Structure et fonctionnement des ecosystemes du haut-rhone francais. 14-etude des preferences alimentaires de la loche franche (noemacheilus barbatulus L.) par une methode des points modifiee

The net effect of zooplankton on phytoplankton productivity was investigated through experiments using natural concentrations of plankton in the epilimnion of Castle Lake, California. Zooplankton enhanced algal productivity during July and August, and nocturnal grazers caused greater proportionate increases than did daytime densities. Consumers had neutral or negative effects in September. Animal biomass was associated significantly with enhanced productivity for the experiment in late July, and with decreased growth rates in the last trial of September. The direct relationship between the activities of zooplankton and primary productivity observed in other lakes is qualified by this seasonal change in net effect. The removal of algae by grazing, increases in the productivity to biomass ratio through nutrient regeneration and temporary photosynthetic inhibition from ammonia excretion appear to have shifted in relative impact during the 3 month experimental period.     

Covariation between zooplankton community composition and cyanobacterial community dynamics in Lake Blaarmeersen (Belgium)

The cyanobacterial community composition in the mesotrophic Lake Blaarmeersen was determined by denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S rRNA gene fragments during two consecutive years to assess the importance of different classes of explanatory variables (bottom-up and top-down factors, physical variables and phytoplankton) in cyanobacterial community dynamics. The most dominant cyanobacteria in Lake Blaarmeersen were Synechococcus (three genotypes), Limnothrix redekei and Anabaena/Aphanizomenon. Analyses of Similarity revealed that the cyanobacterial community in Lake Blaarmeersen differed significantly between the growing season and the winter season as well as between the epilimnion and hypolimnion during the stratified periods. Mantel tests revealed significant correlations between the DGGE data and bottom-up factors, physical variables, the phytoplankton community composition and, interestingly, the zooplankton community composition. In general, the zooplankton community composition (especially the cladoceran community) was more important in structuring the cyanobacterial community than the total zooplankton biomass. This study shows that grazing zooplankton communities can have a relatively strong impact on the cyanobacterial community dynamics and that this impact can be equally important as bottom-up processes regulated by nutrient concentrations and/or physical variables.     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

Seasonal changes in the chemistry and biology of a meromictic lake (Big Soda Lake,Nevada, U.S.A.)

Big Soda Lake is an alkaline, saline lake with a permanent chemocline at 34.5 m and a mixolimnion that undergoes seasonal changes in temperature structure. During the period of thermal stratification, from summer through fall, the epilimnion has low concentrations of dissolved inorganic nutrients (N, Si) and CH₄, and low biomass of phytoplankton (chlorophyll a ca. 1 mgm ^-3). Dissolved oxygen disappears near the compensation depth for algal photosynthesis (ca. 20 m). Surface water is transparent so that light is present in the anoxic hypolimnion, and a dense plate of purple sulfur photosynthetic bacteria (Ectothiorhodospira vacuolata) is present just below 20 m (Bchl a ca. 200 mgm^-3). Concentrations of N H₄ ⁺, Si, and CH₄ are higher in the hypolimnion than in the epilimnion. As the mixolimnion becomes isothermal in winter, oxygen is mixed down to 28 m. Nutrients (NH₄ ⁺, Si) and CH₄ are released from the hypolimnion and mix to the surface, and a diatom bloom develops in the upper 20 m (chlorophyll a > 40 mgm^-3). The deeper mixing of oxygen and enhanced light attenuation by phytoplankton uncouple the anoxic zone and photic zone, and the plate of photosynthetic bacteria disappears (Bchl a ca.10mgm^-3). Hence, seasonal changes in temperature distribution and mixing create conditions such that the primary producer community is alternately dominated by phytoplankton and photosynthetic bacteria: the phytoplankton may be nutrient-limited during periods of stratification and the photosynthetic bacteria are light-limited during periods of mixing.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

Bacterioplankton in a small polyhumic lake with an anoxic hypolimnion

Bacterioplankton biomass and dark fixation of inorganic carbon were measured in the highly humic (water colour up to 550 mg Pt l^?1) and acidic lake, Mekkojärvi. Strong thermal and chemical stratification developed in the water column early in spring and led rapidly to anoxia in the hypolimnion, which extended to less than 1.0 m from the surface. In the epilimnion only small bacteria were abundant. In the anoxic zone both the abundance and the mean size of bacteria were considerably higher than in the epilimnion. These differences are thought to be the result of different grazing pressure from zooplankton in the two zones. In late summer a high concentration of bacteriochlorophyll d in the upper hypolimnion indicated a high density of photosynthetic bacteria. Bacterial biomass was similar to that of phytoplankton in the epilimnion, but 23 times higher in the whole water column. In August, dark fixation of inorganic radiocarbon in the anaerobic zone was 51% of the total ¹⁴C-incorporation and the contribution of light fixation was only 5.4%. In the polyhumic Mekkojarvi, bacterioplankton was evidently a potentially significant carbon source for higher trophic levels, but bacterioplankton production could not be supported by phytoplankton alone. Allochthonous inputs of dissolved organic matter probably support most of the bacterial production.     

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.     

Nutrient regeneration by zooplankton: effects on nutrient limitation of phytoplankton in a eutrophic lake

We tested the hypothesis that excretion of nutrients by zooplanktoncan reduce the severity of nutrient limitation of phytoplankton,and determine whether the phytoplankton community is limitedby nitrogen or phosphorus. In situ experiments were conductedin eutrophic Lake Mendota (Wisconsin, USA) during the summerof 1988, where phytoplankton were limited by N and P, but periodsof nutrient limitation were transitory Increased zooplanktonbiomass and the consequent increased excretion of nutrientsby zooplankton reduced P limitation (as measured by specificalkaline phosphatase activity) in all experiments Excretionof nutrients also reduced N limitation (as measured by ammoniumenhancement response) in one of three experiments. In additionalexperiments in the more highly eutrophic Lake Wingra, excretionof nutrients by zooplankton reduced both N and P limitationThese results support the hypothesis that zooplankton have potentiallyimportant indirect effects on phytoplankton communities throughrecycling of nutrients     

Diel and vertical variability of seston food quality and quantity in a small subalpine oligomesotrophic lake

Vertical profiles of seston food quality and quantity were measuredin subalpine Castle Lake for particulate carbon, chlorophylla, fatty acids and phosphorus in addition to abiotic parametersincluding water temperature and dissolved oxygen levels. Fieldand laboratory incubation experiments were employed to manipulateDaphnia rosea growth environments. Sestonic eicosapentaenoicacid (EPA) content was much lower, whereas carbon to phosphorus(C:P) ratios were much higher in the epilimnion compared tothe deep-water chlorophyll maximum (16–20 m). In a temperature-controlledincubation, Daphnia grew faster when fed seston from the deep-waterchlorophyll maximum. In in situ cage incubations, D. rosea grewfastest in the epilimnion. Daphnia rosea in a migrating treatmentexhibited intermediate growth rates between the ones for epilimnionand hypolimnion. A projection of D. rosea growth rates by growthmodels without regard to water temperature showed much highergrowth potentials in the hypolimnion. However, with inclusionof water temperature, epilimnetic water always gave higher potentialgrowth rates. In this lake, warmer temperatures of epilimneticwater override the higher food quality and quantity providedby the deep-water chlorophyll maximum.     

Nutrients associated with terrestrial dissolved organic matter drive changes in zooplankton:phytoplankton biomass ratios in an alpine lake

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The stoichiometry of N and P in the pelagic zone of Castle Lake, California

We measured the concentrations, as well as lake-wide amounts,of nitrogen (N) and phosphorus (P) in dissolved, seston andzooplankton pools throughout the water column of Castle Lake,California, during summer, 1991. This allowed us to determinethe stoichiometric ratios of important elements in each pool(C:N, C:P, N:P) as well as for the entire lake. Dissolved andseston pools were the predominant storage compartments for bothN and P; zooplankton never contained >5% of N or 10% of Plake wide. However, by late summer, the concentrations of Pin seston and in zooplankton were similar in the upper portionsof the water column, suggesting that changes in food web structurethat alter zooplankton biomass and community composition (andhence elemental storage in the zooplankton) may produce significantshifts in nutrient storage among pelagic pools. Lake-wide levelsof dissolved N were largely constant over the study period;however, lake-wide dissolved P increased. These dynamics suggestedthat the majority of nutrients stored in dissolved pools wereunavailable for phytoplankton growth. N:P and C:P ratios indicatedthat Castle Lake phytoplankton became severely deficient inP during the course of our observations. These ratios also greatlyexceeded recently reported threshold values for elemental constraintson growth and reproduction for several species of zooplankton.The ratio of N to P in the zooplankton pool was relatively constantand consistently lower than that in the sestion. As a result,the predicted N:P ratio of zooplankton-regenerated nutrientsexceeded the N:P ratio of the seston, implying that zooplanktonnutrient regeneration further skewed N and P supply ratios,and potentially enhanced P limitation of phytoplankton in CastleLake. ¹Present address: Department of Biology, Box 19498, Universityof Texas at Arlington, Arlington, TX 76019, USA     

Role of nutrients and zooplankton in regulation of phytoplankton in Flathead Lake (Montana, U.S.A.), a large oligotrophic lake

1. Increased primary production in Flathead Lake during the 1980s has been variously attributed to increased nutrient loadings and/or decreases in zooplankton abundance resulting from the introduction of Mysis relicta . In order to assess the importance of these two factors in regulating the phytoplankton community in Flathead Lake, we manipulated zooplankton abundance and nutrient availability in a series of 5‐day enclosure experiments.
 2. Chlorophyll a levels were stimulated by simultaneous addition of nitrogen and phosphorus. At ambient nutrient levels, alteration of zooplankton density had no effect on chlorophyll a levels. Top‐down control through zooplankton grazing could only be demonstrated in treatments supplemented with nutrients. Under these conditions, there was a significant negative correlation between zooplankton abundance and final chlorophyll a levels.
 3. These results suggest that the phytoplankton community in Flathead Lake is regulated primarily by bottom‐up controls. Consequently, future management activities aimed at preventing further increases in algal growth in the lake should focus on nutrient abatement. Alteration of the upper trophic levels does not appear to have significantly affected phytoplankton abundance in the lake. Should nutrient levels increase in the future, then top‐down controls may become more important.
 4. A conceptual model is presented illustrating the relative importance of top‐down and bottom‐up controls across a trophic gradient.     

Analysis of the change in dominant phytoplankton species in unstratified Lake Oshima-Ohnuma estimated by a bottle incubation experiment

The densities and growth rates of dominant phytoplankton in Lake Oshima-Ohnuma were determined during May to July 1996 to clarify the mechanism of change from spring-dominant to summer-dominant phytoplankton species, examining the influence of zooplankton grazing and nutrient limitation. The dominant phytoplankton in April and May were Asterionella gracillima and Nitzschia spp. Their growth rates were low in May and their densities fell thereafter. The bottle experiment suggests that the main reason for decreases in dominant species has been nutrient deficiency, and zooplankton (sized from 30 to 160 μm) have supported the growth of A. gracillima to recycle nutrients in May. The maximum growth rate was shown by Melosira (Aulacoseira) ambigua living at 2 m in May and June. Although the growth rate of the summer-dominant species Melosira (Aulacoseira) granulata was not as high as that of M. ambigua in May and June, the ambient density increased from late June. Survival in the lower zone of the unstratified euphotic layer might have contributed to the abundance of M. granulata, since the growth rate of this species was not so low as that of M. ambigua at 6 m in June and July. The bottle experiment suggests that the growth of M. granulata was also nutrient limited in June and July and that zooplankton grazing was partially responsible for repressing the biomass of M. granulata before it began growing rapidly in late June. Received: January 31, 2000 / Accepted: September 6, 2000     

Seston food quality and Daphnia production efficiencies in an oligo-mesotrophic Subalpine Lake

Because of major biochemical imbalances between plants and animals, ecological efficiency at this interface may have a major impact on overall energy flow in ecosystems. In order to study relationships between seston food quality and energy transfer between primary producers and herbivores, we conducted five microcosm experiments in Castle Lake, California, USA during the summer of 1996. We simultaneously performed life table experiments to determine the effects of highly unsaturated fatty acids (HUFA) on Daphnia rosea growth, reproduction and survival. The results of these experiments suggest strong energy limitation of D. rosea growth in Castle lake during the study. D. rosea production was coupled with primary production in Castle Lake and in the microcosm experiments. D. rosea production efficiencies, i.e., the ratios of D. rosea productivity to primary productivity, decreased towards the end of the summer. A food quality index based on phytoplankton species composition and seston carbon to phosphorus (C:P) ratio were good predictors of D. rosea production efficiencies. The predicted D. rosea production pattern based on phytoplankton composition and primary productivity matched the zooplankton biomass dynamics in Castle Lake during 1991. Life table experiments showed HUFA effects on D. rosea population growth rates, reproduction and survival in support of the HUFA limitation hypothesis.     

Zooplankton-phytoplankton interactions in a eutrophic lake

Enclosure experiments were made in a cyanobacteria dominatedlake (Lake Rotongaio) to assess the impact of zooplankton (>150µm) grazing on algal growth rates and determine the effectof diel and vertical changes in zooplankton grazing intensityand nutrient (NH₄-N) regeneration upon abundance of phytoplankton.The filamentous cyanobacterium Anabaena minutissima var. attenuataand diatom Cyclotella meneghiniana showed a negative linearchange in abundance with a gradient in zooplankton grazing intensity.Phytoflagellates were not grazed and showed a positive linearchange in abundance with increasing zooplankton biomass. Theseeffects, as well as shortening of filament length of Anabaena,were caused by raptorial feeding by the alanoid copepod Boeckellapropinqua which dominated the zooplankton. Phytoplankton growthwas not stimulated by addition of nutrients, suggesting nutrientregeneration was not important. Diel and vertical changes infeeding and NH₄-N regeneration rates were measured in Marchand June 1988. Diel differences were more pronounced in Marchwhen the water column was stratified. Specific feeding rateswere more important than vertical changes in zooplankton biomassin determining community grazing rates in March, but in Junewhen the water column was mixed, vertical distribution of zooplanktonbiomass was important. Zooplankton grazing was an importantloss process for phytoplankton in the lower part of the epilimnionin Lake Rotongaio.     

Summer dynamics of the deep chlorophyll maximum in Lake Tahoe

Vertical profiles of chlorophyll and phytoplankton biomass weremeasured in Lake Tahoe from July 1976 through April 1977. Adeep chlorophyll maximum (DCM) persisted during summer and earlyautumn (July—October) near 100 m, well below the mixedlayer and at the upper surface of the nitracline. The DCM coincidedwith the phytoplankton biomass maximum as determined from cellcounts. In addition, the composition of the phytoplankton assemblagewas highly differentiated with respect to depth. Cyclotellastelligera was the predominant species in the mixed layer whilethe major species in the DCM layer included C. ocellata andseveral green ultraplanktonic species. In situ cell growth playsa substantial role in maintaining the DCM, but sinking of cellsfrom shallower depths and zooplankton grazing above the DCMmay contribute to the maintenance of the DCM. Calculations supportthe interpretation that the summer DCM persists at the boundarybetween an upper, nutrient-limited phytoplankton assemblageand a deeper, light-limited assemblage.     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.