the types of defences, their costs and benefits, and their consequences at the community level.https://doi.org/10.1007/s10750-010-0395-4Parasitic chytrids could promote copepod survival by mediating material transfer from inedible diatoms
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.https://doi.org/10.1007/s10750-010-0274-zReversal in competitive dominance of a toxic versus non-toxic cyanobacterium in response to rising CO2
Climate change scenarios predict a doubling of the atmospheric CO2 concentration by the end of this century. Yet, how rising CO2 will affect the species composition of aquatic microbial communities is still largely an open question. In this study, we develop a resource competition model to investigate competition for dissolved inorganic carbon in dense algal blooms. The model predicts how dynamic changes in carbon chemistry, pH and light conditions during bloom development feed back on competing phytoplankton species. We test the model predictions in chemostat experiments with monocultures and mixtures of a toxic and non-toxic strain of the freshwater cyanobacterium Microcystis aeruginosa. The toxic strain was able to reduce dissolved CO2 to lower concentrations than the non-toxic strain, and became dominant in competition at low CO2 levels. Conversely, the non-toxic strain could grow at lower light levels, and became dominant in competition at high CO2 levels but low light availability. The model captured the observed reversal in competitive dominance, and was quantitatively in good agreement with the results of the competition experiments. To assess whether microcystins might have a role in this reversal of competitive dominance, we performed further competition experiments with the wild-type strain M. aeruginosa PCC 7806 and its mcyB mutant impaired in microcystin production. The microcystin-producing wild type had a strong selective advantage at low CO2 levels but not at high CO2 levels. Our results thus demonstrate both in theory and experiment that rising CO2 levels can alter the community composition and toxicity of harmful algal blooms.https://doi.org/10.1038/ismej.2011.28Effects of nutrient additions and macrophyte composition on invertebrate community assembly and diversity in experimental ponds.
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,weorthogonally 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 3mg 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.https://doi.org/10.1016/j.baae.2011.05.001Chytrid infections and diatom spring blooms: paradoxical effects of climate warming on fungal epidemics in lakes?
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 °C, offering a disease-free window of opportunity for growth of Asterionella. Climate warming has reduced periods in which water temperature remainshttps://doi.org/10.1111/j.1365-2427.2010.02565.xClimate-driven changes in the ecological stoichiometry of aquatic ecosystems
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.https://doi.org/10.1890/080178Maximizing growth rate at low temperatures: RNA:DNA allocation strategies and life history traits of Arctic and temperate Daphnia
Many short-lived or univoltine organisms at high latitudes and altitudes face the challenge to complete their life-cycle within a brief growing season. This means that they need to maintain a high growth rate at low temperatures, and one way of doing this is to allocate limiting resources like phosphorus to RNA in order to maximize protein synthesis. We here explore the allocations of phosphorus to RNA relative to DNA, and the potential bearings on growth rate and life history traits of polyploid (high-Arctic) and diploid (temperate) Daphnia pulex. The polyploid clone matured earlier at low temperature (8°C) but later than the diploid clone at high temperature (18°C). Juveniles of Arctic Daphnia had both higher specific levels of RNA and higher growth rates at low temperature compared with the temperate clone of Daphnia. We hypothesize that Arctic Daphnia may overcome growth constraints posed by low temperature and polyploidy by increasing their allocation of resources to RNA. The prevalence of polyploidy in Arctic populations strongly suggests that the potential drawbacks of polyploidy are counteracted by an increased allocation of resources to RNA to keep a high rate of protein synthesis even under low temperatures.https://doi.org/10.1007/s00300-010-0814-zCombined and single effects of pesticide carbaryl and toxic Microcystis aeruginosa on the life history of Daphnia pulicaria
The combined influence of a pesticide (carbaryl) and a cyanotoxin (microcystin LR) on the life history of Daphnia pulicaria was investigated. At the beginning of the experiments animals were pulse exposed to carbaryl for 24 h and microcystins were delivered bound in Microcystis’ cells at different, sub-lethal concentrations (chronic exposure). In order to determine the actual carbaryl concentrations in the water LC–MS/MS was used. For analyses of the cyanotoxin concentration in Daphnia’s body enzyme-linked immunosorbent assay (ELISA) was used. Individual daphnids were cultured in a flow-through system under constant light (16 h of light: 8 h of dark), temperature (20°C), and food conditions (Scenedesmus obliquus, 1 mg of C l−1). The results showed that in the treatments with carbaryl egg numbers per female did not differ significantly from controls, but the mortality of newborns increased significantly. Increasing microcystin concentrations significantly delayed maturation, reduced size at first reproduction, number of eggs, and newborns. The interaction between carbaryl and Microcystis was highly significant. Animals matured later and at a smaller size than in controls. The number of eggs per female was reduced as well. Moreover, combined stressors caused frequent premature delivery of offspring with body deformations such as dented carapax or an undeveloped heart. This effect is concluded to be synergistic and could not be predicted from the effects of the single stressors.https://doi.org/10.1007/s10750-010-0130-1Microcystins do not provide anti-herbivore defence against mixotrophic flagellates
While most experiments investigating zooplankton grazing on harmful cyanobacteria have been carried out with metazoan plankton, several protozoa can also feed efficiently on cyanobacteria. We investigated grazing by the mixotrophic flagellate Ochromonas sp. on the toxic cyanobacterium Microcystis aeruginosa. Ochromonas sp. grew rapidly on M. aeruginosa and had a strong impact on the population density of its prey. However, specific growth rates of Ochromonas sp. decreased over time, possibly indicating a negative impact on Ochromonas sp. mediated by M. aeruginosa. Grazing did not have any effect on the intracellular microcystin content of M. aeruginosa, and the ingested microcystins did not accumulate within Ochromonas sp. We studied the functional and numerical response of Ochromonas sp. grazing on the microcystin-producing strain M. aeruginosa PCC 7806 and its microcystin-deficient mutant. Ochromonas sp. showed a Type 3 functional response of very similar shape on both the toxic and non-toxic M. aeruginosa strain. Ingestion rates of Ochromonas sp. were even slightly higher on the toxic M. aeruginosa strain. We therefore found no indication of microcystins acting as a defence against mixotrophic flagellates.https://doi.org/10.3354/ame01395Pulsed nitrogen supply induces dynamic changes in the amino acid compositionand microcystin production of the harmful cyanobacterium Planktothrix agardhii
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.https://doi.org/10.1111/j.1574-6941.2010.00958.xEffect of macrophyte community composition and nutrient enrichment on plant biomass and algal blooms.
Submerged freshwater macrophytes decline with increasing eutrophication. This has consequences for ecosystem processes in shallow lakes and ponds as macrophytes can reduce algal blooms under eutrophic conditions. We hypothesize that the productivity of submerged vegetation, biomass change under eutrophication and the suppression of algal blooms may be affected by macrophyte community composition. To test our hypothesis, we established three macrophyte community types in 36 fishless experimental ponds: one dominated by the oligotrophic species Chara globularis, one dominated by the eutrophic species Potamogeton pectinatus and a diverse vegetation which became co-dominated by Elodea nuttallii and C. globularis, and we fertilized half of the ponds. The macrophyte communities produced different amounts of biomass and they responded differently to fertilization. The community dominated by Potamogeton produced the lowest overall biomass, but was not affected by nutrient addition. The communities dominated by Chara and co-dominated by Elodea and Chara produced more than four-fold the amount of biomass produced in Potamogeton communities under oligotrophic conditions, but were strongly negatively affected by nutrient addition. Phytoplankton abundance did not differ significantly among the plant community types, but showed large variation within community types. There was a significant negative relationship between spring macrophyte biomass and the probability of summer algal blooms. The occurrence of algal blooms coincided with low daphnid densities and high pH (>10). We conclude that the macrophyte community composition, characterized by the dominant species, strongly affected the amount of biomass production as well as the short-term response of the vegetation to nutrient enrichment. Macrophyte community composition had no direct effect on algal blooms, but can affect the occurrence of algal blooms indirectly as these occurred only in ponds with low (https://doi.org/10.1016/j.baae.2010.06.005Local deepening of large shallow peat lakes: a measure to improve their ecological status
Fine organic suspended sediments (refractory detritus) play an important role in the underwater light attenuation of large shallow lakes with a peat origin and a eutrophication history. Wind driven resuspension of this material, its flocculation in the water column and the settlement of the formed flocs are the main processes governing the dynamics of this material. For restoration of the macrophyte community in such lakes, reduction of this refractory detritus to improve underwater light climate is the key process when eutrophication reduction measures alone are not effective enough. The shallow Lake Loosdrecht (The Netherlands) was used as case study to illustrate the effects of artificially created deepenings on suspended matter concentrations and the consequences for the underwater light climate. Suspended sediment balances were created for the current situation and the situation with deepenings. Field measurements were taken to quantify various processes and results of the calculations from the suspended sediment balances were used to quantify the effects on light climate and potential habitat for macrophytes. These calculations show that creating deepenings (three sections with a total surface area of 120 ha and a depth of 12 m on a 10% section of the lake) decreases the concentration of organic detritus by 25% and decreases attenuation coefficients from 2.5 m-1 to 2.2 m-1. P-load reductions affecting chlorophyll-a levels lead to a change of attenuation coefficients from 2.5 m-1 to 2.0 m-1. The combination of deepenings with P-load reduction measures gives the most optimal result and leads to a predicted attenuation coefficient of 1.6 m-1. These improvements of the underwater light climate are a first step to the recovery of the submerged macrophyte community.https://doi.org/10.4081/jlimnol.2010.126The effect of mixotrophic chrysophyte on toxic and colony-forming cyanobacteria
1. In order to test the effect of Ochromonas sp., a mixotrophic chrysophyte, on cyanobacteria, grazing experiments were performed under controlled conditions. We studied grazing on three Microcystis aeruginosa strains, varying in toxicity and morphology, as well as on one filamentous cyanobacterium, Pseudanabaena sp. Furthermore, we analysed the co-occurrence of Ochromonas and Microcystis in natural systems in relation to various environmental parameters (TP, TN, DOC, temperature, pH), using data from 460 Norwegian lakes. 2. Ochromonas was able to feed on all four cyanobacterial strains tested, and grew quickly on all of them. The chrysophyte caused net growth reductions in all three Microcystis strains (the very toxic single-celled strain PCC 7806; the less toxic colony-forming Bear AC and the less toxic single-celled Spring CJ). The effect of Ochromonas was strongest on the Spring CJ strain. Although the effect of Ochromonas grazing on the growth of Pseudanabaena was relatively smaller, it also reduced the net 3. After 4 days of incubation with Ochromonas the total amount of cyanotoxins in the three Microcystis strains was reduced by 91.1–98.7% compared with the controls. 4. Ochromonas occurred in similar densities across all 460 Norwegian lakes. Microcystis occurred only at higher TN, TP, temperature and pH values, although its density was often several orders of magnitude higher than that of Ochromonas. Ochromonas co-occurred in 94% of the samples in which Microcystis was present. 5. From our study it is not clear whether Ochromonas could control Microcystis blooms in natural lakes. However, our study does demonstrate that Ochromonas usually occurs in lakes with Microcystis, and our small scale experiments show that Ochromonas can strongly reduce the biomass of Microcystis and its toxin content.https://doi.org/10.1111/j.1365-2427.2009.02227.xAlgal defenses, population stability and the risk of herbivore extinctions: a chemostat model and experiment
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.https://doi.org/10.1007/s11284-009-0596-3Climate change and the future of freshwater biodiversity in Europe: a primer for policy-makers
Earth’s climate is changing, and by the end of the 21st century in Europe, average temperatures are likely to have risen by at least 2 °C, and more likely 4 °C, with associated effects on patterns of precipitation and the frequency of extreme weather events. Attention among policy-makers is divided about how to minimise the change, how to mitigate its effects, how to maintain the natural resources on which societies depend and how to adapt human societies to the changes. Natural systems are still seen, through a long tradition of conservation management that is largely species-based, as amenable to adaptive management, and biodiversity, mostly perceived as the richness of plant and vertebrate communities, often forms a focus for planning. We argue that prediction of particular species changes will be possible only in a minority of cases but that prediction of trends in general structure and operation of four generic freshwater ecosystems (erosive rivers, depositional floodplain rivers, shallow lakes and deep lakes) in three broad zones of Europe (Mediterranean, Central and Arctic-Boreal) is practicable. Maintenance and rehabilitation of ecological structures and operations will inevitably and incidentally embrace restoration of appropriate levels of species biodiversity. Using expert judgement, based on an extensive literature, we have outlined, primarily for lay policy makers, the pristine features of these systems, their states under current human impacts, how these states are likely to alter with a warming of 2 °C to 4 °C and what might be done to mitigate this. We have avoided technical terms in the interests of communication, and although we have included full referencing as in academic papers, we have eliminated degrees of detail that could confuse broad policy-making.https://doi.org/10.1608/FRJ-2.2.1The ecological stoichiometry of toxins produced by harmful cyanobacteria: an experimental test of the carbon-nutrient balance hypothesis
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 CO2. 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 CO2 and nitrogen enrichment will affect the microcystin composition of harmful cyanobacteria.https://doi.org/10.1111/j.1461-0248.2009.01383.xLakes as sentinels of climate change
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.https://doi.org/10.4319/lo.2009.54.6_part_2.2283Assessing ecological quality of shallow lakes: Does knowledge of transparency suffice?
The European Water Framework Directive (WFD) requires that all aquatic ecosystems in their member states should reach ‘good’ ecological quality by 2015. To assess ecological quality, the WFD requires the definition of reference conditions using biological, physical and chemical indicators and the assignment of each water body to one of five quality classes using these indicators. Elaborate assessment schemes using large sets of variables are now being developed. Here we address the question whether all this is really needed and what the simplest assessment approach would be for the case of shallow lakes. We explore the relationships between the quality class assigned to a lake by experts in shallow lake ecology and a rich set of biological, physical, and chemical data. Multinomial logistic regression analyses were carried out based on data from 86 shallow lakes throughout Europe that were sampled in 2000 and/or 2001. Ecological quality of shallow lakes judged by experts was strongly correlated to physical and chemical variables associated with light regime and nutrients and much less to biological variables. Our regression model showed that ecological quality of this set of shallow lakes judged by experts could be predicted quite well from water transparency expressed as Secchi depth and that other variables did not contribute to it significantly. According to the WFD, lakes should at least have a ‘good’ ecological quality. Quality judged by experts and predicted quality were similar for 78% of the lakes with respect to meeting this standard. As a cautionary note we stress that Secchi depth alone wihttps://doi.org/10.1016/j.baae.2007.12.009Interaction between the macrophyte Stratiotes aloides and filamentous algae: does it indicate allelopathy?
The aquatic macrophyte Stratiotes aloides Linnaeus, which has recently received attention in studies on allelopathy, has been shown to suppress phytoplankton growth. In the Netherlands, S. aloides often co-occurs with floating filamentous algae. However, filamentous algae are generally absent in close proximity to S. aloides, resulting in gaps in filamentous algae mats. We analyzed whether those gaps may be caused by allelopathic substances excreted by S. aloides or by nutrient depletion. We studied in a field survey the colonization of natural S. aloides by filamentous algae and determined in situ nutrient concentrations in natural S. aloides stands. To analyze the relative importance of allelopathy and nutrient competition in the interaction between S. aloides and filamentous algae, we carried out field experiments. Introduction of artificial (non-allelopathic) plants in natural S. aloides stands enabled us to compare the colonization by filamentous algae of both Stratiotes sp. and artificial plants. The filamentous algae were absent in close vicinity to S. aloides. Significantly lower concentrations of ortho-phosphate and potassium were observed close to S. aloides as compared with the filamentous algae. In the field experiments the artificial plants were rapidly colonized by filamentous algae, mainly Cladophera Kützing and Spirogyra Link, while all natural plants remained free of such algae. Additionally, most nutrient concentrations did not significantly differ in the proximity of artificial or natural stands of S. aloides. The concentrations of the major growth-limiting nutrients, phosphate and nitrate, were significantly higher and nonlimiting in natural Stratiotes stands. Our main conclusion is that, although allelopathic interactions between S. aloides and filamentous algae do occur under natural conditions, nutrient competition between the two can also be an important factor.https://doi.org/10.1007/s10452-008-9194-7Re-oligotrophication by phosphorus reduction and effects on seston quality in lakes
We review published data of re-oligotrophication studies in different types of lakes and discuss whether re-oligotrophication by phosphorus reduction measures can affect seston quality in lakes due to an increase in C:P ratios of food for zooplankton. We analyze whether such an increase in elemental ratio may give rise to a stoichiometric constraint or bottleneck for regulation of phytoplankton by zooplankton and thus cause a decrease in trophic transfer efficiency. Because seston C:P ratios of seston tend to decrease with increasing P concentration in lakes, i.e. by increasing eutrophication, the reverse may be true, i.e. the C:P ratio of the seston will increase when lakes become re-oligotrophicated by P reduction. This may imply lower food quality for P-demanding seston grazers, like Daphnia, which have both relatively low C:P ratios and are quite homeostatic with regard to C:P ratios. Consequently, the reduced grazer biomass may lead to lower grazing pr Most investigated lakes – both deep and shallow – responded by improvement in underwater light climate due to reduction of P loadings. Several compensatory mechanisms prevented the occurrence of a stoichiometric bottleneck for top-down control. From a few documented cases of both deep and shallow lakes, where seston stoichiometry, the presence of Daphnia and the trophic transfer efficiency were concurrently measured during the process of re-oligotrophication, we may conclude that the anticipatedhttps://doi.org/10.1016/j.limno.2008.05.005Adaptation of the Fungal Parasite Zygorhizidium planktonicum During 200 Generations of Growth on Homogeneous and Heterogeneous Populations of Its Host, the Diatom Asterionella formosa
We followed adaptation of the chytrid parasite Zygorhizidium planktonicum during 200 generations of growth on its host, the freshwater diatom Asterionella formosa, in a serial passage experiment. Evolution of parasite fitness was assessed both on a homogenous and heterogeneous host population, consisting of respectively a single new and ten different new host strains. These 10 host strains were genetically different and also varied in their initial susceptibility to the parasite. Parasite fitness increased significantly and rapidly on the new, genetically homogenous host population, but remained unaltered during 200 generations of growth on the heterogeneous host population. Enhanced parasite fitness was the result of faster and more efficient transmission, resulting in higher values of R0 (number of secondary infections). Consequently, parasites that evolved within the uniclonal host population infected significantly more of these hosts than did their ancestors. We thus provide experimental evidence for the widely held view that host genetic diversity restricts evolution of parasites and moderates their harmful effects. Genetically uniform host populations are not only at increased risk from fungal epidemics because they all share the same susceptibility, but also because new parasite strains are able to adapt quickly to new host environments and to improve their fitness.https://doi.org/10.1111/j.1550-7408.2008.00306.xEffects of nutrients and fish on periphyton and plant biomass across a European latitudinal gradient
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 València (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.https://doi.org/10.1007/s10452-007-9126-yLake restoration studies: failures, bottlenecks and prospects of new ecotechnological measures
This paper critically reviews the published works on lake restoration in north-western Europe, with the aim to highlight the causes of failures of lake biomanipulation, and to identify the main bottlenecks that have impeded progress. More importantly, we explore the prospects of applying new ecotechnological measures to lakes with a focus on shallow lakes. These complementary measures are: (1) reduction of sediment resuspension; (2) water-level management; and (3) the use in shallow lakes of bivalves as effective grazers on lake seston, especially when cyanobacteria are dominant. If the sustainability of the positive effects of biomanipulation is considered over a decade, there are probably more cases of failures than successes. The failures can be ascribed to several bottlenecks that include: (1) inadequate reduction of allochthonous phosphorus (P) and an increase in autochthonous P inputs, i.e. release of P from the lake sediments following reductions of external P inputs; (2) poor edibility of filamentous and colonial cyanobacteria to daphnids; (3) inadequate coverage of the lake area by macrophytes partly due to foraging on the macrophytes by both fish and birds; (4) ineffective reduction of planktivorous fish biomass and our inability to maintain the fish mass to a ‘low level’ for longer periods; and (5) failure of northern pike (Esox lucius) after its transplantation to the lakes to develop a population level that can control planktivorous fish to desired low levels. Three potentially complementary ecotechnological measures are discussed. The first such measure concerns prevention of sediment resuspension in lakes by creating islands in order to minimise the wind fetch to reduce the wave amplitude. The second measure involves allowing greater water-level fluctuations (WLF) in lakes as planned in lowland countries like the Netherlands; WLF are likely to allow more space for water, and may lead to improved water quality and higher biodiversity. The third ecotechnological measure relates to grazer populations that complement herbivorous zooplankton to regulate phytoplankton, particularly to control cyanobacterial blooms. For this, the bivalve Dreissena polymorpha appears to be a good potential candidate for grazing on phytoplankton, especially in shallow eutrophic lakes that are dominated by filamentous and toxic cyanobacteria (e.g. Planktothrix agardhii and Microcystis aeruginosa).https://doi.org/10.1016/j.limno.2008.05.008Functional response of Anodonta anatina feeding on a green alga and four strains of cyanobacteria, differing in shape, size, and toxicity
We studied the functional response of the freshwater unionid bivalve Anodonta anatina, feeding on five phytoplankton strains differing in food quality: the small green alga Scenedesmus obliquus, a toxic and a non-toxic strain of the filamentous cyanobacterium Planktothrix agardhii and a toxic and a non-toxic strain of the coccoid cyanobacterium Microcystis aeruginosa. On S. obliquus, A. anatina had a type II functional response with a maximum mass-specific ingestion rate (IRmax) of 5.24 mg C g DW−1 h−1 and a maximum mass-specific clearance rate (CRmax) of 492 (±38) ml g DW−1 h−1, the highest values for all the phytoplankton strains that were investigated. On toxic and non-toxic P. agardhii filaments, A. anatina also had a type II functional response, but IRmax and CRmax were considerably lower (IRmax 1.90 and 1.56 mg C g DW−1 h−1; CRmax 387 (±97) and 429 (±71) ml g DW−1 h−1, respectively) than on S. obliquus. Toxicity of P. agardhii had no effect on the filtration rate of the mussels. On the non-toxic M. aeruginosa (small coccoid cells), we also observed a type II functional response, although a type I functional response fitted almost as good to these data. For the colonial and toxic M. aeruginosa, a type I functional response fitted best to the data: IR increased linearly with food concentration and CR remained constant. CRmax and IRmax values for the (colonial) toxic M. aeruginosa (383 (±40) ml g DW−1 h−1; 3.7 mg C g DW−1 h−1) demonstrated that A. anatina filtered and ingested this cyanobacterium as good as the other cyanobacterial strains. However, on the non-toxic M. aeruginosa we observed the lowest CRmax of all phytoplankters (246 (±23) ml g DW−1 h−1, whereas IRmax was similar to that on toxic M. aeruginosa. The high maximum ingestion rates on S. obliquus and M. aeruginosa indicate a short handling time of these phytoplankton species. The high clearance rates on S. obliquus, toxic M. aeruginosa and P. agardhii reflect a high effort of the mussels to filter these particles out of the water column at low concentrations. The low clearance rates on non-toxic M. aeruginosa may be explained by the small size and coccoid form of this cyanobacterium, which may have impaired A. anatina to efficiently capture the cells. Although A. anatina had relatively high maximum clearance rates on non-toxic and toxic P. agardhii, this cyanobacterium does not seem to be a good food source, because of the observed high rates of pseudofaeces production and hence low ingestion rates.https://doi.org/10.1007/s10750-007-0580-2Parasitic chytrids: their effects on phytoplankton communities and wood-web dyamics
Many phytoplankton species are susceptible to fungal parasitism. Parasitic fungi of phytoplankton mainly belong to the Chytridiomycetes (chytrids). Here, we discuss the progression made in the study of chytrids that parasitize phytoplankton species. Specific fluorescent stains aid in the identification of chytrids in the field. The established culturing methods and the advances in molecular science offer good potential to gain a better insight into the mechanisms of epide-zic development of chytrids and coevolution between chytrids and their algal hosts. Chytrids are often considered to be highly host-specific parasites, but the extent of host specificity has not been fully investigated. Chytrids may prefer larger host cells, since they would gain more resources, but whether hosts are really selected on the basis of size is not clear. The dynamics of chytrids epidemics in a number of studies were partly explained by environmental factors such as light, temperature, nutrients, pH, turbulence and zooplankton grazing. No generalization was made about the epidemic conditions; some state unfavorable conditions for the host growth support epidemic development, while others report epidemics even under optimal growth conditions for the host. Phytoplankton is not defenseless, and several mechanisms have been suggested, such as a hypersensitivity response, chemical defense, maintaining a high genetic diversity and multitrophic indirect defenses. Chytrids may also play an important role in food webs, because zoospores of chytrids have been found to be a good food source for zooplankton.https://doi.org/10.1007/s10750-006-0438-zMacrophyte-phytoplankton interactions: the relative importance of allelopathy versus other factors
Abstract Submerged aquatic macrophytes are important in shallow aquatic ecosystems because they stabilize the macrophyte-dominated state by increasing water transparency in various ways. One of these is the excretion of allelopathic substances inhibitory to phytoplankton, but it is still controversial whether this mechanism can be important in the field. We developed a model that describes phytoplankton growth including the effects of two different macrophytes (Chara sp. and Stratiotes aloides), which ar Both macrophyte species had a considerable effect on phosphorus levels in the water. Shading of phytoplankton by Chara sp. was estimated to be negligible. This macrophyte species has a growth form close to the sediment. The allelopathic activity of Chara sp. caused a phytoplankton growth reduction of ca. 5–10%, while the most pronounced effect seems to be related to reduction of resuspension. S. aloides floats on the water surface during spring and summer. Therefore, it has a potentially high shading effect. Additionally, this species has a relatively strong allelopathic capacity, sometimes resulting in a reduction of phytoplankton between 50 and 80%. The model suggests that the relative contribution of allelopathy in situ is low or negligible for charophytes. Only for macrophyte species that have a high allelopathic potential, like S. aloides, it may be a significant effect.https://doi.org/10.1016/j.ecolmodel.2006.12.020Chemical information transfer in freshwater plankton
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.https://doi.org/10.1016/j.ecoinf.2007.03.002Goose-mediated nutrient enrichment and planktonic grazer control in arctic freshwater ponds
A dramatic increase in the breeding population of geese has occurred over the past few decades at Svalbard. This may strongly impact the fragile ecosystems of the Arctic tundra because many of the ultra-oligotrophic freshwater systems experience enrichment from goose feces. We surveyed 21 shallow tundra ponds along a gradient of nutrient enrichment based on exposure to geese. Concentrations of total phosphorus (P) and dissolved inorganic nitrogen (DIN) in the tundra ponds ranged from 2–76 to 2–23 μg l−1 respectively, yet there was no significant increase in phytoplankton biomass (measured as chlorophyll a; range: 0.6–7.3 μg l−1) along the nutrient gradient. This lack of response may be the result of the trophic structure of these ecosystems, which consists of only a two-trophic level food chain with high biomasses of the efficient zooplankton grazer Daphnia in the absence of fish and scarcity of invertebrate predators. Our results indicate that this may cause a highly efficient grazing control of phytoplankton in all ponds, supported by the fact that large fractions of the nutrient pools were bound in zooplankton biomass. The median percentage of Daphnia–N and Daphnia–P content to particulate (sestonic) N and P was 338 and 3009%, respectively, which is extremely high compared to temperate lakes. Our data suggest that Daphnia in shallow arctic ponds is heavily subsidized by major inputs of energy from other food sources (bacteria, benthic biofilm), which may be crucial to the persistence of strong top–down control of pelagic algae by Daphnia.https://doi.org/10.1007/s00442-007-0770-7The influence of Myriophyllum verticillatum and artificial plants on some life history parameters of Daphnia magna
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.https://doi.org/10.1007/s10452-007-9091-5Daphnia growth rates in Arctic ponds: limitation by nutrients or carbon?
Arctic organisms with annual life cycles experience a strong selective pressure to fulfill their life cycle at low temperatures within a short seasonal window. Yet, apart from low temperature, the factors that constrain or promote growth rates in high arctic systems are still poorly understood. A substantial part of the freshwater systems in the arctic consist of shallow, fish-free ponds with the crustacean Daphnia as the key grazer. This grazer has high demands for phosphorus (P) for RNA-synthesis and subsequently protein synthesis for growth. In this study, we compared growth of juvenile Daphnia that were fed seston from two high-Arctic (79°N) ponds on Svalbard in 2004, which differed strongly in P-content and C:P-ratios. In both ponds, Daphnia growth was limited by food quantity (carbon) rather than by P or N. The study also suggests that in absence of predators, infection level of epibionts might be an important factor regulating growth rate and population development of Daphnia growth in these systems.https://doi.org/10.1007/s00300-006-0177-7Relaxed circadian rhythm in zooplankton along a latitudinal gradient
To test whether aquatic invertebrates are able to adjust their diel migratory cycle to different day length and presence of predators, we performed standardized enclosure experiments in shallow lakes at four different latitudes from southern Spain, with strong night–day cycles, to Finland where daylight is almost continuous during summer. We show here that nearly continuous daylight at high latitudes causes a relaxation in diel migratory behaviour in zooplankton irrespective of predation risk. At lower latitudes, however, similar conditions lead to pronounced diel rhythms in migration. Hence, zooplankton may show local behavioural adaptations in their circadian rhythm. They are also able to make risk assessments as to whether diel migration is beneficial or not, manifested in a lack of diel migration at near constant daylight, irrespective of predator presence. Our results provide an additional explanation to previous knowledge regarding diel migrations among aquatic invertebrates by showing that both physical (light) and biological (predation) factors may affect the migratory behaviour.https://doi.org/10.1111/j.2006.0030-1299.15754.x,Grazing on microcystin-producing and microcystin-free phytoplankters by different filter-feeders: implications for lake restoration
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.https://doi.org/10.1007/s00027-007-0916-zAllelopathic effect of the aquatic macrophyte, Stratiotes aloides, on natural phytoplankton
A survey of different Dutch Stratiotes stands showed that the density of phytoplankton (except cyanobacteria) was always higher outside S. aloides than between the rosettes of S. aloides. Analyses of water samples revealed that nutrient limitation was unlikely to have caused the lower phytoplankton biomass in the vicinity of S. aloides. 2. An in situ incubation experiment in the Danube Delta, Romania, indicated allelopathic activity against phytoplankton in S. aloides stands. The growth rate of natural phytoplankton populations exposed to water from S. aloides stands was significantly lower than that of populations that had not been in contact with S. aloides exudates. 3. A laboratory microcosm experiment showed a significantly lower phytoplankton biomass in treatments with S. aloides exudates. Nutrient concentrations and the light intensity were high enough that the lower phytoplankton biomass could not be explained by nutrient or light limitation. [KEYWORDS: allelopathy ; exudates ; field experiment ; infochemical ; natural phytoplankton]https://doi.org/10.1111/j.1365-2427.2006.01510.xInfochemicals structure marine, terrestrial and freshwater food webs: implications for ecological informatics
Here we consider how information transfer shapes interactions in aquatic and terrestrial food webs. All organisms, whether they are dead or alive, release certain chemicals into their environment. These can be used as infochemicals by any other individual in the food web that has the biological machinery to sense and process such information. Such machinery has evolved in bacteria, plants and animals and has thus become an inextricable part of the mechanisms that underlie feeding relations in food webs. Organisms live in environments suffused with infochemicals and this information network can be tapped into by both predators and their prey. However, it also opens doors to confusion in the face of a bewildering abundance and complexity of information. Infochemical mixing, masking, crypsis and mimicry could cause such confusion, especially in species-rich communities. We provide a point of entry into this field of enquiry by identifying seminal papers and major reviews and by discussing research lines that might enhance our mechanistic understanding of interactions in food webs. We highlight empirical work on the ways in which individuals use infochemicals and discuss model results on how this mediates patterns of population dynamics. We consider implications for ecosystem management and indicate how classical models and novel approaches from ecological infor [KEYWORDS: Biodiversity ; Biological control ; Climate ; Dimethyl sulphide ; Global warming ; Individual based models ; Information networks in ecosystems ; Integration of laboratory and field data ; Learning ; Linking levels of ecological organization ; Lake restoration ; Phenotypic plasticity ; Trait-mediated interactions]https://doi.org/10.1016/j.ecoinf.2005.06.001Contrasting life history responses to fish released infochemicals of two co-occurring Daphnia species that show different migration behaviour
In a previous field study (Flik & Vijverberg 2003) we showed that in an oligotrophic-mesotrophic lake in the Netherlands (L. Maarsseveen) two co-occurring Daphnia species, Daphnia pulicaria and Daphnia galeata × hyalina performed distinct vertical migration behaviour during summer which is induced by a high fish biomass of young perch. During late spring and summer, D. galeata × hyalina exhibits diel vertical migration, whereas D. pulicaria is staying down day and night in the hypolimnion. This difference in migration behaviour results in contrasting predation mortalities caused by fish, low for D. pulicaria and relatively high for D. galeata × hyalina. In the present study, we measured in the laboratory the effects of fish released infochemicals on five life history traits in four genetically distinct D. galeata × hyalina and three genetically distinct D. pulicaria clones collected during summer at day-time from the hypolimnion. We tested the hypothesis that the species which behaviour is providing the best protection against fish predation (i.e. D. pulicaria) is less protected by life history traits induced by fish released infochemicals than the species which by its behaviour is less well protected against fish predation (i.e. D. galeata × hyalina). Our results show that D. galeata × hyalina responded in three out of five traits differently to fish infochemicals than D. pulicaria. In all these three traits D. galeata × hyalina responded significantly to fish-released infochemicals, whereas D. pulicaria did not show any significant response at all. We conclude that in D. pulicaria behavioural defences trade-off against life history defences.https://doi.org/10.1127/0003-9136/2006/0167-0089Comparing grazing on lake seston by Dreissena and Daphnia: lessons for biomanipulation
Biomanipulation measures in lakes, taken to diminish algal blooms, have mainly been restricted to the reduction of zooplanktivorous fish with the aim to stimulate the grazing pressure by native filter feeders such as Daphnia. However, larger filter feeders like the exotic zebra mussel, Dreissena polymorpha, have been suggested as an optional tool because of their high filtering capacity. We compared grazing by two filter feeders, D. polymorpha and Daphnia galeata, offered seston from Lake IJsselmeer, the Netherlands in two consecutive years: 2002 and 2003. The seston in both years was dominated by the colony-forming cyanobacterium Microcystis aeruginosa. The grazing studies were performed under controlled conditions in the laboratory and samples were analyzed on a flow cytometer, making it possible to quantify grazing on different seston components and size fractions, including cyanobacteria, other phytoplankton (green algae, diatoms, etc.), and detritus. No differences in clearance rates, on a per weight basis, were found between the two grazer species. The clearance rate on cyanobacteria (especiallyhttps://doi.org/10.1007/s00248-004-0147-6Seasonal seston stoichiometry: effects on zooplankton in cyanobacteria-dominated lakes
Seasonal dynamics in elemental composition [carbon (C), nitrogen and phosphorus (P)] of seston and zooplankton were studied over several years in three hypereutrophic Dutch lakes with persistent dominance and high biomass of cyanobacteria. In all three lakes, there was a strong pattern with decreased P-content and increased C:P ratio in seston (https://doi.org/10.1093/plankt/fbi018Allelopathic inhibition of phytoplankton by exudates from Stratiotes aloides
The allelopathic potential of exudates from the aquatic macrophyte Stratiotes aloides on the growth of phytoplankton was investigated. A selection of phytoplankton species, occurring in habitats similar to that of Stratiotes, was used: two cyanobacterial strains (toxic and non-toxic Microcystis aeruginosa), one green alga (Scenedesmus obliquus) and one eustigmatophyte (Nannochloropsis limnetica). The results indicate allelopathic effects of Stratiotes on phytoplankton in six of the eight cases, expressed in an extended duration of the initial biovolume doubling time. The overall inhibitory effect (8–51%) was strain-specific for the two cyanobacteria. We also studied the effect of irradiance on the allelopathic potential of exudates from Stratiotes. Irradiance influenced the response of Scenedesmus only. The inhibitory effect of Stratiotes exudates on the growth of this green alga was stronger at 35 µmol m-2 s-1 than at 105 µmol m-2 s-1. We conclude that Stratiotes has allelopathic effects on phytoplankton, and that irradiance can, but does not always determine the extent of the allelopathic inhibition. In our experiments, the sensitivity of cyanobacteria to Stratiotes exudates was not higher than for other phytoplankton strains, but within cyanobacteria, the toxic strain was more sensitive than the non-toxic one. [KEYWORDS: Allelopathy ; Exudate ; Growth inhibition ; Initial biovolume doubling time ; Irradiance ; Lag phase]https://doi.org/10.1016/j.aquabot.2005.04.001Allelopathic growth inhibition and colony formation of the green alga Scenedesmus obliquus by the aquatic macrophytes Stratiotes aloides
Laboratory experiments were conducted to elucidate the allelopathic effects of exudates from the aquatic macrophyte Stratiotes aloides on the growth and morphology of the green alga Scenedesmus obliquus. Both water originating from a S. aloides culture and water that had not been in contact with S. aloides was simultaneously inoculated with S. obliquus cells. In all experiments we observed a highly significant inhibitory effect of Stratiotes water on the green alga. The duration of the lag phase as well as the initial biovolume doubling time of S. obliquus were extended. The growth rate of S. obliquus was only significantly inhibited by the younger Stratiotes plants. Furthermore, the mean particle volume (MPV) of Scenedesmus increased significantly in the presence of Stratiotes water in all three experiments. Microscopic analyses confirmed that S. obliquus forms more colonies in the presence of water from a S. aloides culture. Colonies of phytoplankton have higher sinking rates than single phytoplankton cells of the same species and disappear faster from the upper water layers, reducing the competition for light between algae and macrophytes. [KEYWORDS: Colony formation ; Growth inhibition ; Infochemical ; Macrophytes ; Sedimentation]https://doi.org/10.1007/s10452-004-1021-1The role of climate in shaping zooplankton communities of shallow lakes
We analyzed data from 81 shallow European lakes, which were sampled with standardized methods, for combined effects of climatic, physical, and chemical features of food-web interactions, with a specific focus on zooplankton biomass and community structure. Multiple-regression analysis showed that total phosphorus (TP) generally was the most important predictor of zooplankton biomass and community structure. Climate was the next most important predictor and acted mainly through its effect on pelagic zooplankton taxa. Benthic and plant-associated taxa (typically almost half the total zooplankton biomass) were, however, affected mainly by macrophyte coverage. Neither climate nor TP affected the relation between small and large taxa, and we found only a weak trend with increasing TP of increasing mean crustacean body mass. Dividing the data set into three climate zones revealed a pronounced difference in response to lake productivity between cold lakes, with long periods of ice cover, and the two warmer lake types. These ‘‘ice’’ lakes differed from the others with respect to the effect of TP on chlorophyll a, the zooplankton : chlorophyll a ratio, the chlorophyll a :TP ratio, and the proportion of cyclopoids in the copepod community. Our data suggest that bottom-up forces, such as nutrient concentration, are the most important predictors of zooplankton biomass. In addition, climate contributes significantly—possibly by affecting top-down regulation by fish—and may interact with productivity in determining the zooplankton standing biomass and community composition. Hence, the present study suggests that food-web dynamics are closely linked to climatic features.https://doi.org/10.4319/lo.2005.50.6.2008Grazing on colonial and filamentous, toxic and non-toxic cyanobacteria by the zebra mussel Dreissena polymorpha
Colony forming and toxic cyanobacteria form a problem in surface waters of shallow lakes, both for recreation and wildlife. Zebra mussels, Dreissena polymorpha, have been employed to help to restore shallow lakes in the Netherlands, dominated by cyanobacteria, to their former clear state. Zebra mussels have been present in these lakes since they were created in the 19th century by the excavation of peat and are usually not considered to be an invasive species. Most grazing experiments using Dreissena have been performed with uni-cellular phytoplankton laboratory strains and information on grazing of larger phytoplankton taxa hardly exists. To gain more insight in to whether D. polymorpha is indeed able to decrease cyanobacteria in the phytoplankton, we therefore performed grazing experiments with zebra mussels and two species of cyanobacteria, that greatly differ in shape: colony forming strains of Microcystis aeruginosa and the filamentous species Planktothrix agardhii. For both species a toxic and a non-toxic strain was selected. We found that zebra mussels cleared toxic Planktothrix at a higher rate than non-toxic Planktothrix, toxic or non-toxic Microcystis. Clearance rates between the other strains were not significantly different. Both phytoplankton species, regardless of toxicity, size and shape, were found in equal amounts (based on chlorophyll concentrations) in the excreted products of the mussels (pseudofaeces). The results show that zebra mussels are capable of removing colonial and filamentous cyanobacteria from the water, regardless of whether the cyanobacteria are toxic or not. This implies that the mussels may be used as a biofilter for the removal of harmful cyanobacterial blooms in shallow (Dutch) lakes where the mussels are already present and not a nuisance. Providing more suitable substrate for zebra mussel attachment may lead to appropriate mussel densities capable of filtering large quantities of cyanobacteria.https://doi.org/10.1093/plankt/fbi008Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment
1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year-to-year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 g TP L1) when grazer biomass was high (>8090 g dry mass L1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (1723 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation. [KEYWORDS: eutrophication ; fish predation ; littoral zooplankton ; mesocosm experiments ; nutrient enrichment]https://doi.org/10.1111/j.1365-2427.2004.01300.xEffects of fish and nutrient additions on food-web stability in a charophyte-dominated lake
1. The response of major food-web constituents to combinations of nutrient addition and zooplanktivorous fish abundance was tested during two subsequent years in the shallow charophyte-dominated lake Naardermeer in the Netherlands, using in situ enclosures. 2. Treatment effects differed sharply between study years. In 1998, when the summer temperature was low (1721 °C), high algal biomass only developed at high nutrient levels in the presence of fish, but with no major effect on Chara biomass. In 1999, when the summer temperature was relatively high (2025 °C), algal blooms occurred at high nutrient levels regardless of fish abundance, and were associated with a drastic decline in Chara biomass. 3. Differences between years in temperature and initial zooplankton composition and biomass were likely to contribute to the varying relative importance of top-down and bottom-up effects in these enclosure experiments. 4. The results suggest that when nutrient loads are increased towards levels where the macrophyte-dominated state is being destabilised, a 'switch' is more likely to occur in warm summers. [KEYWORDS: enclosures ; multiple stable states ; shallow lakes ; submerged macrophytes ; turbidity]https://doi.org/10.1111/j.1365-2427.2004.01301.xGenetic variation in Asterionella formosa (Bacillariophyceae) is it linked to frequent epidemics of host-specific parasitic fungi?
Understanding of the genetic basis for susceptibility and resistance is still lacking for most aquatic host-parasite systems, for instance, for phytoplankton and their fungal parasites. Fungal parasites can have significant effects on phytoplankton populations, mainly through their ability to decimate algal host populations during epidemics. We used random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP) analysis to study levels of genetic variation within a population of the freshwater diatom Asterionella formosa Hassall in relation to parasitism by the obligate, host-specific, fungal parasite Zygorhizidium planktonicum Canter. The level of genetic variation within the A. formosa population in Lake Maarsseveen, The Netherlands was found to be high despite the presumed absence or very low frequency of sexual reproduction in this species, the limited gene flow, and the severity of parasite attack that would purge the population from susceptible genotypes. RAPD analysis revealed four distinct banding patterns, with 3 of 21 markers (14%) being polymorphic. In AFLP analysis, every single isolate of A. formosa showed a unique banding pattern, and 120 of the 210 AFLP markers (57%) were found to be polymorphic. Furthermore, character compatibility analysis revealed that sexual reproduction may be one of the mechanisms that generates and maintains genetic variation in the A. formosa population in Lake Maarsseveen. The presence of genetic variation in A. formosa was reflected in infection experiments, which showed that genetically different A. formosa strains differed in their susceptibility to various Z. planktonicum strains and that parasite strains differed in their ability to infect particular host strains. [KEYWORDS: AFLP; genetic variation; parasitism; RAPD; Red Queen; sexual reproduction]https://doi.org/10.1111/j.1529-8817.2004.04006.xDaphnia can protect diatoms from fungal parasitism
Many phytoplankton species are susceptible to chytrid fungal parasitism. Much attention has been paid to abiotic factors that determine whether fungal infections become epidemic. It is still unknown, however, how biotic factors, such as interactions with zooplankton, affect the fungal infection process. Because the size of fungal zoospores is well within the preferred range of food particle size for Daphnia, we hypothesize that Daphnia will affect the fungal infection process by reducing the abundance of fungal zoospores. To examine the effects of zooplankton on the fungal parasitism of phytoplankton, we performed grazing experiments. These experiments revealed that Daphnia significantly decreased fungal parasitism by grazing on the fungal zoospores. Daphnia also had a small positive effect on fungal infection, probably by increasing the encounter rate between fungi and host phytoplankton cells. These results suggest that Daphnia can affect the seasonal succession of chytrids and their host phytoplankton species. In addition, these results imply that zoospore-producing fungi may play an ecological role as food sources for Daphnia in aquatic food webs.https://doi.org/10.4319/lo.2004.49.3.0680Assimilation and depuration of microcystin–LR by the zebra mussel, Dreissena polymorpha
Zebra mussels (Dreissena polymorpha) are an important component of the foodweb of shallow lakes in the Netherlands, amongst others in Lake IJsselmeer, an international important wetland. Large numbers of ducks feed on these mussels in autumn and winter. The mussels are filter feeders and are exposed to high densities of cyanobacteria in summer and autumn. Mussels and cyanobacteria both thrive in Lake IJsselmeer. Apparently the mussels are somehow protected against accumulation of harmful quantities of cyanobacterial toxins. In this study, we investigated the assimilation of the cyanobacterial toxin microcystin–LR (MC–LR) in zebra mussels when fed the toxic cyanobacterium Microcystis aeruginosa as sole food or in a mixture with the eustigmatophyte Nannochloropsis limnetica. After 3 weeks of assimilation we studied the depuration of MC–LR during 3 weeks when the food of the mussels was free of cyanobacteria. These assimilation/depuration experiments were combined with grazing experiments, using the same food treatments. Microcystins were analyzed using liquid chromatography–mass spectrometry (LC–MS); in addition, covalently bound MC were analyzed using the MMPB method. The mussels showed higher clearance rates on Microcystis than on Nannochloropsis. No selective rejection of either phytoplankton species was observed in the excretion products of the mussels. Zebra mussels fed Microcystis as single food, assimilated microcystin–LR relatively fast, and after 1 week the maximum value of free unbound microcystin assimilation (ca. 11 µg g DW-1) was attained. For mussels, fed with the mixed food, a maximum of only 3.9 µg g DW-1 was recorded after 3 weeks. Covalently bound MC never reached high values, with a maximum of 62% of free MC in the 2nd week of the experiment. In the depuration period microcystin decreased rapidly to low values and after 3 weeks only very low amounts of microcystin were detectable. The amount of toxin that accumulated in the mussels would appear to be high enough to cause (liver) damage in diving ducks. However, death by exposure to microcystin seems unlikely. Mussels seem efficient in minimizing the assimilation of microcystin. If it were not for this, mass mortalities of ducks in shallow lakes in the Netherlands would presumably occur on a much more widespread scale than is currently observed. [KEYWORDS: Clearance rate; Cyanotoxins; Diving ducks; Food web; LC–MS; Microcystis; Phytoplankton; Pseudo-faeces]https://doi.org/10.1016/j.aquatox.2004.06.004Continental-scale patterns of nutrient and fish effects on shallow lakes: synthesis of a pan-European mesocosm experiment
1. Results are analysed from 11 experiments in which effects of fish addition and nutrient loading on shallow lakes were studied in mesocosms. The experiments, five in 1998, six in 1999, were carried out in six lakes, distributed from Finland to southern Spain, according to a standard protocol. 2. Effects of the treatments on 29 standard chemical, phytoplankton and zooplankton variables are examined to assess the relative importance of bottom-up (nutrient enrichment) and top-down (fish predation) effects. For each year, the experiments in different locations are treated as replicates in a meta-analysis. Results of individual experiments are then compared in terms of the patterns of significant influences of nutrient addition and fish predation with these overall results (the baseline), 3. The overall meta-analysis gave consistent results across the 2 years, with nutrient loading influencing all of the chemical variables, and on average 31% of primary producer and 39% of zooplankton variables. In contrast, fish influenced none of the chemical variables, 11% of the primary producer and 44% of the zooplankton variables. Nutrient effects on the system were thus about three times greater than fish effects, although fish effects were not inconsiderable. 4. The relative importance of nutrients and fish in individual experiments often differed between years at the same location and effects deviated to varying degrees from the baseline. These deviations were treated as measures of consistency (predictability) of conclusions in repeat experiments. Consistency increased southwards and this is interpreted as a consequence of more variable annual weather northwards. 5. The influence of nutrient loading was greater southwards and this was probably manifested through naturally greater annual macrophyte abundance in warmer locations in consequence of the longer plant growing-season. There was no trend in the relative importance of fish effects with latitude but this may partly be an artefact of the simple fish community used. These findings suggest that nutrient control should be a greater priority than biomanipulation in the restoration of eutrophicated shall 6. Starting conditions affected the outcome of experiments. High initial concentrations of total phosphorus and planktonic chlorophyll a concentration (created by local conditions prior to the experiment) led to de-emphasis of the importance of nutrient loading in the experiment. [KEYWORDS: continental gradient ; fish ; meta-analysis ; nutrients ; weather variation]https://doi.org/10.1111/j.1365-2427.2004.01304.xInducible colony formation within the Scenedesmaceae: adaptive responses to infochemicals from two different herbivore taxa
We studied the occurrence of colony formation within 40 different strains of Scenedesmaceae (Chlorococcales, Chlorophyta) in response to grazing-released infochemicals from the herbivorous zooplankters Brachionus calyciflorus Pallas (Rotifera) and Daphnia magna Strauss (Cladocera). With the exception of two strains, all strains showed similar responses to both B. calyciflorus and D. magna infochemicals, either no response or inducible colony formation. Colony size was found to increase with B. calyciflorus infochemical concentration and could be described by a sigmoid function. The increase in colony size was more pronounced in the Scenedesmus species tested than in Desmodesmus species, which was probably due to higher threshold infochemical concentrations for colony induction in Desmodesmus. Therefore, the adaptivity of colony formation to the herbivory threat only holds above the threshold concentration for colony induction and as long as maximum colony size has not been attained. Taking this into account, our results suggest that inducible colony formation is a common adaptive response of many Scenedesmaceae to the threat of herbivory. [KEYWORDS: Brachionus calyciflorus; Daphnia magna; Desmodesmus; herbivory; inducible defences; infochemicals ; Scenedesmus]https://doi.org/10.1111/j.1529-8817.2004.04007.xContinental-scale patterns of nutrient and fish effects on shallow lakes: introduction to a pan-European mesocosm experiment
1. Shallow lake ecosystems are normally dominated by submerged and emergent plants. Biological stabilising mechanisms help preserve this dominance. The systems may switch to dominance by phytoplankton, however, with loss of submerged plants. This process usually takes place against a background of increasing nutrient loadings but also requires additional switch mechanisms, which damage the plants or interfere with their stabilising mechanisms. 2. The extent to which the details or even major features of this general model may change with geographical location are not clear. Manipulation of the fish community (biomanipulation) has often been used to clear the water of algae and restore the aquatic plants in northerly locations, but it is again not clear whether this is equally appropriate at lower latitudes. 3. Eleven parallel experiments (collectively the International Mesocosm Experiment, IME) were carried out in six lakes in Finland, Sweden, England, the Netherlands and Spain in 1998 and 1999 to investigate the between-year and large-scale spatial variation in relationships between nutrient loading and zooplanktivorous fish on submerged plant and plankton communities in shallow lakes. 4. Comparability of experiments in different locations was achieved to a high degree. Cross-laboratory comparisons of chemical analyses revealed some systematic differences between laboratories. These are unlikely to lead to major misinterpretations. 5. Nutrient addition, overall, had its greatest effect on water chemistry then substantial effects on phytoplankton and zooplankton. Fish addition had its major effect on zooplankton and did not systematically change the water chemistry. There was no trend in the relative importance of fish effects with latitude, but nutrient addition affected more variables with decreasing latitude. 6. The relative importance of top-down and bottom-up influences on the plankton differed in different locations and between years at the same location. The outcome of the experiments in different years was more predictable with decreasing latitude and this was attributed to more variable weather at higher latitudes that created more variable starting conditions for the experiments. [KEYWORDS: alternative stable states ; community structure ; eutrophication ; fish ; large-scale variation ; nutrients]https://doi.org/10.1111/j.1365-2427.2004.01302.xSelective grazing by adults and larvae of the zebra mussel (Dreissena polymorpha): application of flow cytometry to natural seston
1. Selective grazing of adults and larvae of the zebra mussel (Dreissena polymorpha) on phytoplankton and detritus from both laboratory cultures and natural seston was quantified using flow cytometry. 2. Mean clearance rate of adult zebra mussels was higher on a mixture of the green alga Scenedesmus and the cyanobacterium Microcystis than when Scenedesmus was offered as single food, suggesting selective feeding by the mussels. 3. Feeding on lake seston both adults and larvae showed a higher clearance rate on phytoplankton than on detritus particles, suggesting that zebra mussels select for phytoplankton. Furthermore, it was noted that adults preferred seston particles in the 0-1 and 30-100 m size ranges. 4. In our study, zebra mussels did not discriminate against cyanobacteria, and our results indicate that they may even ingest them preferentiallyhttps://doi.org/10.1046/j.1365-2426.2003.01173.xResponses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment
1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte-dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L1 P and 10 mg L1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within-treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD <50 m). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes. [KEYWORDS: fish ; food-web interactions ; mesocosm experiments nutrients ; phytoplankton composition]https://doi.org/10.1111/j.1365-2427.2004.01307.xHost parasite interactions between freshwater phytoplankton and chytrid fungi (Chytridiomycota)
Some chytrids are host-specific parasiticfungithat may have a considerable impact on phytoplankton dynamics. The phylum Chytridiomycota contains one class, the Chytridiomycetes, and is composed of five different orders. Molecular studies now firmly place the Chytridiomycota within the fungal kingdom. Chytrids are characterized by having zoospores, a motile stage in their life cycle. Zoospores are attracted to the host cell by specific signals. No single physical-chemical factor has been found that fully explains the dynamics of chytrid epidemics in the field. Fungal periodicity was primarily related to host cell density. The absence of aggregated distributions of chytrids on their hosts suggested that their hosts did not vary in their susceptibility to infection. A parasite can only become epidemic when it grows faster than the host. Therefore, it has been suggested that epidemics in phytoplankton populations arise when growth conditions for the host are unfavorable. No support for such a generalization was found, however. Growth of the parasitic fungus Rhizophydium planktonicum Canter emend, parasitic on the diatom Asterionella formosa Hassal, was reduced under stringent nutrient limitation,because production and infectivity of zoospores were affected negatively. A moderate phosphorous or light limitation favored epidemic development, however. Chytrid infections have been shown to affect competition between their algal hosts and in this way altered phytoplankton succession. There is potential for coevolution between Asterionella and the chytrid Zygorhizidium planktonicum Canter based on clear reciprocal fitness costs, absence of overall infective parasite strains, and possibly a genetic basis for host susceptibility and parasite infectivity. [KEYWORDS: chytrids ; coevolution ; diatoms ; epidemics ; food webs ; hosts; parasites; phytoplankton; succession; zoospores]https://doi.org/10.1111/j.1529-8817.2004.03117.xDifferential sensitivity of green algae to allelopathic substances from Chara
Three short-term laboratory experiments were conducted to investigate allelopathic effects of a mixture of Chara globularis var. globularis Thuillier and Chara contraria var. contraria A. Braun ex Kützing on three different green algae. Single phytoplankton species were exposed to filtered water originating from charophyte cultures. Phytoplankton growth was monitored by determination of chlorophyll concentrations in batch cultures. The change in chlorophyll concentration during the experiments was analysed with a logistic growth model, resulting in an estimate of the exponential growth rate and the duration of the lag phase of the single green algae. The results indicate allelopathic effects of Chara on the growth of the green algae Selenastrum capricornutum Printz and Chlorella minutissima Fott et Nováková, whereas Scenedesmus obliquus (Turpin) Kützing did not seem to be affected. The exponential growth rate of S. capricornutum decreased 7% in the presence of water from a charophyte culture, while the growth rate of C. minutissima decreased with 3%. The allelopathic effect of Chara did not increase when the green alga C. minutissima was P-limited. The effect of Chara was different when young sprouts were used. With young sprouts the duration of the lag phase of C. minutissima was extended (25%), whilst for old plants the growth rate of this green alga decreased. Although the inhibiting effect of charophytes on specific phytoplankton species is rather small, the differential sensitivity of the species to Chara might influence the composition and biomass of phytoplankton communities in the field [KEYWORDS: allelopathy, charophytes, Chara globularis, Chara contraria, stoneworts, growth limitation]https://doi.org/10.1023/A:1024483704903The determination of ecological quality in shallow lakes - a tested system (ECOFRAME) for implementation of the European Water Framework Directive.
1. The European Water Framework Directive requires the determination of ecological status in European fresh and saline waters. This is to be through the establishment of a typology of surface water bodies, the determination of reference (high status) conditions in each element (ecotype) of the typology and of lower grades of status (good, moderate, poor and bad) for each ecotype. It then requires classification of the status of the water bodies and their restoration to at least good status in a specified period. 2. Though there are many methods for assessing water quality, none has the scope of that defined in the Directive. The provisions of the Directive require a wide range of variables to be measured and give only general guidance as to how systems of classification should be established. This raises issues of comparability across States and of the costs of making the determinations. 3. Using expert workshops and subsequent field testing, a practicable pan-European typology and classification system has been developed for shallow lakes, which can easily be extended to all lakes. It is parsimonious in its choice of determinands, but based on current limnological understanding and therefore as cost-effective as possible. 4. A core typology is described, which can be expanded easily in particular States to meet local conditions. The core includes 48 ecotypes across the entire European climate gradient and incorporates climate, lake area, geology of the catchment and conductivity. 5. The classification system is founded on a liberal interpretation of Annexes in the Directive and uses variables that are inexpensive to measure and ecologically relevant. The need for taxonomic expertise is minimized. 6. The scheme has been through eight iterations, two of which were tested in the field on tranches of 66 lakes. The final version, Version 8, is offered for operational testing and further refinement by statutory authorities [KEYWORDS: lakes Water Framework Directive typology ecotypes ecological status quality]https://doi.org/10.1002/aqc.592Colony formation in Scenedesmus: a literature overview and further steps towards the chemical characterisation of the Daphnia kairomone
Semiochemicals play an important role in interactions between living organisms in aquatic environments. Although the presence of chemical cues is confirmed in more and more systems, the chemical structures remain predominantly elusive. To create more accurate prey–predator interaction models and to advance the research on chemical communication, it is essential to identify these compounds. A literature overview of cues involving Daphnia (either as producer or receiver) is given and the progress towards their isolation and structure elucidation is described. Most of the research so far has concentrated on the elucidation of kairomones produced by predators of Daphnia (especially Chaoborus and several species of fish). Although some progress has been made, these cues have not been isolated and identified yet. Additionally new results on the isolation and identification of the kairomone responsible for the colony formation in Scenedesmus using differential diagnosis and bioassay-directed fractionation of Daphnia exudates are presented. The importance of suitable and well performing bioassays herein cannot be underestimated. Some preliminary results with solid-phase extraction with C18 proved to be reproducible for extracting the active compound from Daphnia water, although it was not possible to get the biological activity into a single fraction. The cue was not extractable with an anion exchanger (SAX). Subjecting the extract to HPLC led to one active fraction [KEYWORDS: semiochemicals, Scenedesmus, Daphnia, Solid-Phase Extraction, chemical communication, bioassay]https://doi.org/10.1023/A:1024414515222Molecular techniques in phytoplankton research: from allozyme electrophoresis to genomics
Molecular techniques have become a valuable tool in phytoplankton studies over the past decades. The appropriate choice of a technique from an increasing array of methods can be rather complex, because different techniques are suitable for different questions or problems in ecology and evolution. Each technique has its particular strengths and weaknesses and is based upon different (theoretical) assumptions. Our aim is to give a better insight in the (correct) use of various molecular techniques in phytoplankton research, with special emphasis on the fields of strain identification, differentiation of populations and the establishment of phylogenetic relationships. The basic steps in the development of molecular techniques like allozyme electrophoresis, RFLP, DGGE, SSCP, RAPD, AFLP and microsatellites, and the application of these techniques in phytoplankton research, are discussed. Furthermore, recent developments in molecular biology, that have so far only found limited application in phytoplankton studies, such as single-cell PCR, PCR assays combined with molecular probes (Heteroduplex Mobility Assays or DNA arrays), Real-time PCR, complete genome sequencing, multi-gene expression studies using microarrays, and Single Nucleotide Polymorphism (SNPs), are discussed. We emphasise the relevance of fundamental and applied molecular studies on phytoplankton for a wider community of ecologists and evolutionary biologists [KEYWORDS: algae, cyanobacteria, genetic variation, phylogenetics, AFLP, microsatellites, microarrays, genomics, review]https://doi.org/10.1023/A:1024473620792Response of Daphnia to substances released from crowded congeners and conspecifics
The effects of chemicals released from crowded congeners and conspecifics on life history parameters of the freshwater zooplankters Daphnia cucullata and Daphnia pulex were examined. Length and age at maturity of D. pulex were affected by crowding chemicals. Reproduction was lower in crowded medium, and ephippia were produced. Newborn D. pulex in crowded medium were significantly longer than the controls. The intrinsic rate of population increase of D. pulex was 14 and 25% lower than the control when exposed to crowded medium from D. cucullata and D. pulex, respectively. Neither urea nor ammonia (at 1 mg l-1) seemed to be responsible for these effects in D. pulex. In D. cucullata, no significant effect of crowding infochemicals on length and age at maturity was found. However, crowding chemicals reduced reproduction. No ephippia were produced in crowded medium, but up to 83% non-developing eggs were observed in D. cucullata. Newborns were similarly sized in crowded and standard medium. The intrinsic rate of population increase of D. cucullata was 44 and 96% lower than the control when exposed to crowded medium from D. cucullata and D. pulex, respectively. Clearance rates of D. pulex were significantly reduced in crowded media compared with standard medium, which could partly explain why the animals exposed to crowding chemicals reacted as if they were food limitedhttps://doi.org/10.1093/plankt/25.8.967Factors controlling hydrochemical and trophic state variables in 86 shallow lakes in Europe
In order to disentangle the causes of variations in water chemistry among European shallow lakes, we performed standardised sampling programs in 86 lakes along a latitudinal gradient from southern Spain to northern Sweden. Lakes with an area of 0.1 to 27 000 ha and mean depth of 0.4–5.6 m located in low to high altitudes were investigated within the EC project ECOFRAME 1–4 times during June–October 2000–2001. Several variables like conductivity, alkalinity, abundance of submerged plants, concentrations of suspended solids, total nitrogen and phosphorus were latitude-dependent decreasing from south to north. Secchi depth, concentrations of total nitrogen, total phosphorus, suspended solids, and chlorophyll a correlated strongly with the presumed quality classes of the lakes. We came to the conclusion that the variability of shallow lakes in Europe is still mostly dependent on natural differences. Variables connected to lake morphometry, seasonality, basin geology and climate explained altogether nearly half of the total variability of lakes. The trophic state factor, describing mostly the human influence on lakes, was the strongest single factor responsible for nearly a quarter of the total variability of the studied European lakes [KEYWORDS: ecological status, European shallow lakes, hydrochemistry, latitude]https://doi.org/10.1023/B:HYDR.0000008541.36537.51Influence of toxic and non-toxic phytoplankton on feeding and survival of Dreissena polymorpha (Pallas) larvae
Grazing and survival of larvae of the zebra mussel, Dreissena polymorpha, on a green alga and cyanobacteria were studied in laboratory experiments. Clearance rates of the larvae were determined for Chlamydomonas reinhardtii (green alga), two non-toxic and two toxic Microcystis aeruginosa strains (Cyanobacteria). Clearance rates of larvae on non-toxic Microcystis were significantly higher than on toxic Microcystis. The clearance rate on Chlamydomonas reinhardtii was in between the clearance rates on toxic and non-toxic Microcystis strains and not significantly different from them. Effects of toxicity of Microcystis on the survival of zebra mussel larvae was investigated in a short-term experiment. Survival of larvae fed toxic Microcystis was lower than that of larvae fed non-toxic Microcystis, but higher than that of starved larvae. This may imply that, for survival of zebra mussel larvae, it is better to have bad quality (toxic) food than no food [KEYWORDS: zebra mussel, phytoplankton, toxic cyanobacteria, survival, grazing, Microcystis]https://doi.org/10.1023/A:1024467708880Impact of submerged macrophytes including charophytes on phyto- and zooplankton communities: allelopathy versus other mechanisms
Submerged macrophytes are crucial for the stabilization of the clear water state in shallow, mesotrophic and eutrophic lakes. Especially, charophytes often play an important role because they are typically rapid colonizers and because charophyte meadows are believed to have a particularly strong positive effect on water transparency compared to other macrophytes. Several mechanisms may contribute to the impact of submerged macrophytes on the planktonic food web. In this paper, the available literature on these mechanisms is briefly reviewed and special attention is paid to the impact of charophytes on the structure and dynamics of phyto- and zooplankton communities. The paper focuses on allelopathy, and possible differences between charophytes and other macrophytes, as well as gaps in our knowledge are discussed. [KEYWORDS: Macrophyte; Charophyte; Allelopathy; Zooplankton and phytoplankton]https://doi.org/10.1016/S0304-3770(01)00205-4Lakes in the Netherlands, their origin, eutrophication and restoration: state-of-the-art review
This article starts with a brief description of the origin and eutrophication of shallow Dutch lakes, followed by a review of the various lake restoration techniques in use and the results obtained. Most freshwater lakes in the Netherlands are very shallow (https://doi.org/10.1023/A:1021092427559Short-term and long-term effects of zooplanktivorous fish removal in a shallow lake: a synthesis of 15 years of data from Lake Zwemlust
1. Removal of zooplanktivorous fish (mainly bream) in 1987 from a shallow eutrophic lake in the Netherlands, Lake Zwemlust, resulted in a quick switch from a turbid state with cyanobacteria blooms to a clear state dominated by macrophytes. 2. The clear state was not stable in the long term, however, because of high nutrient loadings. 3. In 1999, another removal of zooplanktivorous fish (mainly rudd) had similar effects as in 1987, although macrophytes returned more slowly. 4. In the years directly following both interventions there was a 'transition period' of very clear water with high densities of zooplanktonic grazers in the absence of macrophytes; low oxygen concentrations indicate that during those years primary production was low relative to heterotrophic activity. 5. The transition period appears to provide the light climate necessary for the return of macrophytes. 6. Reduction of nutrient loading is necessary to improve water quality in Lake Zwemlust in the long term. In the short term, repeated fish stock reduction is a reasonable management strategy to keep Lake Zwemlust clear.https://doi.org/10.1046/j.1365-2427.2002.01006.xComparing grazing by Dreissena polymorpha on phytoplankton in the presence of toxic and non-toxic cyanobacteria
The feeding behaviour of the zebra mussel (Dreissena polymorpha) was studied in the laboratory on different combinations of food, including a green alga Chlamydomonas reinhardtii) and toxic and non-toxic strains of the cyanobacterium Microcystis aeruginosa. 2. The highest clearance rate of phytoplankton by zebra mussels was found when the mussels were feeding on a mixture of Chlamydomonas and non-toxic Microcystis, the lowest on a mixture of Chlamydomonas and toxic Microcystis. 3. The differences found in the clearance rates between food combinations can be partly explained by the production of pseudofaeces containing live phytoplankton cells. Zebra mussels expelled significantly more live phytoplankton cells in the presence of toxic Microcystis than in the presence of non-toxic Microcystis. The pseudofaeces contained predominantly live Chlamydomonas cells. Proportionately much less live Microcystis cells were encountered in the pseudofaeces. 4. Consequently, grazing of zebra mussels on a combination of Chlamydomonas and Microcystis may finally result in a dominance of Chlamydomonas over Microcystis. The presence of toxic Microcystis may even strengthen this shift.https://doi.org/10.1046/j.1365-2427.2002.00933.xDaphnia food limitation in three hypereutrophic Dutch lakes: Evidence for exclusion of large-bodied species by interfering filaments of cyanobacteria
The Loosdrecht Lakes comprise three shallow, hypereutrophic lakes in The Netherlands. Research conducted over the past two decades suggests that absence of large-bodied Daphnia from these lakes can be explained by size-specific effects of both poor food quality and predatory mortality. The phytoplankton is dominated by filamentous cyanobacteria which interfere with the feeding of large-bodied Daphnia. Moreover, dense populations of planktivorous fish are expected to feed selectively on larger prey. More recent research, however, suggests that the growth of the small native Daphnia species, Daphnia cucullata, is limited by low dietary phosphorus (P). In this study, we compared the growth and feeding rates of four Daphnia taxa representing a gradient in body size. In each of 10 growth experiments. native D. cucullata feeding in natural seston exhibited improved growth in response to additions of phosphate and the P-rich cyanobacterium Syncehococcus. The two largest species, Daphnia magna and Daphnia galeata, however, failed to grow in natural seston, even when the seston was supplemented with phosphate, Synechococcus, or Scenedesmus A D. galeata X cucullata hybrid exhibited weak growth in natural seston and no response to the supplements. Feeding experiments with polystyrene beads show that feeding inhibition in natural seston increased markedly with increased Daphnia size. Our results verify that Daphnia differing in size face different food quality constraints. The growth of native D. cucullata (adult size 0.6-0.8 mm) is reduced by P deficiency, whereas larger Daphnia appear to be excluded by interfering filaments of cyanobacteria. [KEYWORDS: Blue-green filaments; loosdrecht lakes eutrophic lake zooplankton; netherlands; selection; consumers; carbon]https://doi.org/10.4319/lo.2001.46.8.2054Differential sensitivity to natural ultraviolet radiation among phytoplankton species in Arctic lakes (Spitsbergen, Norway)
Incubation experiments demonstrated a differential sensitivity to natural UV-radiation among the dominant phytoplankton species from three Arctic lakes, situated near Ny-Angstrom lesund, Spitsbergen (79 degrees N). The growth of small chlorophytes, diatoms and picocyanobacteria from two oligotrophic lakes was inhibited primarily by the shorter wavelength UV components, while the growth of the larger colony-forming species (cyanobacteria, Planktothrix sp., Woronichinia sp. and the chrysophyte, Uroglena americana Calkins) apparently was stimulated. These colonies (not easily eaten by daphnids) dominated at the end of the experiment in those treatments where the short wavelength UV components were not excluded. For the two oligotrophic localities, 70 and 61%, respectively, of total phytoplankton biovolume were edible in the treatments excluding short wavelength UV, compared to only 13 and 19%, respectively, in the treatments including such radiation. For the third, more productive and less transparent lake, the percentage of edible species in the treatments with and without short wavelength UV radiation did not differ (ca. 75% for both treatments). [KEYWORDS: cell size; cyanobacteria; daphnids; foodweb; grazers; inedible algae; Svalbard; UV-B Uv-b radiation; ozone depletion; chlamydomonas-reinhardtii; community responses; solar uvb; inhibition; growth; assemblages; algae; zooplankton]https://doi.org/10.1023/A:1012978328768Effects of dietary phosphorus deficiency on the abundance, phosphorus balance, and growth of Daphnia cucullata in three hypereutrophic Dutch lakes
We studied population dynamics, phosphorus balance, and individual growth to evaluate limitation of Daphnia cucullata by dietary phosphorus (P) in three interconnected, hypereutrophic Dutch lakes. Consistent with the P-limitation hypothesis, declines in egg production and population density coincided with seasonal increases in the seston C:P ratio from 300 to 400-500 (molar) in all three lakes. Adult Daphnia collected from Lake Breukeleveen exhibited a linear increase in specific P content during the first 36 h of feeding on a P-rich diet in the laboratory (r(2) = 0.99), which confirms that field Daphnia experienced P deficits in their tissues. Laboratory experiments tested the growth of juvenile D. cucullata in controls with natural seston and in seston supplemented with phosphate or the P-rich cyanobacterium Synechococcus. Growth was also estimated in filtered lake water with Scenedesmus (2 mg C L-1) as food. This design allowed us to partition differences between the control and Scenedesmus growth rates into limitation by phosphorus, energy and residual factors. In each of 10 experiments, D. cucullata exhibited improved growth in response to the phosphate and Synechococcus supplements. In agreement with predictions, Daphnia exhibited the lowest control growth rate and the strongest response to P addition in seston from the lake with the highest seston C:P ratio. However, animals in seston from two lakes with very similar seston C:P ratios differed in their growth responses. Our results confirm dietary P limitation but show that energy limitation and residual factors also contributed to between- lake differences in Daphnia growth. [KEYWORDS: Unsaturated fatty-acids; food quality; mineral limitation green-algae; zooplankton; carbon; cyanobacterium; phytoplankton; nitrogen; nutrition]https://doi.org/10.4319/lo.2001.46.8.1871Feeding in Daphnia galeata on Oscillatoria limnetica and on detritus derived from it
Oscillatoria spp. are generally very abundant in many shallow, eutrophic lakes in the Netherlands. However, this is less true for Daphnia galeata. The main aim of this study was to investigate whether the edibility of live Oscillatoria limnetica by Daphnia galeata, and of the detritus derived from this filamentous cyanobacterium, was, among others, an important limiting factor for the daphnids. We measured the consumption and assimilation rates of Daphnia using dual- labelling radio-tracer techniques (C-14 and P-32) to label separately the live Oscillatoria filaments and detritus prepared from these filaments. The two food types were mixed in different proportions, and both the food ingestion and food incorporation rates by daphnids were measured. The main findings of this study were that specific clearance rates of Daphnia on shorter Oscillatoria filaments were significantly higher than on the longer filaments, in other words the weight- specific ingestion rates were higher on the shorter Oscillatoria filaments than on the longer filaments from the batch cultures, The longer Oscillatoria filaments are more like v to clump and, therefore, are more liable to be rejected by Daphnia during the food collection and ingestion processes. The shorter filaments, in comparison, are apparently less prone to clumping and, therefore, are cleared by the daphnids at higher rates than the longer filaments. Feeding the daphnids on double-labelled (C-14 and P-32) shorter filaments revealed that the assimilation efficiency of shorter Oscillatoria filaments was generally higher for P than for C, probably because of a high CIP ratio of Oscillatoria. Daphnia (0.75-1.85 mm in size)fed significantly better on dead (detritus) Oscillatoria filaments than on live Oscillatoria filaments, even if the relative proportion of detritus in the food was only about one- quarter that of the live filaments. This preference for detritus over live Oscillatoria, as indicated by Chesson's selectivity coefficient a, was apparently a passive process, rather than a case of active food selection. This 'selective' feeding on detritus was apparently facilitated by the relatively greater rejection of the live filaments than the detrital filaments. At very high food concentrations (15-25 mg Cl-1), with the share of the live Oscillatoria filaments 2 to 2.5 times greater than that of the detritus, the ingestion rates increased proportionally less with increase in daphnid size than was expected on the basis of the allometric relationship between the length and weight of these animals. This implies that the larger animals had greater interfering effects of the Oscillatoria filaments on the food collection and ingestion processes. [KEYWORDS: Zooplankton-cyanobacteria interactions; lake lake loosdrecht; eutrophic lake; filamentous cyanobacteria; fresh-water; long-term; algal; discrimination; cladocerans; food]https://doi.org/10.1093/plankt/23.7.705Sequence analysis of the its-2 region: a tool to identify strains of Scenedesmus (Chlorophyceae)
The genetic distances between several strains of Senedesmus obliquus (Turp,) Kutz,, S, acutus Hortobagyi, and S, naegelii Chod. calculated from ITS-2 sequences were found to be smaller than the genetic distances within other strains of Scenedesmus-that is, in S, acuminatus (Lagerh,) Chod, and S, pectinatus Meyen. These results confirm that the studied strains were not properly identified and should be renamed S, obliquus, as already suggested in other studies. [KEYWORDS: ITS, phylogeny, Scenedesmus acutus, Scenedesmus naegelii, Scenedesmus obliquus]https://doi.org/10.1046/j.1529-8817.2000.99190.xImpact of the fungicide carbendazim in freshwater microcosms. II. Zooplankton, primary producers and final conclusions
Effects of chronic application of the fungicide Derosal(R) (active ingredient carbendazim) were studied in indoor macrophyte-dominated freshwater microcosms. The concentrations (0, 3.3, 33, 100, 330 and 1000 mu g/l) were kept at a constant level for 4 weeks. This paper is the second of a series of two; it describes the effects on zooplankton and primary producers and presents an overall discussion. The zooplankton community was negatively affected by the three highest treatment levels (NOECcommunity = 33 mu g/l). At higher treatment levels Cladocera taxa were completely eliminated, while Copepod numbers were reduced. Rotatoria taxa decreased (Keratella quadrata and Lecane sp.) or increased in abundance (Testudinella parva) at the highest treatment level only. Due to the reduced grazing pressure, the abundance of some phytoplankton taxa and the chlorophyll-a content of the phytoplankton increased at the three highest treatment levels (NOECcommunity = 33 mu g/l). This effect was not observed for the periphyton, most probably because the reduced grazing pressure was compensated by the increased abundance of some snail species such as Lymnaea stagnalis and Physella acuta. At the end of the experimental period the biomass of the macrophyte Elodea nuttallii was significantly elevated at the two highest treatment levels. It is hypothesised that carbendazim might have caused, directly or indirectly, the removal of pathogene organisms from the macrophyte. [KEYWORDS: carbendazim; aquatic microcosm; pesticides; ecosystem effects; ecological effect chain]https://doi.org/10.1016/S0166-445X(99)00037-5Grazer-induced colony formation in Scenedesmus: are there costs to being colonial?
Grazer-induced colony formation in the common green alga Scenedesmus acutus may be interpreted as an anti-grazer defense. Costs are to be expected because otherwise the protected colonial morph would be the norm. Analysis of growth rates, light harvesting in terms of photosystem II (PSII) efficiency using Xe-PAM fluorescence measurements and sedimentation rates were compared among unicellular and induced colonial populations. No differences in growth rates and PSII efficiency were detected between unicellular and colonial populations. However, colonial populations had higher settling velocities than unicellular populations. Hence, costs may be attributed to enhanced sinking out of the euphotic zone resulting in reduced growth due to lower light and temperature in deeper water layers. [KEYWORDS: Morphological-changes; inducible defenses; crucian carp; green- algae; in-situ; phytoplankton; daphnia; chlorophyceae; sinking; growth]https://doi.org/10.1034/j.1600-0706.2000.880113.xField research for the authorisation of pesticides
On request of the Dutch government a committee of the Health Council of the Netherlands has reviewed the role that results of field research in its broadest sense (i.e., including multi- species toxicity tests in the laboratory, research on model ecosystems et cetera) can play in ecotoxicological risk assessment for the authorisation of pesticides. The Committee believes that field research can provide valuable additional data about the exposure of non-target organisms and the resultant effects at population, community and ecosystem level. However, it frequently is unclear how these data might be used in reaching a decision about authorisation. To solve this problem, it is necessary to specify what is understood by "unacceptable damage". Both more clearly formulated protection goals of the government and a better understanding of the ecological significance of effects are needed to clarify this. Furthermore, the Committee points out that the statistical power of field trials must be sufficient to allow for the detection of changes that might be regarded as ecologically relevant. Finally, it recommends keeping a finger on the pulse in relation to authorised pesticides by monitoring their presence in environmental compartments and by investigating their role in suddenly occurring mortality among conspicuous animal species, such as birds, fish and honeybees. This kind of research forms a safety net for substances that have been wrongly authorised. [KEYWORDS: authorisation; field research; model ecosystems; monitoring; pesticides]https://doi.org/10.1023/A:1008920401180Grazer-induced colony formation in Scenedesmus acutus (Chlorophyceae): Ecomorph expression at different temperatures
Scenedesmus acutus Meyen was cultured at four temperatures (9.5 degrees, 16.5 degrees, 24 degrees, and 29 degrees C) in standard medium or in medium with filtered water from a Daphnia culture, Growth was significantly reduced at low temperatures. At 9.5 degrees C it took more than a week before formation of eight-celled coenobia occurred in both the absence and presence of water from a Daphnia culture, At higher temperatures, formation of four- and eight-celled coenobia occurred more rapidly and was already observed in the presence of Daphnia water within 2 days, As cultures aged, also in the absence of Daphnia water, four-celled coenobia became dominant. At cold temperature, cell volume initially was significantly larger but declined after 3-4 weeks. Grazer-induced colony formation had occurred independent of the incubation temperature, but the number of cells per colony was increased with declining temperature. The morphological expression may be interpreted as a cyclomorphosis driven by nutrients, temperature, and chemical cues from gazers. [KEYWORDS: colony formation; cyclomorphosis; Daphnia; ecomorph; growth; infochemicals; phenotypic plasticity; Scenedesmus acutus; temperature Phenotypic plasticity; morphological-changes; daphnia-pulex; phytoplankton; growth; cyclomorphosis; light; transformation; infochemicals; consequences]https://doi.org/10.1046/j.1529-8817.1999.3561120.xEffects of artificial ultraviolet-B radiation on experimental aquatic microcosms
1. The effects of prolonged ultraviolet-B (UVB) radiation on freshwater communities were studied in indoor microcosms (600 L) with artificial light sources, simulating a clear, shallow, mesotrophic aquatic ecosystem. A range of six intensities (in duplicate) of UVB radiation, ranging from 0 (control) to 9.56 kJ m(-2) day(-1) at the water surface, was applied for 8 weeks. The UVB radiation levels, attenuation, shading and scattering were comparable to those in Dutch shallow freshwater systems. Physical, chemical and biological variables were monitored weekly. 2. The WE treatment did not affect the abundance, species composition or biovolume of the phytoplankton or zooplankton communities, nor did it affect the periphyton or the macroinvertebrate community. A few species showed a significant response on some of the sampling dates, but there was no negative UVB effect at the community level. Overall, the ecosystems in the microcosms were not affected by the UVB treatment. 3. In a bio-assay, a laboratory clone of Daphnia pulex, not subjected to UVB radiation, was fed with seston from the microcosms. Daphnia pulex feeding on seston from the control microcosms grew faster, had better survival and better reproduction than D. pulex feeding on seston from the UVB treated microcosms. The phytoplankton-zooplankton interaction may have been influenced by the UVB treatment. 4. The dissolved oxygen content (DOC) concentrations in the microcosms were around 5 mg L-1. The DOC levels in Dutch systems rarely fall below 10 mg L-1. This might provide sufficient protection against the detrimental effects of increased UVB radiation. [KEYWORDS: ecosystem, macroinvertebrates, macrophytes, microcosm, phytoplankton, UVB, zooplankton]https://doi.org/10.1046/j.1365-2427.1999.00478.xA model study on the stability of the macrophyte-dominated state as affected by biological factors
The transition of shallow lake ecosystems between the clear-water, macrophyte-dominated state and the turbid state dominated by phytoplankton depends on both physico-chemical and biological factors. In this study, the impact of some of these interactions on the stability of the macrophyte-dominated state of a lake are studied by means of the integrated eutrophication model PCLake. The model describes phytoplankton, macrophytes and a simplified food web, within the framework of closed nutrient cycles. The aim of the study is to evaluate the impact of herbivory by birds and fish on the transition from clear to turbid state, including the influence of variability in other biological parameters. The model was applied to the data of a small, biomanipulated lake, dominated by macrophytes, showing signs of a transition back to the turbid state. Simulations were carried out for the lake as well as for an experimental situation where herbivory was impeded. A parameter variation study was performed for 10 parameters, affecting the zooplankton, fish and macrophytes behaviour, to determine the sensitivities and the model uncertainty. The model reproduced adequately the transition of the lake from phytoplankton dominance before the biomanipulation, via dominance of rooted perennial plants in the first years after it, to a state characterized by turion-forming plants in early summer and phytoplankton in autumn. It is shown that the probability of the transition back to phytoplankton dominance is mainly enhanced by herbivory by birds. This caused a shift towards inedible plant species with a shorter natural growing season, allowing the return of a phytoplankton bloom in autumn. If herbivory was impeded, this shift did not occur and phytoplankton remained low due to nitrogen limitation. The model results were quite sensitive to the zooplankton filtering rate and, in the presence of herbivory only, to the macrophytes growth parameters. The impact of the fish parameters showed to be less important. The model may be used to evaluate the relative importance of different assumptions or factors in the success of biomanipulation measures in lakes. [KEYWORDS: biomanipulation, herbivory, hysteresis, lake ecosystem, Lake Zwemlust, macrophytes, model, phytoplankton, sensitivity analysis, uncertainty analysis]https://doi.org/10.1016/S0043-1354(98)00049-9Altered cell wall morphology in nutrient-deficient phytoplankton and its impact on grazers
Grazing experiments were performed with the zooplankters Daphnia pulex and Daphnia magna feeding on nitrogen- and phosphorus-limited cells of two green algae (Chlamydomonas reinhardtii and Selenastrum capricornutum). To analyze the role of the cell wall structure in digestibility of the algae by Daphnia, the same experiments were carried out with both wild-type C. reinhardtii and a cell wall-deficient mutant. The nonlimited algae were efficiently assimilated, whereas P- and N-limited algal cells were not. Especially P-limited cells passed mostly intact and viable through the gut and were thus spared from heavy grazing pressure. In life-history experiments, D. pulex grazing on nonlimited algae reached the largest body size, whereas animals fed N- or P-limited algae exhibited reduced growth. Cells of the wall-deficient mutant of Chlamydomonas, grown under both nutrient-limited and nonlimited conditions [KEYWORDS: FRESH-WATER ZOOPLANKTON, FOOD QUALITY, DAPHNIA-MAGNA, NITROGEN-CONTENT, GROWTH-RATE, LIMITATION, DISCRIMINATION, SELECTION, NUTRITION, COPEPOD]https://doi.org/10.4319/lo.1997.42.2.0357Changes in food quality of the green alga Scenedesmus induced by Daphnia infochemicals: biochemical composition and morphology
1. The effects of Daphnia infochemicals on the morphology and biochemical composition of Scenedesmus were studied and subsequently their influence on Daphnia life history. Three species of Scenedesmus (S. acutus, S. obliquus and S. subspicatus) were tested for Daphnia-induced colony formation. Life history experiments were performed with Daphnia cucullata and D. magna feeding on unicellular or colonial S. acutus. 2. Colony formation was promoted when S. acutus and S. obliquus were exposed to filtered water from a D. magna culture. S. subspicatus did not form colonies when exposed to culture water, but showed some colony formation when exposed to live D. magna. 3. No clear differences were found in total lipid, protein or carbohydrate content between the unicellular and colonial ecomorphs of the three Scenedesmus species. However, fatty acid (FA) concentration and composition were changed. Total FA as a percentage of frozen dry weight (DW) were increased in colonies. The ratio of monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA) and the ratio of omega 3:omega 6 fatty acids decreased in colonies. 4. A short-term grazing experiment with three size classes of Scenedesmus revealed lower clearance rates for small Daphnia when feeding on large colonial Scenedesmus. 5. Life history parameters of both Daphnia species were influenced by food type. The intrinsic rate of population increase (r) was significantly lower for daphnids feeding on colonies. 6. The negative influence of colonial Scenedesmus on population growth of Daphnia seems to be determined by algal morphology rather than biochemical composition. [KEYWORDS: Life-history shifts; mineral limitation; herbivore daphnia; flos-aquae; growth; zooplankton; phytoplankton; cladoceran; inhibition; magna]https://doi.org/10.1046/j.1365-2427.1997.00225.xEffects of UV-B irradiated algae on life history traits of Daphnia pulex
1. The impact of ultraviolet-B (UVB)-irradiated phytoplankton on the life history parameters of Daphnia was studied. Three species of Chlorophyceae (Chlamydomonas reinhardtii, Scenedesmus acutus and S. subspicatus) and one species of Cryptophyceae (Cryptamonas pyrenoidifera) were cultured with and without exposure to UVB radiation (biologically effective dose of 3 kJ m(-2) day(-1) DNA weighted) and fed to Daphnia pulex. 2. The daphnids feeding on UVB-irradiated algae tended to be smaller in size, to produce less and smaller offspring, and to have a reduced population growth rate. 3. Fatty acid (FA) analysis of the phytoplankton showed a reduced total FA content under UVB radiation for all four species. Changes in the content of unsaturated fatty acids (UFA) were species specific. 4. The results indicate that the energy transfer between the primary producers' and grazers' level might be negatively influenced by UVB radiation. Direct effects on food quality are modest but detectable, changes in quantity or species composition may be more important in the energy transfer. [KEYWORDS: ULTRAVIOLET-B RADIATION, FATTY-ACID COMPOSITION, OZONE DEPLETION, CHLAMYDOMONAS-REINHARDTII, HERBIVORE DAPHNIA, FOOD QUALITY, PHYTOPLANKTON, GROWTH, COMMUNITY, RESPONSES]https://doi.org/10.1046/j.1365-2427.1997.00221.xDefences in phytoplankton against grazing induced by nutrient limitation, UV-B stress and infochemicals
It is becoming increasingly evident that the efficiency of zooplankton grazing on algae is not only a matter of quantity of the grazer relative to its food. Planktonic primary producers are not defenseless food-particles that are easily harvested by the consumers. Several algal species are able to adjust their phenotype (colony formation, spines, size) in such a way that it results in a reduced grazing pressure. It was recently demonstrated that morphological changes in the cell wall of green algae, induced by nutrient limitation and UV-B stress, may reduce their digestibility. A high fraction of induced cells pass intact and viable through the gut of the zooplankters, such that the grazing impact on the population is strongly reduced. It was also found that the presence of exudates (infochemicals) released by daphnids may change the morphology of algae. Unicellular green algae of the genus Scenedesmus were induced to form eight-cell coenobial types, heavily armed with spines, within three to five days after adding filtered water from an algal culture with Daphnia present. Both defence mechanisms may play an important role in zooplankton production and competition, and may serve as an example of highly efficient strategies to resist heavy grazing pressure. [KEYWORDS: algal cell wall, colony formation, Daphnia, digestibility, grazing resistance, induced defence]https://doi.org/10.1023/A:1009951622185Life history consequences for Daphnia pulex feeding on nutrient-limited phytoplankton
1. The growth and feeding of Daphnia pulex De Geer on different algal species was examined. The green algae Chlamydomonas reinhardtii Dangeard and Scenedesmus acutus Meyen, the diatom Synedra tenuissima Kutzing, the cryptophyte Cryptomonas pyrenoidifera Geitler and the cyanobacterium Microcystis aeruginosa Kutzing were cultured in nonlimiting and in N- or P-limiting medium and used as food for D. pulex. 2. Growth limitations were reflected in the elemental and biochemical composition and the morphological characteristics of the algal resources. 3. The clearance rates of D. pulex feeding on nutrient-limited algae were reduced. This was not observed when nutrient-limited mutant Chlamydomonas cells without cell walls were used as food, indicating that the cell wall may play an important part. 4. In comparison with animals grown on nutrient-sufficient cells, nutrient-limited algae resulted in smaller body length, reduced brood sizes, reduced size at maturity, increased age at first reproduction and, consequently, in reduced Daphnia population growth rates. 5. Daphnia population growth rates (r) were negatively correlated with the C:P ratio and the carbohydrate content of the food. Moreover, significant correlations between r and clearance rates were found. 6. The observed differences in the grazing and the life history parameters of Daphnia feeding on non-limited and nutrient-limited algae may be the result of both reduced nutritional value and reduced digestibility of nutrient-limited algae. [KEYWORDS: FRESH-WATER ZOOPLANKTON, FOOD QUALITY, CHEMICAL-COMPOSITION, MINERAL LIMITATION, VARYING QUALITY, GROWTH-RATE, DISCRIMINATION, MAGNA, ALGAE, SCENEDESMUS]https://doi.org/10.1046/j.1365-2427.1997.00242.xMorphological changes in Scenedesmus induced by infochemicals released in situ from zooplankton grazers
Biochemical substances released from Daphnia galeata induced colony formation in the green alga Scenedesmus acutus. Normally this strain consisted mainly of single cells in cultures. However, if exposed for 48 h to either water with live Daphnia or to O.1-mu m filtered water from a culture with Daphnia present, these unicellular "Chodatella" stages were induced to form colonies (coenobia). Colony induction was not unique to Daphnia; other zooplankters (rotifers and copepods) were able to induce colonies in S. acutus as well. This morphological response could also be evoked when Scenedesmus was exposed to 0.1-mu m filtered lake water during high zooplankton abundances. Especially during early spring, a clear relationship was found between rotifer abundance and colony formation in our test alga in the laboratory. Filtered lake water, incubated nonaxenically for at least 2 d at 20 degrees C, did not induce colony formation, possibly due to microbial degradation. The morphological changes in Scenedesmus could promote grazing resistance in small zooplankters and can be interpreted as an adaptive antipredator strategy. [KEYWORDS: DIEL VERTICAL MIGRATION; BLUE-GREEN-ALGAE; DAPHNIA; INDUCTION; CHLOROPHYCEAE; DEFENSES; FISH]UV-induced changes in phytoplankton cells and its effects on grazers
This review addresses the effects of UV-radiation on the morphology and biochemistry of phytoplankton and the potential effects on grazers. UVA and UVB radiation inhibit the uptake of inorganic nutrients in phytoplankton. Reduced rates of ammonium and nitrate uptake in marine diatoms, and reduced uptake of phosphorus in freshwater flagellates are reported. The effects on cell stoichiometry are not settled. UVA and UVB radiation promote increased cell volumes owing to a decoupling between the photosynthetic processes and cell division. Loss or inactivation of flagellae and loss of motility are also reported for a number of phytoplankton species. UVA and UVB radiation may affect major biochemical constituents. Accumulation of intracellular, photosynthetic products (lipids or carbohydrates) is a common, although not unique, property of UV-stressed algae. Fatty acid (FA) profiles seem susceptible to UV radiation. A relative increase of short-chained, and a decrease in polyunsaturated FA (PUFA) are reported. The important membrane FA like eicosapentaenoic acid (EPA, 20:5 omega 3) and docosahexaenoic acid (DHA, 22:6 omega 3) seem particularl UV-related responses are highly dependent on taxonomy, cell-cycle stage, nutrient-limitation and the UV:PAR (photosynthetic active radiation)-ratio. Nutrient deficiency, cell size, cell wall properties and FA can all have significant impacts on grazers. Thus the reported effects on cell morphology and biochemical constituents could have profound effects on grazers and energy transfer in aquatic foodwebs. [KEYWORDS: ULTRAVIOLET-B RADIATION, FATTY-ACID COMPOSITION, EUGLENA-GRACILIS, OZONE DEPLETION, CHLAMYDOMONAS-REINHARDTII, MONOCHROMATIC LIGHT, NITROGEN STARVATION, NUTRIENT LIMITATION, N-15-NITRATE UPTAKE, ISOCHRYSIS-GALBANA]https://doi.org/10.1046/j.1365-2427.1997.00223.xA multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake
1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986-94. 2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992-94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures. 3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface :volume ratio and rapidly reproducing algae. However, in mid-summer of 1992-94, nutrient availability and cladoceran grazing on edible algae favoured cyanophyte. 4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore-macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore- dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community. 5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl. [KEYWORDS: Top-down; environmental-factors; community structure;hypertrophic lake; trophic levels; bottom-up; stability; fish; manipulation; restoration]https://doi.org/10.1046/j.1365-2427.1996.d01-511.xTransition of a lake to turbid state six years after biomanipulation: Mechanisms and pathways
Six years after application of biomanipulation in 1987, Lake Zwemlust (The Netherlands) returned during the summer from a clear water state dominated by aquatic vegetation to a turbid state characterized by high algal biomass. Herbivory and growth of epiphytes on macrophytes were the main factors triggering the switch. Selective herbivory by coots (Fulica atra) and rudd (Scardinius erythrophthalmus) caused a change in macrophyte species composition from a dominance of Elodea nuttallii in 1988/1989 to Ceratophyllum demersum in 1990/1991, and finally to Potamogeton berchtholdii in 1992/1993/1994. Observations revealed a general lack of epiphytes associated with Elodea and Ceratophyllum, while Potamogeton showed a progressive coverage with epiphytes, causing Potamogeton to decline markedly during late summer. Phytoplankton blooms, dominated by cyanobacteria, appeared again during three consecutive autumns, 1992, 1993 and 1994, with chlorophyll-a concentrations reaching 60-240 mu g l(-1). [KEYWORDS: biomanipulation; coots; epiphytes; herbivory; Lake Zwemlust; macrophytes Submerged plants; aquatic plant; stable states; shallow lakes; water; macrophytes; eutrophication; phytoplankton; ecosystem;community]https://doi.org/10.1016/0273-1223(95)00699-0
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Sublethal Effects of Pharmaceuticals on Aquatic Food Web Functioning (Infodisrupt)
Pharmaceuticals are therapeutic agents contaminating aquatic systems and hence included in the Water Framework Directive’s watch list of Contaminants of Emerging Concern. High global consumption of pharmaceuticals has led to their increased occurrence in aquatic systems. As they are designed to be therapeutically active at ng/L to μg/L concentrations, pharmaceuticals can affect non-target organisms in aquatic system exhibiting sub-lethal effects. Sub-lethal effects caused by pharmaceuticals can be direct when there are similarities in the mode of action. Indirect effects triggered by pharmaceuticals can be density-mediated and/or trait-mediated including the once caused by the disruption and/or mimicking of infochemicals involved in chemical communication between organisms.
In this project, we explore the sub-lethal effects of pharmaceuticals on aquatic food web functioning. Firstly, by understanding the impact of sub-lethal concentrations of pharmaceuticals on trophic interactions. Secondly, we assess the significance of these sub-lethal effects on aquatic food web functioning. Therapeutic drugs with high global occurrence, environmental relevance and persistence are used for all the experiments. The observations and findings of this project will inform the water managers about the significance of considering the sub-lethal effects of environmental concentrations of pharmaceuticals on aquatic ecosystems while determining their environmental risk limits.
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