FO-spectra of chlorophyll fluorescence for the determination of zooplankton grazing

In the PHYTO-PAM phytoplankton analyzer the minimal fluorescence of dark-adapted samples (F₀) was assessed, which gives direct information on the chlorophyll-a content. Clearance rates (CR) of Daphnia and Brachionus were calculated from a decrease in chlorophyll-a concentration using the PHYTO-PAM fluorometer for non-sacrificial sampling of chlorophyll-a. Clearance rates of Daphnia were measured and compared with those based on the cell-counts method using an electronic particle counter (Coulter counter). Chlorophyll fluorescence-based CR for Daphnia magna were very strongly correlated with Coulter-based CR, signifying the potential suitability of the PHYTO-PAM in grazing experiments. A procedure for determination of rotifer clearance rates was developed and the effects of rotifer density, duration of the grazing period, and food concentration on CR were investigated. Between 10 and 30 rotifers in 2.5 ml food suspension (i.e. 4–12 rotifers per ml) appeared optimal for calculating CR. The application of the deconvolution of F₀-spectra in food selectivity experiments was evaluated using various mixtures of the green alga Scenedesmus obliquus and the cyanobacterium Microcystis aeruginosa fed to Brachionus. CR for Brachionus on M. aeruginosawere lower than on S. obliquusbut this was not caused by toxicity, because no mortality was observed. The higher CR on Scenedesmus than on Microcystis in the mixtures suggested selectivity. The importance of digital suppression of background fluorescence is highlighted in additional experiments with Daphnia feeding on mixtures of Microcystis and Scenedesmus, or on Microcystis alone. Without background correction of filtered samples, negative clearance rates were obtained for the `blue' Microcystis signal. Soluble fluorescing compounds of cyanobacterial origin, phycocyanin, were released from the Daphniaand contributed 40% to the overall-fluorescence. Deconvolution of F₀-spectra for the determination of chlorophyll-a using the PHYTO-PAM appears to be a suitable tool for determination of rotifer CR even at very low food concentrations. A drawback of the method is that rather high rotifer densities are required. The required grazing period, however, is shorter than for cell-count methods, the method is sensitive, clearance rates can be measured at low food concentrations (< 0.1 mg C l^–1) and information on selective feeding can be obtained.     

Type-3 functional response in limnetic suspension-feeders,as demonstrated by in situ grazing rates

Field-measured grazing rates (ml/animal/d) of cladocerans (mostly daphniids) and diaptomids were assembled from various published studies and plotted as a function of corresponding phytoplankton concentration (μg l⁻¹ f.w.). Filtering rates of both zooplankton groups initially increased with seston concentration until maximal grazing rates were observed at approximately 4 × 10² and 1 × 10² μg l⁻¹ for cladocerans and copepods, respectively; at higher algal concentrations, filtering rates of both declined as a function of food concentration. The shape of these curves are most consistent with Holling's (1966) Type 3 functional response. We found little support for the Type 3 functional response in published laboratory studies of Daphnia; most investigators report either a Type 1 or Type 2 response. The one study in which the Type 3 response was observed involved experiments where animals were acclimated at low food concentrations for 24 h, whereas those studies associated with response Types 1 or 2 had acclimation periods of only 1 to 3 h. We therefore assembled relevant data from the literature to examine the effect of acclimation period on the feeding rates of Daphnia at low food concentrations. In the absence of any acclimation, animals filtered at extremely low rates. After 2 h of acclimation, however, filtering rates increased 4 to 5-fold but declined again with longer durations; after > 70 h of pre-conditioning, filtering rates were almost as low as they had been with no acclimation. We also found little support for the Type 3 functional response in published studies of copepods. The only study associated with a Type 3 response involved a marine copepod that had been subjected to a starvation period of 48 h; however, an analysis of the effects of acclimation period did not yield conclusive evidence that filtering rates of freshwater copepods (Diaptomus and Eudiaptomus) decrease significantly with acclimation duration. The low filtering rates associated with long acclimation periods in laboratory experiments appears to be a direct result of animals becoming emaciated from prolonged exposure to low food concentrations, a situation which renders them incapable of high filtering rates. This may explain the Type 3 functional response for field cladocerans, since zooplankton in food-limiting situations are constantly exposed to low food concentrations, and would therefore have low body carbon and consequently less energy to filter-feed. We cannot, however, use this to explain the Type 3 response for field diaptomids, since copepods in the laboratory did not appear to lose body carbon even after 72 h of feeding at very low food levels, and there was inconclusive evidence that either Diaptomus or Eudiaptomus decrease their filtering rates with acclimation period. Although Incipient Limiting Concentrations (ILC) for Daphnia ranged from 1 to 8.5 × 10³ μg 1⁻¹, more than half of these fell between 1 and 3 × 10³ μg l⁻¹, bracketing the value of 2.7 × 10² μg l⁻¹ for field cladocerans. There was, however, a great deal of variation in reported maximum ingestion rates (MIR), maximum filtering rates (MFR) and ILC values for Daphnia magna. ILC values from the few laboratory studies of freshwater copepods ranged between 0.5 to 2.8 × 10³ μg 1⁻¹, and was higher than the ILC value of approximately 0.2 × 10³ μg l⁻¹ calculated for field populations of D. minutus. Generally, there was considerable agreement among laboratory studies regarding the shape of grazing-rate and ingestion-rate curves when data were converted to similar units and presented on standardized scales.     

Impact of different developmental stages of <Emphasis Type="Italic">Daphnia</Emphasis> <Emphasis Type="Italic">magna</Emphasis> (Straus) on the plankton community under different trophic conditions

In situ 24-h incubation experiments were performed to analyse the grazing effects of Daphnia magna on a planktonic microbial community. Three field grazing experiments under different nutrient concentrations were carried out on treated effluents of a wastewater treatment plant. The grazing effects of three different D. magna size classes (small (0.6–1.6 mm), medium (1.7–2.5 mm) and large individuals (2.6–3.7 mm)) were compared. The different sizes classes had similar effects on the plankton community. However, our results showed big differences in effects among experiments. Our findings suggest that in spite of D. magna’s non-selective feeding behaviour and the fact that different developmental stages (i.e. its size) had similar effects on the microbial planktonic community, these effects can differ according to the initial structure and composition of the community and the resulting cascading trophic interactions. Moreover, D. magna effects can be direct through grazing (as is the case with ciliates), or indirect through trophic cascade interactions (as is the case with bacteria).     

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     

Does excess dietary carbon affect respiration of <Emphasis Type="Italic">Daphnia</Emphasis>?

Like many invertebrate herbivores, Daphnia frequently face diets with excess carbon (C) relative to elements like phosphorus (P), and with limited ability to store C-rich compounds. To cope with this relative surplus of C they may either regulate the net uptake of C or dispose of excess assimilated C via increased release of dissolved organic carbon or CO₂. Here we investigate whether juvenile Daphnia magna use respiration as a means of stoichiometrically regulating excess C. Growth rate and respiration were measured under different algal food qualities (P-replete and P-depleted algae). Growth rate was strongly reduced by P-depleted food, implying a stoichiometric disposal of excess ingested C. Respiration rates of feeding animals were measured after short- (0.5 h), medium- (12 h) and long- (five days) term acclimation to P-limited food. The respiration rates of animals during active feeding were not affected by the acclimation period per se, whereas food quality had a significant effect; respiration rates of feeding animals increased slightly in individuals receiving low-P food under all acclimation regimes. Respiration was also measured on nonfeeding and fasting animals that had been acclimated for five days to P-limited food. Respiration rates of these animals were strongly affected by feeding conditions but not by food quality; feeding individuals had higher respiration rates than those deprived of food, which again had higher respiration than fasting animals. Although animals grown on low-P food had strongly reduced growth and thus were expected to have decreased respiration rates due to reduced growth-related costs, this seems to be canceled out by increased stoichiometric respiration under P-deficiency. These results indicate that D. magna partly releases excess C as CO₂, but other means of stoichiometric regulation most likely add to this.     

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.     

Evolutionary history of Daphnia drives divergence in grazing selectivity and alters temporal community dynamics of producers

Consumers with different seasonal life histories encounter different communities of producers during specific seasonal phases. If consumers evolve to prefer the producers that they encounter, then consumers may reciprocally influence the temporal composition of producer communities. Here, we study the keystone consumer Daphnia ambigua, whose seasonal life history has diverged due to intraspecific predator divergence across lakes of New England. We ask whether grazing preferences of Daphnia have diverged also and test whether any grazing differences influence temporal composition patterns of producers. We reared clonal populations of Daphnia from natural populations representing the two diverged life history types for multiple generations. We conducted short‐term (24 hr) and long‐term (27 days) grazing experiments in equal polycultures consisting of three diatom and two green algae species, treated with no consumer, Daphnia from lakes with anadromous alewife, or from lakes with landlocked alewife. After 24 hr, life history and grazing preference divergence in Daphnia ambigua drove significant differences in producer composition. However, those differences disappeared at the end of the 27‐day experiment. Our results illustrate that, despite potentially more complex long‐term dynamics, a multitrophic cascade of evolutionary divergence from a predator can influence temporal community dynamics at the producer level.     

Growth of Daphnia longispina L. in a polyhumic lake under various availabilities of algal,bacterial and detrital food

The availability and importance of food sources for growth of Daphnia longispina L. from a highly coloured fishless lake with anoxic hypolimnion were assessed by combining in situ and laboratory experiments. In in situ experiments populations were enclosed in tubes with natural temperature stratification and with or without anoxic hypolimnion. In the laboratory experiments the importance of food source (littoral zone vs pelagic epilimnion) was assessed by enclosing moss thalli and a natural zooplankton population in a large-scale flow-through system supplying food for experimental Daphnia. Growth of juveniles of Daphnia in epilimnetic water was determined in batch culture experiments and the importance of increasing concentrations of bacteria and algae for their growth and development was investigated with a small-scale flow-through system. Access to the anoxic hypolimnion enhanced the growth of Daphnia by 23–24%. Growth rates in the tubes with anoxic hypolimnion were 0.36 and 0.16 d^–1 in July and August respectively. In tubes without anoxia the corresponding values were 0.29 and 0.13. In batch-cultures the highest growth rate determined was 0.16 and the overall rates were lower than in in situ experiments. In batch culture Daphnia was able to grow in darkness for 10 days with a rate of 0.16. In the large-scale flow-through system Daphnia population fed with littoral water reproduced well despite the low concentration of algae and increased its number by a factor of c. 32 in 10 days. However, the animals were small and net production of Daphnia population thus lower under the littoral influence than in the control treatment. Population could survive and grew slowly on pelagial water processed by a natural zooplankton community and with very little algae left. It is thus possible that bacteria serve as a life-support system enabling the population survival over periods of algal shortage. Small-scale flow-through experiments revealed that Daphnia longispina is able to mature and reproduce on a bacterial diet if the food concentration is high enough and Daphnia on bacterial food could achieve growth rates similar to those on an algal diet. The threshold food level for Daphnia longispina was estimated to be c. 18.5 g C 1^–1. Detrital material is of limited value in nutrition of Daphnia even in a lake where more than 75% of carbon is bound in particulate detritus.     

Effects of Microcystis aeruginosa on interference competition between Daphnia pulex and Keratella cochlearis

Laboratory experiments tested the hypothesis that a toxic strain of Microcystis aeruginosa decreases the ability of Daphnia pulex to interfere with Keratella cochlearis. To test a variety of conditions, juvenile and adult Daphnia were exposed to the cyanobacterium for different time periods prior to, and during the experiments. Adult Daphnia not only suppressed rotifers over successive two-day intervals, but also had a significant impact within a 24-hour period. However, the presence of Microcystis (5 × 105 cells ml^–1) decreased the Daphnia effect in both experiments. Although juvenile Daphnia also significantly suppressed Keratella population growth, the presence of Microcystis (10⁵ and 5 × 10⁵ cells ml^–1) caused a significant reduction in daphniid body size and decreased the ability of both nonacclimated and acclimated daphniids to suppress rotifers. Keratella inhalation and mortality are positively correlated with filtering rates and body size of Daphnia. Therefore, the feeding rates and size structure of a Daphnia population will determine its potential to interfere with vulnerable rotifers. In all experiments the presence of Microcystis significantly decreased the ability of Daphnia to interfere with this rotifer despite the fact that Keratella was also inhibited. In the field this effect might be augmented if Microcystis colonies are more easily ingested by cladocerans than by the rotifers.     

High acclimation potential in floating Macrocystis pyrifera to abiotic conditions even under grazing pressure – a field study

The persistence of floating seaweeds, which depends on abiotic conditions but also herbivory, had previously been mostly tested in outdoor mesocosm experiments. In order to investigate if the obtained mesocosm results of high seaweed persistence under natural environmental conditions and under grazing pressure can be extrapolated to field situations, we conducted in situ experiments. During two summers (2007 and 2008), Macrocystis pyrifera was tethered (for 14 d) to lines in the presence and absence of the amphipod Peramphithoe femorata at three sites (Iquique, Coquimbo, Calfuco). We hypothesized that grazing damage and seaweed persistence vary among sites due to different abiotic factors. By incubating the sporophytes in mesh bags, we were either able to isolate (grazing) or exclude (control) amphipods. To test for a mesh bag artifact, a set of sporophytes was incubated without mesh bags (natural). Mesh bags used to exclude herbivores influenced sporophyte growth and physiological performance. The chlorophyll a (Chl a) content depended largely on grazers and grazed sporophytes grew less than natural and control sporophytes within the two summers. A decrease in Chl a content was found for the sites with the highest prevailing irradiances and temperatures, suggesting an efficient acclimation to these sea surface conditions. Our field‐based results of sporophyte acclimation ability even under grazing pressure widely align with previous mesocosm results. We conclude that M. pyrifera and other temperate floating seaweeds can function as long‐distance dispersal vectors even with hitchhiking mesoherbivores.     

Contrasting ‘Top-Down’ Effects of Crustacean Zooplankton Grazing on Bacteria and Phytoflagellates

The combined effects of grazing and nutrient regeneration by Daphnia and Eudiaptomus on the growth of Rhodomonas and heterotrophic bacteria was assessed experimentally. The responses of Rhodomonas and bacteria to the grazers were measured as net specific growth rate over the entire experimental periods, as well as production and specific production at the end of the experiments. Both zooplankton species had a negative effect on Rhodomonas net specific growth rate due to grazing and a positive effect on specific primary production due to nutrient regeneration. Daphnia had no effect on bacterial net specific growth rate, bacterial production or specific bacterial production in one of two experiments. In the other experiment, however, both bacterial growth rate and production decreased as a result of grazing. Furthermore, Daphnia had a negative effect on specific bacterial production, but Eudiaptomus had a positive effect on all bacterial parameters due to nutrient regeneration, probably of phosphorus. Positive effects of copepods on bacterial growth has previously been attributed to trophic cascades via protozoa. However, the present experiments show that regeneration of nutrients, especially phosphorus, may account for a large part of the stimulation of bacterial growth.     

The effect of different zooplankton grazing patterns resulting from diel vertical migration on phytoplankton growth and composition: a laboratory experiment

Diel vertical migration (DVM) of herbivorous zooplankton is a widespread behavioural phenomenon in freshwater ecosystems. So far only little attention has been paid to the impact of DVM on the phytoplankton community in the epilimnion. Some theoretical models predict that algal population growth in the epilimnion should depend on the herbivores migration and grazing patterns: even if migrating zooplankton consume the same total amount of algae per day in the epilimnion as non-migrating zooplankton, nocturnal grazing should result in enhanced algal growth and favour algal species with high intrinsic growth rates over species with lower intrinsic growth rates. To test these hypotheses we performed experiments in which several algal species were confronted with different feeding regimes of Daphnia. In the experiments algal growth did not only depend on the absolute time of grazing but was comparatively higher when grazing took place only during the night, even when the grazing pressure was the same. Furthermore, algal species with higher intrinsic growth rates had higher advantages when being grazed upon only discontinuously during the night than algal species with a smaller intrinsic growth rate. The grazing pattern itself was an important factor for relative algal performance.     

Biomanipulation and food-web dynamics — the importance of seasonal stability

Responses of phytoplankton biomass were monitored in pelagic enclosures subjected to manipulations with nutrients (+N/P), planktivore roach (Rutilus rutilus) and large grazers (Daphnia) in 18 bags during spring, summer and autumn in mesotrophic Lake Gjersjøen. In general, the seasonal effects on phytoplankton biomass were more marked than the effects of biomanipulation. Primary top-down effects of fish on zooplankton were conspicuous in all bags, whereas control of phytoplankton growth by grazing was observed only in the nutrient-limited summer situation. The effect of nutrient additions was pronounced in summer, less in spring and autumn; additions of fish gave the most pronounced effect in spring. The phytoplankton/zooplankton biomass ratio remained high (10–100) in bags with fish, with the highest ratios in combination with fertilization. The ratio decreased in bags without fish to<2 in most bags, but a real grazing control was only observed in bags with addition ofDaphnia. No direct grazing effects could be observed on the absolute or relative biomass of cyanobacteria (mainlyOscillatoria agardhii). The share of cyanobacteria in total phytoplankton biomass was lowest in summer (7–26%), higher in spring (39–63%) and more than 90% in the autumn experiment. The development of the cyanobacterial biomass was rather synchronous in all bags in all the three experiments. A high biomass ofDaphnia gave no increase in the pool of dissolved nutrients in spring, a slight increase in summer and a pronounced increase in autumn. While a strong decrease in the P/C-cell quota of the phytoplankton was observed from spring to autumn, no effect of grazing or nutrient release could be related to this P/C-status. The experiments indicate that such systems, with high and stable densities of inedible cyanobacteria, are rather insensitive to short-term (3–4 weeks) biomanipulation efforts. This is supported by observations on the long-term development of the lake.

     

<Emphasis Type="Italic">Daphnia</Emphasis> distribution in Andean Patagonian lakes: effect of low food quality and fish predation

Food quality in terms of carbon (C):phosphorus (P) ratio can constrain the success of highly demanding P herbivores as Daphnia. North Andean Patagonian lakes are ultraoligotrophic with low nutrient concentrations and well-developed euphotic zones. We investigated the distribution of the large Daphnia commutata in relation with food quality (sestonic C:P ratio) and predation risk in these lakes. The predation risk was estimated based on the fish species present and their relative eye diameter and transparency of the lake. The C:P ratios in the lakes were high, varying from 350 to >1,200. The lakes with D. commutata had significantly lower C:P ratio than those without these daphnids. On the other hand, those lakes where Daphnia is present have the lower predation risk than those were Daphnia is absent. In addition, we carried out growth experiments with neonates and natural seston of three lakes with different C:P ratio. The growth rates were inversely related with C:P of the food. Food quality and predation risk together determined the success or failure of large Daphnia populations in these Andean clear ultraoligotrophic lakes.     

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.     

Relative impacts of Daphnia grazing and direct stimulation by fish on phytoplankton abundance in mesocosm communities

• 1 Planktivorous fish were hypothesised to influence the abundance of algal biomass in lakes by changing zooplankton grazing, affecting zooplankton nutrient recycling and by direct recycling of nutrients to phytoplankton. The relative roles of direct fish effects vs. zooplankton grazing were tested in mesocosm experiments by adding to natural communities large grazing zooplankton (Daphnia carinata) and small planktivorous fish (mosquitofish or juveniles of Australian golden perch).
• 2 The addition of Daphnia to natural communities reduced the numbers of all phytoplankton less than 30 µm in size, but did not affect total biomass of phytoplankton as large Volvox colonies predominated.
• 3 The addition of Daphnia also reduced the abundance of some small (Moina, Bosmina, Keratella) and large (adult Boeckella) zooplankton, suggesting competitive interactions within zooplankton.
• 4 The addition of mosquitofish to communities containing Daphnia further reduced the abundance of some small zooplankton (Moina, Keratella), but increased the numbers of Daphnia and adult Boeckella. In spite of the likely increase in grazing due to Daphnia, the abundance of total phytoplankton and dominant alga Volvox did not decline in the presence of mosquitofish but was maintained at a significantly higher level than in control.
• 5 The addition of juveniles of golden perch to communities containing Daphnia reduced the abundance of small zooplankton (Moina), increased the abundance of large zooplankton (adult Boeckella) but had no significant effect on Daphnia and total phytoplankton abundance.
• 6 The results of the present study suggest that some planktivorous fish can promote the growth of phytoplankton in a direct way, probably by recycling nutrients, and even in the presence of large grazers. However, the manifestation of the direct effect of fish can vary with fish species.

     

Density-dependent predation ofChaoborus flavicans onDaphnia longispina in a small lake: the effect of prey size

Functional response curves of fourth instar larvae ofChaoborus flavicans preying on two size classes ofDaphnia longispina were examined throughout three summer seasons in a small forest lake. Data for each size class were fitted to Holling's disc equation. The parametersa (attack rate) andTh (handling time) were calculated for each prey size from these curves. Attack rate was greater and handling time was shorter for small (0.77 mm) than for large (1.82 mm)Daphnia. In 1:1 mixture of these prey size classes the predation rates ofChaoborus on smallDaphnia at prey densities above 20 l^–1 were greater than predicted from the single size-class experiments. The observed predation rates on largeDaphnia were lower than predicted at all prey densities. Since both single size-class and two size-class experiments were run during the same period of time the difference in observed and predicted predation rates could not be attributed to seasonal changes in prey preference ofChaoborus larvae. In experiments with a concentrated mixture of lake zooplankton (dominated byD. longispina)Chaoborus preference forDaphnia decreased as prey body size increased. There was no obvious correlation between selectivity coefficients and size-frequency distributions ofDaphnia. When medium-sizedDaphnia were omitted from calculations the preference of small over large prey did not differ significantly from the predictions of the single size-class model.     

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.     

Elemental composition (C:N:P) and growth rates of bacteria and Rhodomonas grazed by Daphnia

The elemental composition and growth rate of Rhodomonas andheterotrophic bacteria were studied in batch cultures in thepresence and absence of Daphnia and at two different levelsof phosphorus limitation. The elemental content of single cellswas measured with X-ray microanalysis. Simultaneously, dilutionexperiments were performed in order to estimate grazing losses,growth rates and dominant nutrient sources for bacteria andRhodomonas. The phosphorus:carbon (P:C) ratios of the bacteriawere generally higher in the experiment with the stronger Plimitation of the system. High P:C ratios were taken as an indicationthat bacteria were carbon limited. The presence of Daphnia resultedin a further increase in bacterial P:C ratios and increasedspecific growth rates. Thus, grazing increased the availabilityboth of inorganic nutrients and organic substrates, stimulatingthe growth of the bacteria. P:C ratios of Rhodomonas decreasedwith increasing P limitation of the system. Only at strong Plimitation did the presence of Daphnia result in increased P:Cratios of Rhodomonas compared with the control without daphnids.This study shows that the elemental content and growth rateof heterotrophic bacteria and Rhodomonas are influenced by grazingand nutrient regeneration by daphnids. The response is dynamicand depends on the level of nutrient limitation of the system. Present address: Department of Microbiology, University of BergenJahnebakken 5, NO-5020 Bergen, Norway     

Heat tolerance,temperature acclimation,acute oxidative damage and canalization of haemoglobin expression in <Emphasis Type="Italic">Daphnia</Emphasis>

Daphnia is a widespread freshwater zooplankton species, which is both a classic and emerging new model for research in ecological physiology, ecotoxicology and evolutionary biology of adaptation to novel environments. Heat tolerance in Daphnia is known to depend both upon evolutionary history of a genotype and on individuals’ acclimation to elevated temperature and to correlate with the level of haemoglobin expression. We demonstrate the existence of north–south gradient of heat tolerance in North American D. pulex, which is not associated with any parallel changes in haemoglobin expression. Geographically distinct clones differ in the way their haemoglobin expression changes due to acclimation to a sub-stressful (28°C) temperature, but these changes are not correlated with the latitude of clones’ origin. Likewise, the effect of acclimation to sub-stressful temperature is independent from, and cannot be fully explained by, haemoglobin expression changes during acclimation. The degree of oxidative damage to haemoglobin, measured as the ratio of absorbance at 540:576 nm at the acclimation temperature, is a strong predictor of 28°C-acclimated Daphnia survival during an acute heat exposure. The comparison of haemoglobin expression in resistant and tolerant clones acclimated to different temperatures indicates that tolerant clones exhibit canalization of haemoglobin expression, possessing a high level of haemoglobin even at non-stressful temperatures. We discuss the evolutionary biology of adaptation and acclimation to elevated temperatures in an ecologically important component of freshwater ecosystems in the context of global climate change.