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Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. Effects of these food quality indicators vary across studies, and the relative importance of elemental (nitrogen and phosphorus) versus biochemical (fatty acid and sterol) limitation in aquatic food webs has been debated. Here we present results of a meta-analysis using >100 experimental studies, which confirms that limitation by N, P, essential fatty acids, and sterols all have significant negative effects on zooplankton performance. However, effects varied by grazer response (growth versus reproduction), specific manipulation, and across taxa. P limitation had greater effects on zooplankton growth than fatty acids, but P and fatty acid limitation had equal effects on reproduction. Furthermore, we show that nutrient co-limitation in zooplankton occurs, that indirect effects induced by P limitation exceed direct effects of mineral P limitation, that effects of nutrient amendments using laboratory phytoplankton isolates exceed those using natural field communities, and that algal physiology mediates zooplankton responses to nutrient limitation. Our meta-analysis reconciles contrasting views about the role of various food quality indicators, and their interactions, for zooplankton performance, and provides a mechanistic understanding of how environmental change affects trophic transfer.

Underwater light is spatially as well as temporally variable and directly affects phytoplankton growth and competition. Here we systematically (following the guidelines of PRISMA-EcoEvo) searched and screened the published literature resulting in 640 individual articles. We mapped the conducted research for the objectives of (1) phytoplankton fundamental responses to light, (2) effects of light on the competition between phytoplankton species, and (3) effects of climate-change-induced changes in the light availability in aquatic ecosystems. Among the fundamental responses of phytoplankton to light, the effects of light intensity (quantity, as measure of total photon or energy flux) were investigated in most identified studies. The effects of the light spectrum (quality) that via species-specific light absorbance result in direct consequences on species competition emerged more recently. Complexity in competition arises due to variability and fluctuations in light which effects are sparsely investigated on community level. Predictions regarding future climate change scenarios included changes in in stratification and mixing, lake and coastal ocean darkening, UV radiation, ice melting as well as light pollution which affect the underwater light-climate. Generalization of consequences is difficult due to a high variability, interactions of consequences as well as a lack in sustained timeseries and holistic approaches. Nevertheless, our systematic literature map, and the identified articles within, provide a comprehensive overview and shall guide prospective research.

Human-induced salinization caused by the use of road deicing salts, agricultural practices, mining operations, and climate change is a major threat to the biodiversity and functioning of freshwater ecosystems. Yet, it is unclear if freshwater ecosystems are protected from salinization by current water quality guidelines. Leveraging an experimental network of land-based and in-lake mesocosms across North America and Europe, we tested how salinization—indicated as elevated chloride (Cl−) concentration—will affect lake food webs and if two of the lowest Cl− thresholds found globally are sufficient to protect these food webs. Our results indicated that salinization will cause substantial zooplankton mortality at the lowest Cl− thresholds established in Canada (120 mg Cl−/L) and the United States (230 mg Cl−/L) and throughout Europe where Cl− thresholds are generally higher. For instance, at 73% of our study sites, Cl− concentrations that caused a ≥50% reduction in cladoceran abundance were at or below Cl− thresholds in Canada, in the United States, and throughout Europe. Similar trends occurred for copepod and rotifer zooplankton. The loss of zooplankton triggered a cascading effect causing an increase in phytoplankton biomass at 47% of study sites. Such changes in lake food webs could alter nutrient cycling and water clarity and trigger declines in fish production. Current Cl− thresholds across North America and Europe clearly do not adequately protect lake food webs. Water quality guidelines should be developed where they do not exist, and there is an urgent need to reassess existing guidelines to protect lake ecosystems from human-induced salinization. This article contains supporting information online at https://www.pnas.org/lookup/suppl/doi:10.1073/pnas.2115033119/-/DCSupplemental

1: Body size is a decisive functional trait in many organisms, especially for phytoplankton, which span several orders of magnitude in cell volume. Therefore, the analysis of size as a functional trait driving species’ performance has received wide attention in aquatic ecology, amended in recent decades by studies documenting changes in phytoplankton size in response to abiotic or biotic factors in the environment. 2. We performed a systematic literature review to provide an overarching, partially quantitative synthesis of cell size as a driver and sentinel of phytoplankton ecology. We found consistent and significant allometric relationships between cell sizes and the functional performance of phytoplankton species (cellular rates of carbon fixation, respiration, and exudation as well as resource affinities, uptake, and content). Size-scaling became weaker, absent or even negative when addressing C- or volume-specific rates or growth. C-specific photosynthesis and population growth rate peaked at intermediate cell sizes around 100 µm³. 4: Additionally, we found a rich literature on sizes changing in response to warming, nutrients and pollutants. Whereas small cells tended to dominate under oligotrophic and warm conditions, there are a few notable exceptions, which indicates that other environmental or biotic constraints alter this general trend. Grazing seems a likely explanation, which we reviewed to understand both how size affects edibility and how size structure changes in response to grazing. Cell size also predisposes the strength and outcome of competitive interactions between algal species. Finally, we address size in a community context, where size-abundance scaling describes community composition and thereby the biodiversity in phytoplankton assemblages. 5: We conclude that i) size is a highly predictive trait for phytoplankton metabolism at the cellular scale, with less strong and non-linear implications for growth and specific metabolism, and ii) size structure is a highly suitable sentinel of phytoplankton responses to changing environments.

Light is a fundamental resource for phytoplankton. To utilize the available light, most phytoplankton species possess pigments in taxon‐specific combinations and quantities, which in turn result in a specific use of certain wavelengths. This optimizes the light use efficiency, allows for a complementary use of light, and may be an additional driver for community structure. While the effects of light intensity on phytoplankton biomass production and community composition have been intensively studied, here we focused on the effects of specific light spectrum quality (thus light color) on a natural phytoplankton community. In a controlled mesocosm experiment we reduced the supplied wavelength range to its blue, green, or red part of the light spectrum and compared the responses of each treatment to a full spectrum control over 28 d. Highest community growth rates were observed under blue, lowest under red light. Light absorption by the communities showed adaptation toward the supplied wavelength range. Community composition was significantly affected by light quality treatments, driven by Bacillariophyta and Chlorophyta, whereas pigment composition was not. Furthermore, lower species richness but higher evenness occurred when communities were exposed to red light compared to the full spectrum. We expected the response of phytoplankton communities to changes in the light spectrum to be driven by a combination of species sorting and pigment acclimation; however, the effect of species sorting turned out to be stronger. Our study showed that, even if species might acclimate, changes in the available light spectrum affect primary production and phytoplankton community composition.

Increasing human impact on the environment is causing drastic changes in disturbance regimes and how they prevail over time. Of increasing relevance is to further our understanding on biological responses to pulse disturbances (short duration) and how they interact with other ongoing press disturbances (constantly present). Because the temporal and spatial contexts of single experiments often limit our ability to generalize results across space and time, we conducted a modularized mesocosm experiment replicated in space (five lakes along a latitudinal gradient in Scandinavia) and time (two seasons, spring and summer) to generate general predictions on how the functioning and composition of multitrophic plankton communities (zoo-, phyto- and bacterioplankton) respond to pulse disturbances acting either in isolation or combined with press disturbances. As pulse disturbance, we used short-term changes in fish presence, and as press disturbance, we addressed the ongoing reduction in light availability caused by increased cloudiness and lake browning in many boreal and subarctic lakes. First, our results show that the top-down pulse disturbance had the strongest effects on both functioning and composition of the three trophic levels across sites and seasons, with signs for interactive impacts with the bottom-up press disturbance on phytoplankton communities. Second, community composition responses to disturbances were highly divergent between lakes and seasons: temporal accumulated community turnover of the same trophic level either increased (destabilization) or decreased (stabilization) in response to the disturbances compared to control conditions. Third, we found functional recovery from the pulse disturbances to be frequent at the end of most experiments. In a broader context, these results demonstrate that top-down, pulse disturbances, either alone or with additional constant stress upon primary producers caused by bottom-up disturbances, can induce profound but often functionally reversible changes across multiple trophic levels, which are strongly linked to spatial and temporal context dependencies. Furthermore, the identified dichotomy of disturbance effects on the turnover in community composition demonstrates the potential of disturbances to either stabilize or destabilize biodiversity patterns over time across a wide range of environmental conditions.

For aquatic scientists mesocosm experiments are important tools for hypothesis testing as they offer a compromise between experimental control and realism. Here we present a new mesocosm infrastructure-SITES AquaNET-located in five lakes connected to field stations in Sweden that cover a similar to 760 km latitudinal gradient. SITES AquaNet overcomes major hindrances in aquatic experimental research through: (i) openness to the scientific community, (ii) the potential to implement coordinated experiments across sites and time, and (iii) high-frequency measurements (temperature, photosynthetic photon flux density, turbidity and dissolved oxygen, chlorophyll a and phycocyanin concentrations) with an autonomous sensor system. Moreover, the infrastructure provides operational guidance and sensor expertise from technical staff, and connections to a multi-layered monitoring programme ("SITES Water") for each lake. This enables ecological observations from whole lake ecosystems to be compared with experimental studies aiming at disentangling major drivers and mechanisms underlying observed changes. Here we describe the technical properties of the infrastructure along with possibilities for experimental manipulations to tackle pressing issues in aquatic ecology and global change science. As a proof of concept, we also present a first mesocosm experiment across all five field sites with a cross-factorial design to evaluate responses of the sensor measurements to press/bottom-up (constant light reduction) and pulse/top-down (temporary fish predation) disturbances. This demonstrates the suitability of the infrastructure and autonomous sensor system to host modularized experiments and exemplifies the power and advantages of the approach to integrate a network of mecsocosm facilities with manageable costs across large geographic areas.

Ecological stability encompasses multiple dimensions of functional and compositional responses to environmental change. Though no single stability dimension used in isolation can fully reflect the overall response to environmental change, a common vulnerability assessment that integrates simultaneously across multiple stability components is highly desirable for ecological risk assessment. We develop both functional and compositional counterparts of a novel, integrative metric of overall ecological vulnerability (OEV). We test the framework with data from a modularized experiment replicated in five lakes over two seasons, examining functional and compositional responses to both pulse and press disturbances across three trophic groups. OEV is measured as the area under the curve integrated over the entire observation period, with the curve delimiting the difference between the disturbance treatment and undisturbed parallel controls, expressed either as the log response ratio of biomass (functional OEV) or community dissimilarity index (compositional OEV). Both, functional and compositional OEV correlated negatively with functional and compositional ‘resistance’, ‘temporal stability’ and ‘final/extent of recovery’ following both pulse and press disturbances, though less so with ‘resilience’ following a pulse disturbance. We also found a positive correlation between functional and compositional OEV, which reveals the potential to also evaluate the intricate linkage between biodiversity and functional change. Our findings demonstrate that OEV comprises a robust framework to: (a) capture simultaneously multiple functional and compositional stability components, and (b) quantify the functional consequences of biodiversity change. Our results provide the basis for an overarching framework for quantifying the overall vulnerability of ecosystems to environmental change, opening new possibilities for ecological risk assessment and management. Synthesis. Ecological stability comprises multiple dimensions that together encapsulate how ecosystems respond to environmental change. Considering these multiple aspects of stability simultaneously often poses a problem in environmental assessments, which frequently require overarching indicators of risk or vulnerability. While an analysis of multiple dimensions allows for deeper exploration of mechanisms, here we develop and test a new univariate indicator that integrates stability aspects under a broad range of disturbance regimes. Using a modularized experiment in Swedish lakes, we show that this integrative measure captures multiple stability dimensions reflecting compositional and functional vulnerability and their relationships between them.

The sea surface microlayer (SML) is the boundary layer between the ocean and the atmosphere and plays a unique role in marine biogeochemistry. Phytoplankton communities in this uppermost surface layer are exposed to extreme ultraviolet (UV) radiation and potentially high nutrient supplies. In order to understand the response of SML communities to such contrasting conditions, we conducted experiments at three different sites, the North Sea (open ocean) and two sites, outer and middle fjord, in the Sognefjord, Norway, with differing physical and chemical parameters. We manipulated light, nitrogen (N) and phosphorus (P) supply to natural communities collected from the SML and compared their response to that of the underlying water (ULW) communities at 1-m depth. Phytoplankton communities in both SML and ULW responded significantly to N addition, suggesting the upper 1-m surface phytoplankton communities were N-limited. While phytoplankton growth rates were higher with high N and high light supply, biomass yield was higher under low light conditions and with a combined N and P supply. Furthermore, biomass yield was generally higher in the ULW communities compared to SML communities. Nutrient and light effects on phytoplankton growth rates, particulate organic carbon (POC) and stoichiometry varied with geographical location. Phytoplankton growth rates in both SML and ULW at the open ocean station, the site with highest salinity, did not respond to light changes, whereas the communities in the middle fjord, characterized by high turbidity and low salinity, did experience light limitation. This work on the upper surface phytoplankton communities provides new insights into possible effects of coastal darkening and increases understanding of oceanic biogeochemical cycling.

International audience; A central concept for understanding the mechanisms linking diversity and primary production or more general, ecosystem functioning, is resource use efficiency (RUE). It quantifies the amount of biomass production over time relative to unit resource supplied, that is, represents a quota of matter use efficiency. Given anthropogenic alterations of biogeochemical cycles, the consequent changes in supply rate and especially supply ratio of nutrients will change. Using four species of freshwater phytoplankton, and their mixture, we asked how the RUE for nitrogen and phosphorus depends on the stoichiometry of resource supply and how this differs between single species and their mixture. We conducted a factorial laboratory experiment spanning 25 different nutrient supply treatments with differing absolute and relative nitrogen (N) and phosphorus (P) concentrations. N and P supply increased biomass production and decreased C : nutrient ratios and RUE for the respective nutrient, but always significantly affected by the supply of the respective other nutrient. Biomass peaked at molar N : P supply ratios above the Redfield ratio (18–22). Species tended to respond similarly to the resource gradients. Consequently, mixtures outperformed the component species only during early growth responses, but not regarding maximum biomass and RUE. Bioassays performed at the end of the main experiment revealed predominance of N‐limitation, but again strongly depending on the interaction between both nutrient gradients. Our study suggests that stoichiometric constraints of resource incorporation and RUE need to be accounted for when studying the response of phytoplankton to natural and anthropogenic variation in resource availability.

MILAN was a multidisciplinary, international study examining how the diel variability of sea-surface microlayer biogeochemical properties potentially impacts ocean-atmosphere interaction, in order to improve our understanding of this globally important process. The sea-surface microlayer (SML) at the air-sea interface is < 1 mm deep but it is physically, chemically and biologically distinct from the underlying water and the atmosphere above. Wind-driven turbulence and solar radiation are important drivers of SML physical and biogeochemical properties. Given that the SML is involved in all ocean-atmosphere exchanges of mass and energy, its response to solar radiation, especially in relation to how it regulates the air-sea exchange of climate-relevant gases and aerosols, is surprisingly poorly characterised. MILAN (sea-surface MIcroLAyer at Night) was an international, multidisciplinary campaign designed to specifically address this issue. In spring 2017, we deployed diverse sampling platforms (research vessels, radio-controlled catamaran, free-drifting buoy) to study full diel cycles in the coastal North Sea SML and in underlying water, and installed a land-based aerosol sampler. We also carried out concurrent ex situ experiments using several microsensors, a laboratory gas exchange tank, a solar simulator, and a sea spray simulation chamber. In this paper we outline the diversity of approaches employed and some initial results obtained during MILAN. Our observations of diel SML variability, e.g. the influence of changing solar radiation on the quantity and quality of organic material, and diel changes in wind intensity primarily forcing air-sea CO2 exchange, underline the value and the need of multidisciplinary campaigns for integrating SML complexity into the context of air-sea interaction.

While there is a lot of data on interactive effects of eutrophication and warming, to date, we lack data to generate reliable predictions concerning possible effects of nutrient decrease and temperature increase on community composition and functional responses. In recent years, a wide‐ranging trend of nutrient decrease (re‐oligotrophication) was reported for freshwater systems. Small lakes and ponds, in particular, show rapid responses to anthropogenic pressures and became model systems to investigate single as well as synergistic effects of warming and fertilization in situ and in experiments. Therefore, we set up an experiment to investigate the single as well as the interactive effects of nutrient reduction and gradual temperature increase on a natural freshwater phytoplankton community, using an experimental indoor mesocosm setup. Biomass production initially increased with warming but decreased with nutrient depletion. If nutrient supply was constant, biomass increased further, especially under warming conditions. Under low nutrient supply, we found a sharp transition from initially positive effects of warming to negative effects when resources became scarce. Warming reduced phytoplankton richness and evenness, whereas nutrient reduction at ambient temperature had positive effects on diversity. Our results indicate that temperature effects on freshwater systems will be altered by nutrient availability. These interactive effects of energy increase and resource decrease have major impacts on biodiversity and ecosystem function and thus need to be considered in environmental management plans.

1. A resampling of 38 small farmland ponds in Belgium after 10 years revealed a high temporal species turnover for both phytoplankton and zooplankton communities, associated with substantial changes in abiotic factors, especially a reduction in total phosphorus concentration. 2. Across ponds, phytoplankton biomass decreased while evenness and richness increased between the samplings in 2003 and 2013. By contrast, the zooplankton assemblage was characterised by lower biomass, richness and evenness in 2013. Ponds experiencing larger environmental change showed stronger changes in phytoplankton richness and evenness. 3. Resource use efficiency (RUE) of zooplankton increased with greater environmental change and zooplankton evenness, which points to a switch towards species with higher RUE or greater variety in food sources in higher trophic levels. 4. As ponds are important habitats for freshwater biodiversity and ecosystems services, the strong but predictable species turnover and the opposing effects of environmental change on different trophic levels need to be embedded in conservation and management plans.

We established a new indoor mesocosm facility, 12 fully controlled “Planktotrons”, designed to conduct marine and freshwater experiments for biodiversity and food web approaches using natural or artificial, benthic or planktonic communities. The Planktotrons are a unique and custom-tailored facility allowing long-term experiments. Wall growth can be inhibited by a rotating gate paddle with silicone lips. Additionally, temperature and light intensity are individually controllable for each Planktotron and the large volume (600 L) enables high-frequency or volume-intense measurements. In a pilot freshwater experiment various trophic levels of a pelagic food web were maintained for up to 90 d. First, an artificially assembled phytoplankton community of 11 species was inoculated in all Planktotrons. After 22 d, two ciliates were added to all, and three Daphnia species were added to six Planktotrons. After 72 d, dissolved organic matter (DOM, an alkaline soil extract) was added as an external disturbance to six of the 12 Planktotrons, involving three Planktotrons stocked with Daphnia and three without, respectively. We demonstrate the suitability of the Planktotrons for food web and biodiversity research. Variation among replicated Planktotrons (n = 3 minimum) did not differ from other laboratory systems and field experiments. We investigated population dynamics and interactions among the different trophic levels, and found them affected by the sequence of ciliate and Daphnia addition and the disturbance caused by addition of DOM.

This study addresses key processes in high-energy beach systems using an interdisciplinary approach. We assess spatial variations in subsurface pore water residence times, salinity, organic matter (OM) availability, and redox conditions and their effects on nutrient cycles as well as on microbial community patterns and microphytobenthos growth. At the study site on Spiekeroog Island, southern North Sea, beach hydrology is characterized by the classical zonation with an upper saline plume (USP), a saltwater wedge, and a freshwater discharge tube in between. Sediment and pore water samples were taken along a cross-shore transect from the dunes to the low water line reaching sediment depths down to 5 m below sediment surface. Spatial variations in pore water residence time, salinity, and organic matter availability lead to steep redox and nutrient gradients. Vertical and horizontal differences in the microbial community indicate the influence of these gradients and salinity on the community structure. Modeled seawater flux through the USP and freshwater flux through the tube are on average 2.8 and 0.75 m3 per day and meter of shoreline, respectively. Furthermore, ridge sediments at the lower beach discharge seawater at rates of 0.5 and 1.0 m3 per day and meter of shoreline towards the runnel and seaside, respectively. Applying seawater and freshwater fluxes and representative nutrient concentrations for the discharge zones, nutrient fluxes to adjacent nearshore waters are 117 mmol NH4+, 55 mmol PO43 − and 575 mmol Si(OH)4 per day and meter of shoreline. We propose that this nutrient efflux triggers growth of microphytobenthos on sediment surfaces of the discharge zone. A first comparison of nutrient discharge rates of the beach site with a nearby sandy backbarrier tidal flat margin indicates that the beach system might be of less importance in supplying recycled nutrients to nearshore waters than the backbarrier tidal flat area.

We developed an effective fluorometric technique to quantify extracellular carbonic anhydrase (eCA) present in natural seawater samples. The technique includes the separation of eCA from cells to achieve low detection limits through high signal : noise ratios. eCA was efficiently extracted from cell membranes by treatment with 0.1 M phosphate buffer containing 2.5 M NaCl. The free eCA specifically forms a fluorescent complex with dansylamide, and the detection limit of the complex is below 0.1 nM. We applied the technique to samples from different culture solutions and natural seawater collected from the Baltic Sea. We observed eCA concentrations to be in the range of 0.10–0.67 nM in natural seawater. The data indicated that this technique is very sensitive, accurate, and feasible for routine and shipboard measurement of eCA from natural seawater. It is therefore an effective and rapid tool to investigate the carbon acquisition of phytoplankton both in mono culture as well natural communities.

Nutrient availability and temperature are important drivers of phytoplankton growth and stoichiometry. However, the interactive effects of nutrients and temperature on phytoplankton have been analyzed mostly by addressing changes in average temperature, whereas recent evidence suggests an important role of temperature fluctuations. In a laboratory experiment, we grew a natural phytoplankton community under fluctuating and constant temperature regimes across 25 combinations of nitrogen (N) and phosphorus (P) supply. Temperature fluctuations decreased phytoplankton growth rate (r(max)), as predicted by nonlinear averaging along the temperature-growth relationship. r(max) increased with increasing P supply, and a significant temperature x P x N interaction reflected that the shape of the thermal reaction norm depended on nutrients. By contrast, phytoplankton carrying capacity increased with N supply and in fluctuating rather than constant temperature. Higher phytoplankton N:P ratios under constant temperature showed that temperature regimes affected cellular nutrient incorporation. Minor differences in species diversity and composition existed. Our results suggest that temperature variability interacts with nutrient supply to affect phytoplankton physiology and stoichiometry at the community level.

Sea cucumbers are bottom dwelling invertebrates, which are mostly found on subtropical and tropical sea grass beds, sandy reef flats, or reef slopes. Although constantly exposed to fouling communities in these habitats, many species are surprisingly free of invertebrate epibionts and microfouling algae such as diatoms. In our study, we investigated the anti-fouling (AF) activities of different crude extracts of tropical Indo-Pacific sea cucumber species against the fouling diatom Cylindrotheca closterium. Nine sea cucumber species from three genera (i.e., Holothuria, Bohadschia, Actinopyga) were selected and extracted to assess their AF activities. To verify whether the sea cucumber characteristic triterpene glycosides were responsible for the observed potent AF activities, we tested purified fractions enriched in saponins isolated from Bohadschia argus, representing one of the most active anti-fouling extracts. Saponins were quantified by vanillin-sulfuric acid colorimetric assays and identified by LC-MS and LC-MS/MS analyses. We were able to demonstrate that AF activities in sea cucumber extracts were species-specific, and growth inhibition as well as attachment of the diatom to surfaces is dependent on the saponin concentration (i.e., Actinopyga contained the highest quantities), as well as on the molecular composition and structure of the present saponins (i.e., Bivittoside D derivative was the most bioactive compound). In conclusion, the here performed AF assay represents a promising and fast method for selecting the most promising bioactive organism as well as for identifying novel compounds with potent AF activities for the discovery of potentially novel pharmacologically active natural products.

About 60 years ago, the critical depth hypothesis was proposed to describe the occurrence of spring phytoplankton blooms and emphasized the role of stratification for the timing of onset. Since then, several alternative hypotheses appeared focusing on the role of grazing and mixing processes such as turbulent convection or wind activity. Surprisingly, the role of community composition—and thus the distribution of phytoplankton traits—for bloom formation has not been addressed. Here, we discuss how trait variability between competing species might influence phytoplankton growth during the onset of the spring bloom. We hypothesize that the bloom will only occur if there are species with a combination of traits fitting to the environmental conditions at the respective location and time. The basic traits for formation of the typical spring bloom are high growth rates and photoadaptation to low light conditions, but other traits such as nutrient kinetics and grazing resistance might also be important. We present concise ideas on how to test our theoretical considerations experimentally. Furthermore, we suggest that future models of phytoplankton blooms should include both water column dynamics and variability of phytoplankton traits to make realistic projections instead of treating the phytoplankton bloom as an aggregate community phenomenon.

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

Biodiversity can strongly influence trophic interactions. The nutritional quality of prey communities and how it is related to the prey diversity is suspected to be a major driver of biodiversity effects. As consumer growth can be co-limited by the supply of several biochemical components, biochemically diverse prey communities should promote consumer growth. Yet, there is no clear consensus on how prey specific diversity is linked to community biochemical diversity since previous studies have considered only single nutritional quality traits. Here, we demonstrate that phytoplankton biochemical traits (fatty acids and sterols) can to a large extent explain Daphnia magna growth and its apparent dependence on phytoplankton species diversity. We find strong correlative evidence between phytoplankton species diversity, biochemical diversity, and growth. The relationship between species diversity and growth was partially explained by the fact that in many communities Daphnia was co-limited by long chained polyunsaturated fatty acids and sterols, which was driven by different prey taxa. We suggest that biochemical diversity is a good proxy for the presence of high food quality taxa, and a careful consideration of the distribution of the different biochemical traits among species is necessary before concluding about causal links between species diversity and consumer performance.

The intermediate disturbance hypothesis (IDH) predicts that disturbances of moder- ate frequency or intensity result in maximum diversity, while the dynamic equilibrium hypothesis (DEH) proposes that disturbance and productivity interact in their effects on diversity. These hypotheses were tested with a marine ciliate community in a mesocosm study conducted in a Nor- wegian fjord. Two nutrient regimes, 6 disturbance frequencies (undisturbed to daily disturbed) and 2 replicates for each treatment were used. To disturb the system, we repeatedly lowered and raised a disk to mix the water column. The experiment lasted 3 wk, with samples (7 including the initial sample) taken for nutrients and physical parameters, chl a, ciliates and mesozooplankton. The system reacted more strongly to nutrient addition than to our chosen range of disturbance frequencies. Nutrient addition resulted in higher chl a levels and ciliate abundance but reduced ciliate diversity without changing the mean ciliate species richness. The reduction in diversity was due to the very strong, positive response to nutrient addition by the mixotrophic Mesodinium rubrum. In contrast, metazoan predators on ciliates (primarily copepods) did not respond to any of the experimental manipulations, at least partially due to the duration of the experiment rela- tive to their generation times. While disturbance had varied and at times marked effects on the mesocosm communities, the expected unimodal response to disturbance frequency was not seen, nor was the expected interaction between disturbance and productivity. While the IDH and DEH have intuitive appeal, they appear to have limited power to predict natural ciliate community patterns.

Over the past four decades, mesocosm studies have been successfully used for a wide range of applications and have provided a lot of information on trophic interactions and biogeochemical cycling of aquatic ecosystem. However, the setup of such mesocosms (e.g., dimensions and duration of experiments) needs to be adapted to the relevant biological processes being investigated. Mixing of the water column is an important factor to be considered in mesocosm experiments because enclosing water in an artificial chamber always alters the mixing regime. Various approaches have been applied to generate mixing in experimental ecosystems, including pure mechanical mixing (e.g., using a disc), airlifts, bubbling with compressed air, and pumping. In this study, we tested different mixing techniques for outdoor mesocosms and their impact on plankton biomass and community composition. We compared mesocosms mixed with a disc, an airlift-system, and bubbling, and used a nonactively mixed mesocosm as a control. We investigated phytoplankton, ciliate, and zooplankton communities during a 19-d mesocosm experiment. Based on our results, we concluded that mechanical mixing with a disc was the most effective technique due to the undertow produced by lowering and lifting the disc. While no mixing technique affected seston biomass, zooplankton biomass was highest in the treatments mixed with the disc. The airlift treatments had the lowest relative share of small flagellates. However, no further differences in phytoplankton community composition occurred and no differences in zooplankton community composition existed between all actively mixed treatments.

Ptacnik, Robert ... et al.-- 10 pages, 4 figures, 1 table, supplementary information https://dx.doi.org/10.1038/srep29286, Mixotrophs combine photosynthesis with phagotrophy to cover their demands in energy and essential nutrients. This gives them a competitive advantage under oligotropihc conditions, where nutrients and bacteria concentrations are low. As the advantage for the mixotroph depends on light, the competition between mixo- and heterotrophic bacterivores should be regulated by light. To test this hypothesis, we incubated natural plankton from the ultra-oligotrophic Eastern Mediterranean in a set of mesocosms maintained at 4 light levels spanning a 10-fold light gradient. Picoplankton (heterotrophic bacteria (HB), pico-sized cyanobacteria, and small-sized flagellates) showed the fastest and most marked response to light, with pronounced predator-prey cycles, in the high-light treatments. Albeit cell specific activity of heterotrophic bacteria was constant across the light gradient, bacterial abundances exhibited an inverse relationship with light. This pattern was explained by light-induced top-down control of HB by bacterivorous phototrophic eukaryotes (PE), which was evidenced by a significant inverse relationship between HB net growth rate and PE abundances. Our results show that light mediates the impact of mixotrophic bacterivores. As mixo- and heterotrophs differ in the way they remineralize nutrients, these results have far-reaching implications for how nutrient cycling is affected by light, The experiment was financed by the European Union Seventh Framework Program MESOAQUA (FP7/2009-2013) coordinated by JCN under grant agreement no. 228224. R.P. further acknowledges financial support by the German Research Council (DFG Pt 5/3-1), JMG by Spanish project REMEI (CTM2015-70340) and AC by project PROTOS (CTM2009-08783) from the Spanish Ministry of Economy and Competitiveness

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.

Effects of temperature changes on phytoplankton communities seem to be highly context-specific, but few studies have analyzed whether this context specificity depends on differences in the abiotic conditions or in species composition between studies. We present an experiment that allows disentangling the contribution of abiotic and biotic differences in shaping the response to two aspects of temperature change: permanent increase of mean temperature versus pulse disturbance in form of a heat wave. We used natural communities from six different sites of a floodplain system as well as artificially mixed communities from laboratory cultures and grew both, artificial and natural communities, in water from the six different floodplain lakes (sites). All 12 contexts (2 communities × 6 sites) were first exposed to three different temperature levels (12, 18, 24 °C, respectively) and afterward to temperature pulses (4 °C increase for 7 h day−1). Temperature-dependent changes in biomass and community composition depended on the initial composition of phytoplankton communities. Abiotic conditions had a major effect on biomass of phytoplankton communities exposed to different temperature conditions, however, the effect of biotic and abiotic conditions together was even more pronounced. Additionally, phytoplankton community responses to pulse temperature effects depended on the warming history. By disentangling abiotic and biotic effects, our study shows that temperature-dependent effects on phytoplankton communities depend on both, biotic and abiotic constraints. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3693-3) contains supplementary material, which is available to authorized users.

Phytoplankton play an important role as primary producers and thus can affect higher trophic levels. Phytoplankton growth and diversity may, besides other factors, be controlled by seasonal temperature changes and increasing water temperatures. In this study, we investigated the combined effects of temperature and diversity on phytoplankton growth. In a controlled laboratory experiment, monocultures of 15 freshwater phytoplankton taxa (green algae, cyanobacteria, and diatoms) as well as 25 mixed communities of different species richness (2–12 species) and taxa composition were exposed to constant temperatures of 12, 18, and 24 °C. Additionally, they were exposed to short-term daily temperature peaks of +4 °C. Increased species richness had a positive effect on phytoplankton growth rates and phosphorous content at all temperature levels, with maximum values occurring at 18 °C. Overyielding was observed at almost all temperature levels and could mostly be explained by complementary traits. Higher temperatures resulted in higher fractions of cyanobacteria in communities. This negative effect of temperature on phytoplankton diversity following a shift in community composition was most obvious in communities adapted to cooler temperatures, pointing to the assumption that relative temperature changes may be more important than absolute ones. Electronic supplementary material The online version of this article (doi:10.1007/s00442-012-2419-4) contains supplementary material, which is available to authorized users.

Actively motile, flagellated phytoplankton taxa often exploit vertical gradients in the availability of light and nutrients. The ability to move actively usually goes along with low investment in s ...

Both low and high rates of light supply can restrict herbivore growth rates by limiting either the quantity (photosynthetically fixed carbon) or the nutritional quality (nutrient content per fixed carbon) of the herbivores’ food. The ‘‘light–nutrient hypothesis,’’ therefore, predicts that, if phosphorus (P) supply is sufficiently low, production of herbivorous zooplankton should be unimodally related to light intensity. We manipulated the light regime of six different algal communities in a field experiment and investigated the effect of these manipulations on Daphnia growth. The algal communities came from six lakes having different total phosphorus concentrations, ranging from oligotrophic to eutrophic. Seston carbon (C) and seston carbon-to-phosphorus ratios in communities from oligotrophic and mesotrophic lakes increased with higher light availability. Across all lakes, the strength of these responses was related to algal diversity. More diverse algal communities showed a stronger increase than less diverse communities in both their carbon biomass and their C : P ratio with increasing light. Furthermore, in oligotrophic and mesotrophic treatments, Daphnia growth was highest at intermediate light intensities. In contrast, seston parameters and Daphnia growth were only weakly related to light supply in communities from eutrophic lakes.