Can tropical freshwater zooplankton graze efficiently on cyanobacteria?

Zooplankton may at times graze cyanobacteria. However, their top-down effects are considered to be low, particularly in tropical regions dominated by small-size grazers that may be unable to consume efficiently filamentous or colonial species. Recently, cyanobacteria blooms were reported in the Senegal River hydrosystem. We conducted feeding experiments to assess the ability of copepods (Pseudodiaptomus hessei and Mesocyclops ogunnus), cladocerans (Moina micrura and Ceriodaphnia cornuta), and rotifers (Brachionus angularis, B. falcatus, and Keratella sp.) to control different cyanobacteria (Cylindrospermopsis raciborskii, Anabaena solitaria, A. flos-aquae, and Microcystis aeruginosa). None of the zooplankton species ingested M. aeruginosa. Mesocyclops ogunnus did not consume any of the cyanobacteria. Both cladocerans consumed the smallest filaments of cyanobacteria, whereas all the rotifers and P. hessei consumed a broader food-size spectrum. The functional feeding responses suggest that the concentration and size of the filaments are not the sole criteria for food consumption. The high zooplankton community grazing rates, estimated by applying the clearance rates measured in the laboratory to the in situ zooplankton abundance, indicate that grazing by zooplankton potentially constitutes an important controlling factor for the filamentous cyanobacteria in the tropics.     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

Differential zooplankton feeding behaviors,selectivities, and community impacts of two planktivorous fishes

Synopsis We examined the feeding behaviors and selectives of two common planktivorous fishes, pumpkinseeds Lepomis gibbosa and fathead minnows Pimephales promelas in the laboratory. Ingestion rates for both pumpkinseeds and fathead minnows feeding on zooplankton increased as a function of fish length. Pumpkinseeds fed on zooplankton strictly as particulate feeders, with preferences increasing as a function of zooplankton body size regardless of taxonomic identity. Preferences were highest for large Daphnia, intermediate for intermediate-sized copepods, and lowest for small Ceriodaphnia. Fathead minnows displayed the ability to use both particulate-feeding and filter-feeding behaviors. Differential preferences tended to reflect both zooplankton size and taxon, being highest for large, slow-swimming Daphnia, intermediate for small Ceriodaphnia, and lowest for faster-swimming copepods. These differences in prey capture behaviors and preferences of the two fishes are reflected in the zooplankton taxonomic composition of small ponds containing each fish type. The crustacean zooplankton assemblages in ponds containing both pumpkinseeds and fathead minnows were dominated by copepods. Cladocerans were rare. In ponds containing pumpkinseeds, but no fathead minnows, cladocerans were abundant, generally accounting for up to 80% of total crustacean zooplankton biomass. These results suggest that the type of planktivore, and not simply the presence or abundance of planktivores in a system, can determine zooplankton community structure.     

Responses of cyanobacteria to herbivorous zooplankton across predator regimes: who mows the bloom?

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Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Cyanobacterial chemical warfare affects zooplankton community composition

1. Toxic algal blooms widely affect our use of water resources both with respect to drinking water and recreation. However, it is not only humans, but also organisms living in freshwater and marine ecosystems that may be affected by algal toxins. 2. In order to assess if cyanobacterial toxins affect the composition of natural zooplankton communities, we quantified the temporal fluctuations in microcystin concentration and zooplankton community composition in six lakes. 3. Microcystin concentrations generally showed a bimodal pattern with peaks in early summer and in autumn, and total zooplankton biomass was negatively correlated with microcystin concentrations. Separating the zooplankton assemblages into finer taxonomic groups revealed that high microcystin concentrations were negatively correlated with Daphnia and calanoid copepods, but positively correlated with small, relatively inefficient phytoplankton feeders, such as cyclopoid copepods, Bosmina and rotifers. 4. In a complementary, mechanistic laboratory experiment using the natural phytoplankton communities from the six lakes, we showed that changes in in situ levels of microcystin were coupled with reduced adult size and diminished juvenile biomass in Daphnia. 5. We argue that in eutrophic lakes, large unselective herbivores, such as Daphnia, are ‘sandwiched’ between high fish predation and toxic food (cyanobacteria). In combination, these two mechanisms may explain why the zooplankton community in eutrophic lakes generally comprise small forms (e.g. rotifers and Bosmina) and selective raptorial feeders, such as cyclopoid copepods, whereas large, unselective herbivores, such as Daphnia, are rare. Hence, this cyanobacterial chemical warfare against herbivores may add to our knowledge on population and community dynamics among zooplankton in eutrophic systems.     

Seasonal dynamics of crustacean zooplankton, heterotrophic nanoflagellates and bacteria in a shallow, eutrophic lake

1. The seasonal development of crustacean zooplankton, heterotrophic nanoflagellates (HNF) and bacteria was examined in Grosser Binnensee, a shallow, eutrophic lake in northern Germany. The grazing impact of Daphnia on bacteria and nanoflagellates was estimated from field data on population abundances and from clearance rates obtained in laboratory experiments. 2. The seasonal succession of zooplankton showed distinct peaks of Daphnia magna, cyclopopid copepods, Bosmina longirostris and Daphnia galeata and D. hynlina. The population dynamics of Dapfinia had the strongest impact on all sestonic components. Daphnia maxima coincided with clearwater phases, and were negatively correlated with particulate organic carbon (POC), HNF and phytoplankton. Bacterial abundance was only slightly affected although daphnids were at times more important as bacterial consumers than HNF, as estimated from measured bacterial clearance rates. Other crustaceans (copepods, Bosmina) were probably of minor importance as grazers of bacteria and nanoplankton. 3. HNF abundance varied from 550 ml^?1 to more than 30000 ml^?1. HNF appeared to be suppressed by daphnids and reached highest densities when copepods dominated the metazooplankton. The variation in HNF abundance was not reflected in the concentration of heterotrophic bacteria, which fluctuated rather irregularly between 5 and 20 ± 10⁶ ml^?1. Long filamentous bacteria which were probably resistant to protozoan grazing, however, appeared parallel to the development of HNF. These bacterial cells, although small in number, could comprise more than 30% of the total bacterial biomass.     

Chytrids enhance Daphnia fitness by selectively retained chytrid-synthesised stearidonic acid and conversion of short-chain to long-chain polyunsaturated fatty acids

1. Chytrid fungal parasites convert dietary energy and essential dietary molecules, such as long-chain (LC) polyunsaturated fatty acids (PUFA), from inedible algal/cyanobacteria hosts into edible zoospores. How the improved biochemical PUFA composition of chytrid-infected diet may extend to zooplankton, linking diet quality to consumer fitness, remains unexplored.
2. Here, we assessed the trophic role of chytrids in supporting dietary energy and PUFA requirements of the crustacean zooplankton Daphnia, when feeding on the filamentous cyanobacterium Planktothrix.
3. Only Daphnia feeding on chytrid-infected Planktothrix reproduced successfully and had significantly higher survival and growth rates compared with Daphnia feeding on the sole Planktothrix diet. While the presence of chytrids resulted in a two-fold increase of carbon ingested by Daphnia, carbon assimilation increased by a factor of four, clearly indicating enhanced carbon transfer efficiency with chytrid presence.
4. Bulk carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotopes did not indicate any treatment-specific dietary effects on Daphnia, nor differences in trophic position among diet sources and the consumer. Compound-specific carbon isotopes of fatty acids (δ¹³C_FA), however, revealed that chytrids bioconverted short-chain to LC-PUFA, making it available for Daphnia. Chytrids synthesised the ω-3 PUFA stearidonic acid de novo, which was selectively retained by Daphnia. Values of δ¹³C_FA demonstrated that Daphnia also bioconverted short-chain to LC-PUFA.
5. We provide isotopic evidence that chytrids improved the dietary provision of LC-PUFA for Daphnia and enhanced their fitness. We argue for the existence of a positive feedback loop between enhanced Daphnia growth and herbivory in response to chytrid-mediated improved diet quality. Chytrids upgrade carbon from the primary producer and facilitate energy and PUFA transfer to primary consumers, potentially also benefitting upper trophic levels of pelagic food webs.

     

Fish-induced changes in zooplankton grazing on phytoplankton and bacterioplankton: a long-term study in shallow hypertrophic Lake Sobygaard

The impact of fish-mediated changes on the structure and grazingof zooplankton on phytoplankton and bacterioplankton was studiedin Lake Søbygaard during the period 1984–92 bymeans of in vitro grazing experiments (¹⁴C-labelled phytoplankton,³H-labelled bacterioplankton) and model predictions. Measuredzooplankton clearance rates ranged from 0–25 ml l^–1h^–1 on phytoplankton to 0–33 ml l^–1 h^–1on bacterioplankton.The highest rates were found during thesummer when Daphnia spp. were dominant. As the phytoplanktonbiomass was substantially greater than that of bacterioplanktonthroughout the study period, ingestion of phytoplankton was26-fold greater than that of bacterioplankton. Multiple regressionanalysis of the experimental data revealed that Daphnia spp.,Bosmina longirostris and Cyclops vicinus, which were the dominantzooplankton, all contributed significantly to the variationin ingestion of phytoplankton, while only Daphnia spp. contributedsignificantly to that of bacterioplankton. Using estimated meanvalues for clearance and ingestion rates for different zooplankters,we calculated zooplankton grazing on phytoplankton and bacterioplanktonon the basis of monitoring data of lake plankton obtained duringa 9 year study period. Summer mean grazing ranged from 2 to4% of phytoplankton production and 2% of bacterioplankton productionto maxima of 53 and 88%, respectively. The grazing percentagedecreased with increasing density of planktivorous fish caughtin August each year using gill nets and shore-line electrofishing.The changes along a gradient of planktivorous fish abundanceseemed highest for bacterioplankton. Accordingly, the percentagecontribution of bacterioplankton to the total ingestion of thetwo carbon sources decreased from a summer mean value of 8%in Daphnia-dominated communities at lower fish density to 0.7–1.1%at high fish density, when cyclopoid copepods or Bosmina androtifers dominated. Likewise, the percentage of phytoplanktonproduction channelled through the bacteria varied, it beinghighest (5–8%) at high fish densities. It is argued thatthe negative impact of zooplankton grazing on bacterioplanktonin shallow lakes is highest at intermediate phosphorus levels,under which conditions Daphnia dominate the zooplankton community.     

Cascading predation effects of Daphnia and copepods on microbial food web components

1. We performed a mesocosm experiment to investigate the structuring and cascading effects of two predominant crustacean mesozooplankton groups on microbial food web components. The natural summer plankton community of a mesotrophic lake was exposed to density gradients of Daphnia and copepods. Regression analysis was used to reveal top–down impacts of mesozooplankton on protists and bacteria after days 9 and 15. 2. Selective grazing by copepods caused a clear trophic cascade via ciliates to nanoplankton. Medium‐sized (20–40 μm) ciliates (mainly Oligotrichida) were particularly negatively affected by copepods whereas nanociliates (mainly Prostomatida) became more abundant. Phototrophic and heterotrophic nanoflagellates increased significantly with increasing copepod biomass, which we interpret as an indirect response to reduced grazing pressure from the medium‐sized ciliates. 3. In Daphnia‐treatments, ciliates of all size classes as well as nanoflagellates were reduced directly but the overall predation effect became most strongly visible after 15 days at higher Daphnia biomass. 4. The response of bacterioplankton involved only modest changes in bacterial biomass and cell‐size distribution along the zooplankton gradients. Increasing zooplankton biomass resulted either in a reduction (with Daphnia) or in an increase (with copepods) of bacterial biovolume, activity and production. Patterns of bacterial diversity, as measured by polymerase chain reaction–denaturing gradient gel electrophoresis (PCR–DGGE), showed no distinct grouping after 9 days, whereas a clear treatment‐coupled similarity clustering occurred after 15 days. 5. The experiment demonstrated that zooplankton‐mediated predatory interactions cascade down to the bacterial level, but also revealed that changes occurred rather slowly in this summer plankton community and were most pronounced with respect to bacterial activity and composition.     

Top‐down regulation of filamentous cyanobacteria varies among a raptorial versus current feeding copepod across multiple prey generations

1. Although considered a key functional trait, little is known about how zooplankton feeding mode affects top‐down regulation of phytoplankton communities. Indeed, copepods are expected to promote the dominance of toxic phytoplankton by selective removal of their edible competitors; however, empirical evidence comparing the effect among calanoid and cyclopoid copepods is lacking.
2. We compared the top‐down effects of two copepods with contrasting feeding modes—the calanoid Notodiaptomus iheringi (current feeder) and the cyclopoid Thermocyclops decipiens (ambush feeder) — on the relative and absolute biomass of the filamentous cyanobacterium Raphidiopsis raciborskii co‐cultured with the nutritious eukaryotic phytoplankton Cryptomonas obovata in a week‐long laboratory assay.
3. The current feeder had a stronger top‐down effect on the biomass of both prey throughout the experiment, with mass‐specific clearance rates 3–5× higher than ambush feeder. By the end of the experiment, the current feeder significantly reduced cyanobacteria biomass compared to controls while the ambush feeder did not. During the week‐long experiment, the current feeder switched from grazing on edible prey to cyanobacteria as the former became less abundant.
4. Contrary to expectation, neither of the copepod species promoted cyanobacterial dominance by the end of the experiment. This is because both grazers, but especially the current feeder, initially increased but subsequently decreased the relative contribution of cyanobacteria to total phytoplankton biomass. Moreover, both copepods decreased the length of cyanobacteria filaments by c. 70%
5. Current feeders can switch from edible prey to cyanobacteria when the abundance of shortened filaments surpasses the abundance of edible prey. While top‐down regulation of phytoplankton can be stronger for current feeding copepods, ambush feeding copepods can have a significant role during blooms by shortening cyanobacterial filaments. Hence, the broader role of contrasting copepod feeding traits on phytoplankton communities merits further study.

     

Effects of UV-B irradiated algae on zooplankton grazing

We tested the effects of UV-B stressed algae on grazing rates of zooplankton. Four algal species (Chlamydomonas reinhardtii, Cryptomonas sp., Scenedesmus obliquus and Microcystis aeruginosa) were used as food and fed to three zooplankton species (Daphnia galeata, Bosmina longirostris and Brachionus calyciflorus), representing different taxonomic groups. The phytoplankton species were cultured under PAR conditions, and under PAR supplemented with UV-B radiation at two intensities (0.3 W m^–2 and 0.7 W m^–2, 6 hours per day). Ingestion and incorporation experiments were performed at two food levels (0.1 and 1.0 mg C l^–1) using radiotracer techniques. The effect of food concentration on ingestion and incorporation rate was significant for all three zooplankton species, but the effect of UV-B radiation was more complex. The reactions of the zooplankton species to UV-B stressed algae were different. UV-B stressed algae did not affect Daphnia grazing rates. For Bosmina the rates increased when feeding on UV-B stressed Microcystis and decreased when feeding on UV-B stressed Chlamydomonas, compared with non-stressed algae. Brachionus grazing rates were increased when feeding on UV-B stressed Cryptomonas and UV-B stressed Scenedesmus, and decreased when feeding on UV-B stressed Microcystis, compared with non-stressed algae. These results suggest that on a short time scale UV-B radiation may result in increased grazing rates of zooplankton, but also in decreased grazing rates. Long term effects of UV-B radiation on phytoplankton and zooplankton communities are therefore difficult to predict.     

Zooplankton community size structure and taxonomic composition affects size-selective grazing in natural communities

The body size of an individual zooplankton is well related to its grazing rate and to the range of particle sizes it can ingest, and since cladocerans and copepods feed differently, they follow different relationships. Based on these general patterns in individual organisms, we tested whether the size structure and taxonomic composition of more complex natural zooplankton communities are related to their in situ grazing rate and to the range of algal sizes they graze. We compared community grazing rates on individual algal taxa in two communities dominated by small cladocerans, three communities dominated by large cladocerans and three copepod-dominated communities. Small algae were usually grazed most intensively, but grazing rates were poorly related to algal size alone. The range in size of grazed algae increased with increasing mean zooplankton body size, but differed systematically with their taxonomic composition. Communities dominated by Ceriodaphnia or Holopedium grazed a narrower size range of algae [maximum greatest axial length dimension (GALD)=16–36 μm)] than communities with large biomasses of Bosmina or Daphnia (maximum GALD=28–78 μm). Copepod-dominated communities followed the same general relationship as cladocerans. Daphnia-dominated communities grazed the broadest range of algal sizes, and their total grazing rates were up to 2.4 times their grazing rates on small (<35 μm) “highly edible” algae, a difference of similar magnitude to those found in successful trophic cascade biomanipulations. Received: 31 March 1998 / Accepted: 19 October 1998     

Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

A comparison of the trophic transfer of fatty acids in freshwater plankton by cladocerans and calanoid copepods

1. Analyses of zooplankton fatty acid (FA) composition in laboratory experiments and samples collected from lakes in New Zealand spanning a wide gradient of productivity were used to assess the extent to which FAs might infer their diet. We used the cladocerans, Daphnia and Ceriodaphnia, and the calanoid copepod, Boeckella, as test organisms, and monocultures of cryptophytes, chlorophytes and cyanobacteria as food. Based on reproductive success, cryptophytes were the highest food quality, chlorophytes were intermediate and cyanobacteria the poorest. 2. Several FA groups were highly correlated between zooplankton and their diets. They were monounsaturated fatty acids (MUFAs), and ω3 and ω6 polyunsaturated fatty acids (PUFAs) for cladocerans, and saturated fatty acids (SAFAs) and ω3 PUFAs for copepods. Several FAs varied significantly less in the zooplankton than in their monoculture diets, e.g. MUFAs in Daphnia, and ω3 and ω6 PUFAs in Ceriodaphnia, despite clear dietary dependency for these FAs. 3. Zooplankton collected from lakes in New Zealand had more eicosapentaenoic acid (EPA) (Daphnia), more highly unsaturated ω3 and ω6 FAs (C₂₀, C₂₂; Daphnia, Ceriodaphnia, Boeckella) and less ω3 C₁₈ PUFAs (Daphnia, Ceriodaphnia, Boeckella) and ω6 C₁₈ PUFAs (Daphnia, Ceriodaphnia) than measured in the same species reared on phytoplankton in the laboratory. 4. Analyses of FA composition of seston and freshwater zooplankton globally showed that, in general, zooplankton had a significantly higher proportion of arachidonic acid and EPA than seston, and copepods also had a higher percentage of docosahexaenoic acid than seston. 5. These results suggest that zooplankton selectively incorporate the most physiologically important FAs. This could be a consequence of preferential assimilation, selective feeding on more nutritious cells or locating and feeding within higher food quality food patches.     

Seasonal patterns of microcystin-producing and non-producing Planktothrix rubescens genotypes in a deep pre-alpine lake

The filamentous cyanobacterium Planktothrix rubescens produces secondary metabolites called microcystins (MC) that are potent toxins for most eukaryotes, including zooplankton grazers, cattle and humans. P. rubescens occurs in many deep and thermally stratified lakes throughout Europe. In Lake Zurich (Switzerland), it re-appeared in the 1970s concomitant with decreasing eutrophication. Since then, P. rubescens has become the dominant species in this major drinking water reservoir, where it forms massive metalimnetic blooms during late summer. These cyanobacteria harbor subpopulations of non-MC producers, but little is known about the environmental factors affecting the success of such genotypes. The non-MC-producing subpopulation of P. rubescens was studied using a quantitative real-time PCR (qPCR) assay on the MC synthetase (mcy) gene cluster that targets a deletion on the mcyH and mcyA genes, which inactivates MC biosynthesis. Two complementary qPCR assays were used to assess the total population abundance (based on the 16S rDNA gene) and the mcy gene copy number (based on a conserved region in the adenylation domain of the mcyB gene). The objective was to evaluate the seasonal patterns of the share of non-MC-producing filaments in the total P. rubescens population. The mcyHA mutants were present in low proportions (up to 14%) throughout the year. Their highest relative abundances occurred during the winter mixis, when total concentrations of P. rubescens were minimal. The MC deficient mutants seemed to better survive in sparse populations, possibly because of lower grazing pressure and a consequently reduced need for MC-mediated protection. Alternatively, the mutants might cope better with the sub-optimal, stressful pressure and light conditions during the winter mixis. Altogether, our results suggest that subtle trade-offs might seasonally determine the proportions of non-MC producers within P. rubescens populations.     

Zooplankton interactions in an enclosure experiment: insights from stable isotope analyses

1. Density gradients of cladocerans and copepods were generated in an enclosure experiment to compare the impact on the plankton of a filter feeder (Daphnia hyalina × galeata) with that of more selective feeders (calanoid and cyclopoid copepods). The experiment was conducted in situ over 25 days during spring in a mesotrophic lake, Schöhsee, Germany. 2. The plankton community was monitored regularly. Daphniids were able to graze on the phytoplankton present, which mainly consisted of small (<1000 μm³) species, whereas copepods did not show any impact on algae. 3. At the end of the experiment, Daphnia and remaining cyclopoid copepods were harvested and sorted manually, prior to analyses for stable isotopes of carbon and nitrogen. Daphniids from mesocosms stocked purely with differing densities of Daphnia showed little variability in stable isotope values, whereas those that thrived in enclosure bags together with copepods exhibited lower δ¹³C values. 4. The change in Daphniaδ¹³C indicates a change of food sources, modified by the presence of the copepods: the higher the mean abundance of copepods in the enclosures, the more ¹³C‐depleted the daphniids. Increasing abundance of high nucleic acid (HNA) bacteria in the copepod bags may account for the trend in Daphniaδ¹³C via increased grazing on the bacteria themselves, or via grazing on phytoplankton utilising isotopically light CO₂ from respiratory release. 5. Cyclopoid copepod stable isotope signatures were related to Daphnia and copepod abundances in copepod bags, suggesting that cyclopoids preyed on the available zooplankton.     

Zooplankton grazing on Phaeocystis: a quantitative review and future challenges

The worldwide colony-forming haptophyte phytoplankton Phaeocystis spp. are key organisms in trophic and biogeochemical processes in the ocean. Many organisms from protists to fish ingest cells and/or colonies of Phaeocystis. Reports on specific mortality of Phaeocystis in natural plankton or mixed prey due to grazing by zooplankton, especially protozooplankton, are still limited. Reported feeding rates vary widely for both crustaceans and protists feeding on even the same Phaeocystis types and sizes. Quantitative analysis of available data showed that: (1) laboratory-derived crustacean grazing rates on monocultures of Phaeocystis may have been overestimated compared to feeding in natural plankton communities, and should be treated with caution; (2) formation of colonies by P. globosa appeared to reduce predation by small copepods (e.g., Acartia, Pseudocalanus, Temora and Centropages), whereas large copepods (e.g., Calanus spp.) were able to feed on colonies of Phaeocystis pouchetii; (3) physiological differences between different growth states, species, strains, cell types, and laboratory culture versus natural assemblages may explain most of the variations in reported feeding rates; (4) chemical signaling between predator and prey may be a major factor controlling grazing on Phaeocystis; (5) it is unclear to what extent different zooplankton, especially protozooplankton, feed on the different life forms of Phaeocystis in situ. To better understand the mechanisms controlling zooplankton grazing in situ, future studies should aim at quantifying specific feeding rates on different Phaeocystis species, strains, cell types, prey sizes and growth states, and account for chemical signaling between the predator and prey. Recently developed molecular tools are promising approaches to achieve this goal in the future.     

Effects of Daphnia exudates and sodium octyl sulphates on filament morphology and cell wall thickness of Aphanizomenon gracile (Nostocales), Cylindrospermopsis raciborskii (Nostocales) and Planktothrix agardhii (Oscillatoriales)

Grazing is recognized as one of the selective factors shaping the morphology and physiology of cyanobacteria. A recent study has shown that the filamentous cyanobacterium Aphanizomenon gracile strain SAG 31.79 thickened in the presence of Daphnia (Cladocera) and its exudates. The aims of our study were: (1) to determine whether this type of response to Daphnia cues is common for other strains of A. gracile, and other species of filamentous cyanobacteria, (2) to test whether the response is due to nutrients recycled by Daphnia, or kairomone induced, and (3) whether it is related to toxin production. Prior to the experiment, cyanobacterial strains were inspected using chromatographic methods for the presence of two toxins, cylindrospermopsin (CYN) and three homologues of microcystin (MC-RR, MC-YR, MC-LR). HPLC analyses showed that all strains were free of cylindrospermopsin, whereas microcystins were detected only in one strain (Planktothrix agardhii). We then tested whether Daphnia exudates can cause thickening of cyanobacterial filaments, which would suggest the morphological changes in cyanobacterial filaments are caused by recycled nutrients. Cyanobacteria were also exposed to sodium octyl sulphate (a commercially available Daphnia kairomone). Transmission electron microscopy (TEM) was used to check whether Daphnia exudates and sodium octyl sulphate trigger thickening of cyanobacterial cell walls, which would be a defence mechanism against grazing. The TEM analysis revealed no significant effect of either Daphnia exudates or kairomone (sodium octyl sulphate) on the cell wall thickness of cyanobacteria. However, our study showed that Daphnia exudates triggered filament thickening in nostocalean cyanobacteria, while filaments of the oscillatorialean strain P. agardhii did not show this response. It was also demonstrated that sodium octyl sulphate alone can also cause filament thickening, which suggests that this might be a specific defence response to the presence of grazers.     

Effects of Cell-Bound Microcystins on Survival and Feeding of Daphnia spp.

The influence of cell-bound microcystins on the survival time and feeding rates of six Daphnia clones belonging to five common species was studied. To do this, the effects of the microcystin-producing Microcystis strain PCC7806 and its mutant, which has been genetically engineered to knock out microcystin synthesis, were compared. Additionally, the relationship between microcystin ingestion rate by the Daphnia clones and Daphnia survival time was analyzed. Microcystins ingested with Microcystis cells were poisonous to all Daphnia clones tested. The median survival time of the animals was closely correlated to their microcystin ingestion rate. It was therefore suggested that differences in survival among Daphnia clones were due to variations in microcystin intake rather than due to differences in susceptibility to the toxins. The correlation between median survival time and microcystin ingestion rate could be described by a reciprocal power function. Feeding experiments showed that, independent of the occurrence of microcystins, cells of wild-type PCC7806 and its mutant are able to inhibit the feeding activity of Daphnia. Both variants of PCC7806 were thus ingested at low rates. In summary, our findings strongly suggest that (i) sensitivity to the toxic effect of cell-bound microcystins is typical for Daphnia spp., (ii) Daphnia spp. and clones may have a comparable sensitivity to microcystins ingested with food particles, (iii) Daphnia spp. may be unable to distinguish between microcystin-producing and -lacking cells, and (iv) the strength of the toxic effect can be predicted from the microcystin ingestion rate of the animals.