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Abstract Copepods are the most abundant metazoans on Earth, driving cycles of key elements in aquatic systems, most prominently carbon (C), nitrogen (N), and phosphorus (P). One key factor determining nutrient cycling is copepod somatic stoichiometry, which can reflect ecological strategy. We conducted a systematic review that updates the seminal work of Båmstedt (1986) by summarizing the effects of latitude, habitat, life history stage, and taxonomy on C:N, C:P, and N:P ratios of field‐collected copepods. We found that differences among copepod families accounted for the greatest variation, with the Rhincalanidae and Diaptomidae being particularly C‐rich, while the Calanidae were more N‐ and P‐rich. Copepod C:N was higher in inland waters compared with animals from marine environments in both copepodites and adult females, matching the higher C content of seston in many inland freshwaters. For both copepodites and adult females, mid‐latitude animals had higher C:N and C:P than high‐latitude animals, which matched predictions based on the availability of nutrients or adaptation to cold environments. More data must be gathered to fill gaps in our knowledge of copepod stoichiometry, focusing particularly on younger life stages, non‐calanoids, low and high latitudes, the southern hemisphere, and estuarine and some inland water habitats, including large lakes. Such information will help better parameterize models of aquatic ecosystems and improve our understanding of how copepods influence consumer‐driven nutrient cycling and food web dynamics.

The performance of herbivore consumers in freshwater ecosystems is influenced by food quality and quantity, which are crucial factors in understanding energy flow. However, the comprehensive effects of these factors on consumer growth based on natural food research remain unclear. To address this gap, we conducted a growth experiment culturing the Cladocera Daphnia similis in natural lake water collected from 30 subtropical lakes. Our results showed that the seston concentration (SC), seston phosphorus (seston P), and seston carbon: phosphorus ratio (seston C:P, in moles) were the most important factors in influencing the growth rate of D. similis according to the Random Forest model. Specifically, the growth rate of D. similis was significantly positive correlated with SC and seston P, and significantly negative correlated with seston C:P. D. similis exhibited the optimal growth performance within the seston C:P range of 32.8 to 69.8, with a sharp decline in growth rate observed at a break point of seston C:P of 70. The combined effect of food quality and quantity on growth rate was that higher SC (≥0.26 mg C/L), lower seston C:P (≤69.80), and higher P (≥0.11 mmol/g) were associated with significantly higher growth rates. Additionally, the growth rate increased significantly with the biomass of cryptophyte, indicating the importance of food composition. Our study shows that the growth rate of D. similis is co-affected by the quality and quantity of natural food. SC, seston C:P, seston P and algae composition are reliable indicators for assessing the growth rate of consumer in freshwater ecosystems. The inhibitory effect of low C:P on consumer growth should be applied with caution when assessing the development of herbivore consumers in natural lakes.

ABSTRACT Net growth of microbial populations, that is, changes in abundances over time, can be studied using 16S rRNA fluorescence in situ hybridization (FISH). However, this approach does not differentiate between mortality and cell division rates. We used FISH-based image cytometry in combination with dilution culture experiments to study net growth, cell division, and mortality rates of four bacterial taxa over two distinct phytoplankton blooms: the oligotrophs SAR11 and SAR86, and the copiotrophic phylum Bacteroidetes, and its genus Aurantivirga. Cell volumes, ribosome content, and frequency of dividing cells (FDC) co-varied over time. Among the three, FDC was the most suitable predictor to calculate cell division rates for the selected taxa. The FDC-derived cell division rates for SAR86 of up to 0.8/day and Aurantivirga of up to 1.9/day differed, as expected for oligotrophs and copiotrophs. Surprisingly, SAR11 also reached high cell division rates of up to 1.9/day, even before the onset of phytoplankton blooms. For all four taxonomic groups, the abundance-derived net growth (−0.6 to 0.5/day) was about an order of magnitude lower than the cell division rates. Consequently, mortality rates were comparably high to cell division rates, indicating that about 90% of bacterial production is recycled without apparent time lag within 1 day. Our study shows that determining taxon-specific cell division rates complements omics-based tools and provides unprecedented clues on individual bacterial growth strategies including bottom–up and top–down controls. IMPORTANCE The growth of a microbial population is often calculated from their numerical abundance over time. However, this does not take cell division and mortality rates into account, which are important for deriving ecological processes like bottom–up and top–down control. In this study, we determined growth by numerical abundance and calibrated microscopy-based methods to determine the frequency of dividing cells and subsequently calculate taxon-specific cell division rates in situ. The cell division and mortality rates of two oligotrophic (SAR11 and SAR86) and two copiotrophic (Bacteroidetes and Aurantivirga) taxa during two spring phytoplankton blooms showed a tight coupling for all four taxa throughout the blooms without any temporal offset. Unexpectedly, SAR11 showed high cell division rates days before the bloom while cell abundances remained constant, which is indicative of strong top–down control. Microscopy remains the method of choice to understand ecological processes like top–down and bottom–up control on a cellular level.

Climate model-informed experiments indicate that marine plankton food webs may be restructured in the future. There exists a tipping point where a crucial plankton community structure changes under increased nitrogen to phosphorous ratios, pCO2 and temperature.

Climate change combined with anthropogenic stressors (e.g. overfishing, habitat destruction) may have particularly strong effects on threatened populations of coastal invertebrates. The collapse of the population of European lobster (Homarus gammarus) around Helgoland constitutes a good example and prompted a large-scale restocking program. The question arises if recruitment of remaining natural individuals and program-released specimens could be stunted by ongoing climate change. We examined the joint effect of ocean warming and acidification on survival, development, morphology, energy metabolism and enzymatic antioxidant activity of the larval stages of the European lobster. Larvae from four independent hatches were reared from stage I to III under a gradient of 10 seawater temperatures (13–24°C) combined with moderate (∼470 µatm) and elevated (∼1160 µatm) seawater pCO2 treatments. Those treatments correspond to the shared socio-economic pathways (SSP), SSP1-2.6 and SSP5-8.5 (i.e. the low and the very high greenhouse gas emissions respectively) projected for 2100 by the Intergovernmental Panel on Climate Change. Larvae under the elevated pCO2 treatment had not only lower survival rates, but also significantly smaller rostrum length. However, temperature was the main driver of energy demands with increased oxygen consumption rates and elemental C:N ratio towards warmer temperatures, with a reducing effect on development time. Using this large temperature gradient, we provide a more precise insight on the aerobic thermal window trade-offs of lobster larvae and whether exposure to the worst hypercapnia scenario may narrow it. This may have repercussions on the recruitment of the remaining natural and program-released specimens and thus, in the enhancement success of future lobster stocks.

Consumer regulation of lipid composition during assimilation of dietary items is related to their ecology, habitat, and life cycle, and may lead to extra energetic costs associated with the conversion of dietary material into the fatty acids (FAs) necessary to meet metabolic requirements. For example, lipid-rich copepods from temperate and polar latitudes must convert assimilated dietary FAs into wax esters, an efficient type of energy storage which enables them to cope with seasonal food shortages and buoyancy requirements. Lipid-poor copepods, however, tend to not be as constrained by food availability as their lipid-rich counterparts and, thus, should have no need for modifying dietary FAs. Our objective was to test the assumption that Temora longicornis, a proxy species for lipid-poor copepods, does not regulate its lipid composition. Isotopically-enriched (13C) diatoms were fed to copepods during a 5-day laboratory experiment. Compound-specific stable isotope analysis of algae and copepod samples was performed in order to calculate dietary FA assimilation, turnover, and assimilation efficiency into copepod FAs. Approximately 65% of the total dietary lipid carbon (C) assimilated (913 ± 68 ng C ind-1 at the end of the experiment) was recorded as polyunsaturated FAs, with 20 and 15% recorded as saturated and monounsaturated FAs, respectively. As expected, T. longicornis assimilated dietary FAs in an unregulated, non-homeostatic manner, as evidenced by the changes in its FA profile, which became more similar to that of their diet. Copepods assimilated 11% of the total dietary C (or 40% of the dietary lipid C) ingested in the first two days of the experiment. In addition, 34% of their somatic growth (in C) after two days was due to the assimilation of dietary C in FAs. Global warming may lead to increased proportions of smaller copepods in the oceans, and to a lower availability of algae-produced essential FAs. In order for changes in the energy transfer in marine food webs to be better understood, it is important that future investigations assess a broader range of diets as well as lipid-poor zooplankton from oceanographic areas throughout the world’s oceans.

Marine zooplankton are central components of holistic ecosystem assessments due to their intermediary role in the food chain, linking the base of the food chain with higher trophic levels. As a result, these organisms incorporate the inherent properties and changes occurring atall levels of the marine ecosystem, temporally integrating signatures of physical and chemical conditions. For this reason, zooplankton-based biometrics are widely accepted as useful tools for assessing and monitoring the ecological health and integrity of aquatic systems. The European Marine Strategy Framework Directive (EU-MSFD) requires the use of different types of bio-monitors, including zooplankton, to monitor progress towards achieving specific environmental and water quality targets in EU. However, there is currently no comprehensive synthesis of zooplankton indices development, use, and associated challenges. We addressed this issue with a two-step approach. First, we formulated the indicator-metrics-indices cycle (IMIC) to redefine the closely related but often ambiguously utilized terms - indicator, metric and index, highlighting the convergence between them and the iterative nature of their interaction. Secondly, we formulated frameworks for synthesizing, presenting and systematically applying zooplankton indices based on the IMIC framework. The main benefits of the IMIC are twofold: 1). to disambiguate the key elements: indicators, metrics, and indices, revealing their links to an operational ecological indicator system, and 2) to serve as an organizing tool for the coherent classification of indices according to the MSFD descriptors. Using the IMIC framework, we identified and described two broad categories of indices namely the core biodiversity indices already in use in the Baltic Sea and North Atlantic regions, including the ‘Zooplankton Mean Size and Total Stock (zooplankton MSTS)’ and 'Plankton Lifeforms index (PLI)', and stressor-response indices retrieved from the existing literature, elucidating their applicability to different MSFD descriptors. Finally, major challenges of developing new indices and applying existing ones in the context of the MSFD were critically addressed and some solutions were proposed.

In this study, we created a dataset of a continuous three-year 18S metabarcoding survey to identify eukaryotic parasitoids, and potential connections to hosts at the Long-Term Ecological Research station Helgoland Roads. The importance of parasites and parasitoids for food web dynamics has previously been recognized mostly in terrestrial and freshwater systems, while marine planktonic parasitoids have been understudied in comparison to those. Therefore, the occurrence and role of parasites and parasitoids remains mostly unconsidered in the marine environment. We observed high abundances and diversity of parasitoid operational taxonomic units in our dataset all year round. While some parasitoid groups were present throughout the year and merely fluctuated in abundances, we also detected a succession of parasitoid groups with peaks of individual species only during certain seasons. Using co-occurrence and patterns of seasonal occurrence, we were able to identify known host-parasitoid dynamics, however identification of new potential host-parasitoid interactions was not possible due to their high dynamics and variability in the dataset.

Morphological and physiological characteristics of phytoplankton cells are highly sensitive to changes in environmental conditions and, in turn, influence the dynamics of phytoplankton populations and communities. To cope with environmental change, trait variability and phenotypic plasticity may play an important role. Since global change comprises simultaneous changes in abiotic parameters, we assessed the impact of multiple drivers on functional traits of the diatom Thalassiosira (Conticribra) weissflogii by manipulating concurrently temperature, pCO2, and dissolved nitrogen:phosphorus (N:P) ratio. We tested three scenarios: ambient (ambient temperature and atmospheric pCO2; 16 N:P ratio), moderate future scenario (+1.5°C and 800 ppm CO2; 25 N:P ratio), and more severe future scenario (+3°C and 1000 ppm CO2; 25 N:P ratio). We applied flow cytometry to measure on single-cell levels to investigate trait variability and phenotypic plasticity within one strain of diatoms. Growth rates differed significantly between the treatments and were strongly correlated with cell size and cellular chlorophyll a content. We observed a negative correlation of growth rate with chlorophyll a variability among single strain populations and a negative correlation with the phenotypic plasticity of cell size, i.e. when growth rates were higher, the cell size cell-to-cell variability within cultures was lower. Additionally, the phenotypic plasticity in cell size was lower under the global change scenarios. Overall, our study shows that multiple traits are interlinked and driven by growth rate and that this interconnection may partly be shaped by environmental factors.

Marine fungi are an important component of pelagic planktonic communities. However, it is not yet clear how individual fungal taxa are integrated in marine processes of the microbial loop and food webs. Most likely, biotic interactions play a major role in shaping the fungal community structure. Thus, the aim of our work was to identify possible biotic interactions of mycoplankton with phytoplankton and zooplankton groups and among fungi, and to investigate whether there is coherence between interactions and the dynamics, abundance and temporal occurrence of individual fungal OTUs. Marine surface water was sampled weekly over the course of 1 year, in the vicinity of the island of Helgoland in the German Bight (North Sea). The mycoplankton community was analyzed using 18S rRNA gene tag-sequencing and the identified dynamics were correlated to environmental data including phytoplankton, zooplankton, and abiotic factors. Finally, co-occurrence patterns of fungal taxa were detected with network analyses based on weighted topological overlaps (wTO). Of all abundant and persistent OTUs, 77% showed no biotic relations suggesting a saprotrophic lifestyle. Of all other fungal OTUs, nearly the half (44%) had at least one significant negative relationship, especially with zooplankton and other fungi, or to a lesser extent with phytoplankton. These findings suggest that mycoplankton OTUs are embedded into marine food web chains via highly complex and manifold relationships such as parasitism, predation, grazing, or allelopathy. Furthermore, about one third of all rare OTUs were part of a dense fungal co-occurrence network probably stabilizing the fungal community against environmental changes and acting as functional guilds or being involved in fungal cross-feeding. Placed in an ecological context, strong antagonistic relationships of the mycoplankton community with other components of the plankton suggest that: (i) there is a top-down control by fungi on zooplankton and phytoplankton; (ii) fungi serve as a food source for zooplankton and thereby transfer nutrients and organic material; (iii) the dynamics of fungi harmful to other plankton groups are controlled by antagonistic fungal taxa.

Ocean acidification (OA) is affecting marine ecosystems through changes in carbonate chemistry that may influence consumers of phytoplankton, often via trophic pathways. Using a mesocosm approach, we investigated OA effects on a subtropical zooplankton community during oligotrophic, bloom, and post-bloom phases under a range of different pCO2 levels (from ∼400 to ∼1480 μatm). Furthermore, we simulated an upwelling event by adding 650 m-depth nutrient-rich water to the mesocosms, which initiated a phytoplankton bloom. No effects of pCO2 on the zooplankton community were visible in the oligotrophic conditions before the bloom. The zooplankton community responded to phytoplankton bloom by increased abundances in all treatments, although the response was delayed under high-pCO2 conditions. Microzooplankton was dominated by small dinoflagellates and aloricate ciliates, which were more abundant under medium- to high-pCO2 conditions. The most abundant mesozooplankters were calanoid copepods, which did not respond to CO2 treatments during the oligotrophic phase of the experiment but were found in higher abundance under medium- and high-pCO2 conditions toward the end of the experiment, most likely as a response to increased phyto- and microzooplankton standing stocks. The second most abundant mesozooplankton taxon were appendicularians, which did not show a response to the different pCO2 treatments. Overall, CO2 effects on zooplankton seemed to be primarily transmitted through significant CO2 effects on phytoplankton and therefore indirect pathways. We conclude that elevated pCO2 can change trophic cascades with significant effects on zooplankton, what might ultimately affect higher trophic levels in the future.

The meroplanktonic larvae of benthic organisms are an important seasonal component of the zooplankton in temperate coastal waters. The larvae of the reef-building polychaete Lanice conchilega contribute up to 15% of the summer zooplankton biomass in the North Sea. Despite their importance for reef maintenance (which positively affects the benthic community), little is known about the trophic ecology of this meroplanktonic larva. Qualitative and quantitative estimates of carbon (C) transfer between trophic levels and of fatty acid (FA)-specific assimilation, biosynthesis, and bioconversion can be obtained by compound-specific stable isotope analysis of FA. The present work tested the hypothesis that the concept of fatty acid trophic markers (FATM), widely used for studies on holoplankton with intermediate to high lipid contents, is also applicable to lipid-poor organisms such as meroplanktonic larvae. The incorporation of isotopically-enriched dietary C by L. conchilega larvae was traced, and lipid assimilation did not follow FA-specific relative availabilities in the diet. Furthermore, FAs that were unavailable in the diet, such as 22:5(n-3), were recorded in L. conchilega, suggesting their bioconversion by the larvae. The results indicate that L. conchilega larvae preferentially assimilate certain FAs and regulate their FA composition (lipid homeostasis) independently of that of their diet. Their quasi-homeostatic response to dietary FA availability could imply that the concept of FATM has limited application in lipid-poor organisms such as L. conchilega larvae.

Live feed are still necessary for the rearing of larval stages of several fish species, especially marine ones. Compared to Artemia, copepods are of superior quality. This is based on a suite of traits like size, movement, and nutritional value. Copepods are for example usually high in protein and fatty acids. Essential fatty acid profiles reflect to a large degree the fatty acid supply, which provides the opportunity to manipulate fatty acid profiles of, amongst others, copepods. By manipulating nutrient supply of the algae Rhodomonas salina we were able to double essential fatty acid concentrations in naupliar and copepodit life stages of the copepod Acartia tonsa. However, this lead to growth depression rather than to increased growth rates in a series of consumer species, including larval fish. The reason for the growth depression is likely to be mineral deficiencies occurring along with the nutrient manipulation of the algae.

The theories developed in ecological stoichiometry (ES) are fundamentally based on traits. Traits directly linked to cell/body stoichiometry, such as nutrient uptake and storage, as well as the associated trade-offs, have the potential to shape ecological interactions such as competition and predation within ecosystems. Further, traits that indirectly influence and are influenced by nutritional requirements, such as cell/body size and growth rate, are tightly linked to organismal stoichiometry. Despite their physiological and ecological relevance, traits are rarely explicitly integrated in the framework of ES and, currently, the major challenge is to more closely inter-connect ES with trait-based ecology (TBE). Here, we highlight four interconnected nutrient trait groups, i.e., acquisition, body stoichiometry, storage, and excretion, which alter interspecific competition in autotrophs and heterotrophs. We also identify key differences between producer-consumer interactions in aquatic and terrestrial ecosystems. For instance, our synthesis shows that, in contrast to aquatic ecosystems, traits directly influencing herbivore stoichiometry in forested ecosystems should play only a minor role in the cycling of nutrients. We furthermore describe how linking ES and TBE can help predict the ecosystem consequences of global change. The concepts we highlight here allow us to predict that increasing N:P ratios in ecosystems should shift trait dominances in communities toward species with higher optimal N:P ratios and higher P uptake affinity, while decreasing N retention and increasing P storage.

Ocean acidification may affect zooplankton directly by decreasing in pH, as well as indirectly via trophic pathways, where changes in carbon availability or pH effects on primary producers may cascade up the food web thereby altering ecosystem functioning and community composition. Here, we present results from a mesocosm experiment carried out during 113 days in the Gullmar Fjord, Skagerrak coast of Sweden, studying plankton responses to predicted end-of-century pCO2 levels. We did not observe any pCO2 effect on the diversity of the mesozooplankton community, but a positive pCO2 effect on the total mesozooplankton abundance. Furthermore, we observed species-specific sensitivities to pCO2 in the two major groups in this experiment, copepods and hydromedusae. Also stage-specific pCO2 sensitivities were detected in copepods, with copepodites being the most responsive stage. Focusing on the most abundant species, Pseudocalanus acuspes, we observed that copepodites were significantly more abundant in the high-pCO2 treatment during most of the experiment, probably fuelled by phytoplankton community responses to high-pCO2 conditions. Physiological and reproductive output was analysed on P. acuspes females through two additional laboratory experiments, showing no pCO2 effect on females' condition nor on egg hatching. Overall, our results suggest that the Gullmar Fjord mesozooplankton community structure is not expected to change much under realistic end-of-century OA scenarios as used here. However, the positive pCO2 effect detected on mesozooplankton abundance could potentially affect biomass transfer to higher trophic levels in the future.

The acidification of the oceans could potentially alter marine plankton communities with consequences for ecosystem functioning. While several studies have investigated effects of ocean acidification on communities using traditional methods, few have used genetic analyses. Here, we use community barcoding to assess the impact of ocean acidification on the composition of a coastal plankton community in a large scale, in situ, long-term mesocosm experiment. High-throughput sequencing resulted in the identification of a wide range of planktonic taxa (Alveolata, Cryptophyta, Haptophyceae, Fungi, Metazoa, Hydrozoa, Rhizaria, Straminipila, Chlorophyta). Analyses based on predicted operational taxonomical units as well as taxonomical compositions revealed no differences between communities in high CO2 mesocosms (~ 760 μatm) and those exposed to present-day CO2 conditions. Observed shifts in the planktonic community composition were mainly related to seasonal changes in temperature and nutrients. Furthermore, based on our investigations, the elevated CO2 did not affect the intraspecific diversity of the most common mesozooplankter, the calanoid copepod Pseudocalanus acuspes. Nevertheless, accompanying studies found temporary effects attributed to a raise in CO2. Differences in taxa composition between the CO2 treatments could, however, only be observed in a specific period of the experiment. Based on our genetic investigations, no compositional long-term shifts of the plankton communities exposed to elevated CO2 conditions were observed. Thus, we conclude that the compositions of planktonic communities, especially those in coastal areas, remain rather unaffected by increased CO2.

We investigated the trophic level of larvae of two fish species (lesser sandeel. Ammodytes marinus, and dab, Limanda limanda) in spring 2004 by means of stable isotope signatures at the helgoland roads station (54°11.18’N and 07°54.00’ E). the signatures were contrasted with the spring succession of phytoplankton and zooplankton. Phytoplankton biomass remained low until the middle of April, when a bloom developed. the δ15N signature of the seston increased until the bloom started then decreased during the bloom. the δ15N of the larvae of both fish species generally followed the development of the baseline, but the decrease in the fishes’ trophic level (expressed as the ∆δ15N was larger than that of the seston, suggesting that larval fish switched their diet to lower trophic levels. For larval sandeel we found that the switch to feeding on lower trophic levels was accompanied by a decrease in nutritional condition, while this pattern was not apparent in larval dab. hence, larval sandeel were not able to substitute the lack of high trophic level zooplankton prey with prey originating from lower trophic levels; however, at least the smaller size classes of larval dab could successfully switch diets.

Ocean acidification is considered as a crucial stressor for marine communities. In this study, we tested the effects of the IPCC RPC6.0 end-of-century acidification scenario on a natural plankton community in the Gullmar Fjord, Sweden, during a long-term mesocosm experiment from a spring bloom to a mid-summer situation. The focus of this study was on microzooplankton and its interactions with phytoplankton and mesozooplankton. The microzooplankton community was dominated by ciliates, especially small Strombidium sp., with the exception of the last days when heterotrophic dinoflagellates increased in abundance. We did not observe any effects of high CO2 on the community composition and diversity of microzooplankton. While ciliate abundance, biomass and growth rate were not affected by elevated CO2, we observed a positive effect of elevated CO2 on dinoflagellate abundances. Additionally, growth rates of dinoflagellates were significantly higher in the high CO2 treatments. Given the higher Chlorophyll a content measured under high CO2, our results point at mainly indirect effects of CO2 on microzooplankton caused by changes in phytoplankton standing stocks, in this case most likely an increase in small-sized phytoplankton of

Stoichiometric homeostasis is the ability of an organism to keep its body chemical composition constant, despite varying inputs. Stoichiometric homeostasis therefore constrains the metabolic needs of consumers which in turn often feed on resources not matching these requirements. In a broader context, homeostasis also relates to the capacity of an organism to maintain other biological parameters (e.g. body temperature) at a constant level over ambient environmental variations. Unfortunately, there are discrepancies in the literature and ecological and physiological definitions of homeostasis are disparate and partly contradictory. Here, we address this matter by reviewing the existing knowledge considering two distinct groups, regulators and conformers and, based on examples of thermo- and osmoregulation, we propose a new approach to stoichiometric homeostasis, unifying ecological and physiological concepts. We suggest a simple and precise graphical way to identify regulators and conformers: for any given biological parameter (e.g. nutrient stoichiometry, temperature), a sigmoidal relation between internal and external conditions can be observed for conformers while an inverse sigmoidal response is characteristic of regulators. This new definition and method, based on well-studied physiological mechanisms, unifies ecological and physiological approaches and is a useful tool for understanding how organisms are affected by and affect their environment.

Diel vertical migration (DVM) is a common behavior of many pelagic herbivorous zooplankton species in response to predation pressure. It is characterized by a twice daily habitat shift of the zooplankton species: staying in the epilimnion only during night time and migrating down in the crack of dawn in deeper water layers, staying there during the day time. This causes a discontinuous grazing regime and previous studies have shown that the direction and strength of phytoplankton community responses to zooplankton DVM most probably depends on the size of phytoplankton species. To examine the influence of zooplankton DVM on different sized phytoplankton communities, we designed an experiment where we manipulated the size distribution of a natural phytoplankton community a priori in field mesocosms. We investigated the influence of DVM of the cladoceran Daphnia hyalina on two different phytoplankton communities, by the use of deep (10 m) field enclosures. Epilimnetic lake water, containing a summer phytoplankton community, was filtered with two different mesh sizes (11 mm and 64 mm). The 11 mm phytoplankton community (“small”) contained mainly small algal species, while the 64 mm community (“large”) had a wider range of phytoplankton sizes. To simulate zooplankton DVM, D. hyalina were placed in mesh cages that were lowered or raised (“migration”) as dictated by the study design; a “no migration” (representing absence of DVM) treatment was also tested. Phytoplankton abundance was measured using chlorophyll-a and biovolume; size distribution of the algae and nutrient availability was also determined in each treatment. The results indicated that DVM had contrasting effects on the two evaluated phytoplankton communities. Comparison of “migration” and “no migration” zooplankton treatments showed that nutrient availability and total phytoplankton biovolume was higher in (1) “no migration” treatments with phytoplankton communities comprising mainly small algae and (2) “migration” treatments with phytoplankton communities of a broader size spectrum of algae. Hence our study showed two different mechanisms of how zooplankton DVM may influence the phytoplankton community dynamics. Nutrient cycling was an important factor in phytoplankton communities of mainly small algae, whereas the refuge effect was the main driver of phytoplankton dynamics in phytoplankton communities of a large size spectrum of algae.

The issues of food quality and food quantity are crucial for trophic interactions. Although most research has focussed on the primary producer-herbivore link, recent studies have shown that quality effects at the bottom of the food web propagate to higher trophic levels. Negative effects of poor food quality have almost exclusively been demonstrated at higher food quantities. Whether these negative effects have the same impact at low food availability in situations where the majority if not all of the resources are channelled into routine metabolism, is under debate. In this study a tri-trophic food chain was designed, consisting of the algae Rhodomonas salina, the copepod Acartia tonsa and freshly hatched larvae of the European lobster Homarus gammarus. The lobster larvae were presented with food of two different qualities (C:P ratios) and four different quantities to investigate the combined effects of food quality and quantity. Our results show that the quality of food has an impact on the condition of lobster larvae even at very low food quantities. Food with a lower C:P content resulted in higher condition of the lobster larvae regardless of the quantity of food. These interacting effects of food quality and food quantity can have far reaching consequences for ecosystem productivity.