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.     

鲢放养和施肥对盐碱池塘围隔生态系统浮游生物群落的影响

报道了单养链（Hypophthalmichthys molitrix)和施肥对盐碱池塘围隔生态系统浮游生物群落的影响,链的放养使浮游植物丰度,叶绿素a 含量和和初组生产力增大,浮游植物小型化,生物量以小型硅藻和绿藻占优势,裸藻和金藻占有相当比重;浮游动物生物量减少,特别是枝角类的生物量无鱼围隔大于有鱼围隔,且多是较大型的种类,施肥特别是施无机肥能显著地提高浮游植物丰度和初级生产力,浮游动物生物量也增大,施有机肥围隔浮游植物和浮游动物生物量虽高于有鱼对照围隔,但其浮游植物初级生产力,叶绿素a含量,浮游生物多样性指数,P／R系数均较低,链鱼的生长最差,文中讨论了滤食性鱼类和施肥对浮游生物的影响。     

Effects of a filter-feeding fish [silver carp, Hypophthalmichthys molitrix (Val.)] on phyto- and zooplankton in a mesotrophic reservoir: results from an enclosure experiment

SUMMARY 1. Silver carp, Hypophthalmichthys molitrix (Val.), feeds on both phyto- and zooplankton and has been used in lake biomanipulation studies to suppress algal biomass. Because reports on the effects of silver carp on lake food webs have been contradictory, we conducted an enclosure experiment to test how a moderate biomass of the fish (10 g wet weight m⁻³) affects phytoplankton and crustacean zooplankton in a mesotrophic temperate reservoir.
2. Phytoplankton biomass <30 μm and particulate organic carbon (POC) <30 μm were significantly higher in enclosures with silver carp than in enclosures without fish, whereas Secchi depth was lower. Total copepod biomass declined strongly in both treatments during the experiment, but it was significantly higher in fish-free enclosures. Daphnid biomass was also consistently higher in enclosures without fish, although this effect was not significant. However, the presence of fish led to a fast and significant decrease in the size at maturity of Daphnia galeata Sars. Thus, the moderate biomass of silver carp had a stronger negative effect on cladoceran zooplankton than on phytoplankton.
3. Based on these results and those of previous studies, we conclude that silver carp should be used for biomanipulation only if the primary aim is to reduce nuisance blooms of large phytoplankton species (e.g. cyanobacteria) that cannot be effectively controlled by large herbivorous zooplankton. Therefore, stocking of silver carp appears to be most appropriate in tropical lakes that are highly productive and naturally lack large cladoceran zooplankton.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.     

Size-selective dispersal of Daphnia resting eggs by backswimmers (Notonecta maculata)

Freshwater zooplankton is increasingly used to study effects of dispersal on community and metacommunity structure. Yet, it remains unclear how zooplankton disperses. Clearly, birds and wind play a significant role as zooplankton dispersal agents, but they may not always be the main vectors. This experimental study shows that a cosmopolitan aquatic insect, Notonecta, can be an important vector of cladoceran resting eggs (ephippia). Dispersing Notonecta frequently transported ephippia during flight, with a bias towards smaller ephippia in two species. A similar trend was present at the species level: Daphnia species with smaller ephippia were more often dispersed, suggesting that Notonecta could generate specific colonist communities. In addition, buoyancy appeared a critical trait, as non-floating ephippia of Daphnia magna were never dispersed. Our data suggest that Notonecta could be important dispersers of Daphnia, and that knowledge of dispersal dynamics of Notonecta may be used to predict Daphnia dispersal, colonization and resilience to disturbance.     

Correlations between nutrient concentrations and zooplankton populations in a mesotrophic reservoir

1. A year-round study was conducted in a mesotrophic reservoir to determine the dynamics of zooplankton populations as a function of food availability (edible phytoplankton), nutrient concentration, temperature and hydraulic regime.
2. Rotifer biomass was correlated with soluble reactive phosphorus (SRP) concentration. The abundance of the rotifers Keratella cochlearis and Anuraeopsis fissa were not correlated with food availability (measured by chlorophyll and cell counts) but showed a strong dependence on P availability. Another rotifer, Synchaeta oblonga , and crustacean species were not related to nutrient availability but seemed to be dependent on food concentrations, especially of some phytoplankton taxa.
3. In this field study, rotifers seemed more susceptible than Daphnia or copepods to P-limitation. Among rotifer species, Keratella seemed to be more susceptible than Anuraeopsis to P limitation. Different susceptibilities of zooplankton species to nutrient limitation may be important in explaining the dynamics of these organisms in natural situations. Further analyses are warranted to clarify the interactions between nutrient limitation and energy limitation among zooplankton.     

The impact of planktivorous flsh on the structure of a plankton community

SUMMARY. 1. The abundance of pianktivorous juvenile yellow perch, Perca flavescens , was manipulated in three 750 m³ enclosures in a eutrophic lake.
2. There was a significant negative relationship between fish and zoopiankton biomasses. At high fish densities the zooplankton community was dominated by small filter-feeding cladocera. primarily bosmi- nids. At low fish densities the zooplankton community was dominated by large filter-feeding cladocera, primarily daphnids.
3. There was no significant relationship between zooplankton and phytoplankton biomasses when considered over the whole experiment but there was a trend towards lower phytoplankton biomass in the enclosure dominated by daphnids during mid-summer.
4. We conclude that although planktivorous fish have a strong negative impact on zooplankton community biomass and size structure, the relationship at the next lower trophic level, zooplankton and phytoplankton, is much weaker. Therefore, the biomanipulation of planktivorous fish populations as a management technique to control phytoplankton abundance is largely ineffective.     

Effects of eutrophication on seasonal patterns of mesozooplankton in a tropical reservoir: a 4‐year study in Pampulha Lake, Brazil

1. Pampulha Lake has undergone rapid eutrophication. Annual maxima of total phosphorus and conductivity increased steadily from 1993 to 1996. Nitrogen quickly decreased after macrophyte removal in 1994, but increased again in the following years
2. The zooplankton was dominated by Daphnia gessneri , D. laevis , Diaphanosoma birgei and Thermocyclops decipiens . The biomass of major zooplankton organisms, including Daphnia and rotifers, increased during the period covered in this study.
3. Autocorrelation coefficients have confirmed the existence of recurrent seasonal patterns for both chemical and biological variables.
4. Daphnia , calanoid copepods and rotifers showed clear and recurrent seasonal patterns. Cyclopoid copepods and Diaphanosoma had temporal patterns more affected by long‐term trophic changes. Other organisms such as Moina, Ceriodaphnia and Bosmina exhibited high temporal variability with no recurrent patterns.
5. There was a strong and positive correlation between total phosphorus and Daphnia biomass. Seasonal patterns of this cladoceran were also inversely associated with sestonic C:P ratios. Thus, Daphnia may be limited primarily by phosphorus and not by total food availability as expressed by particulate carbon.
6. This study suggests that increasing eutrophication has changed the structure as well as seasonal patterns of the zooplankton community in Pampulha Reservoir.     

The zooplankton consumption of primary production and an assessment of the waterbody trophic state on the basis of its structural and functional characteristics

The patterns of the seasonal dynamics of species composition, abundance, biomass, and ration of zooplankton in the Curonian Lagoon of the Baltic Sea have been considered. It is shown that zooplankton can consume up to 17–21% of the phytoplankton primary production. Of this, 7–8% of the primary production is transformed into secondary production and can be used by invertebrates and fishes. The structural and functional characteristics of zooplankton (index and coefficient of the trophic state, the number of dominating species, and the Shannon index by biomass) can be used to assess the waterbody trophic state.     

Production biology of copepods and cladocerans in three south-east Sri Lankan low-land reservoirs and its comparison to other tropical freshwater bodies

Production, biomass and productivity of the microcrustacean zooplanktonpopulations of three low-land reservoirs, Tissawewa (eutrophic), Ridiyagama(moderately eutrophic), and Muruthawela (mesotrophic) in South-east SriLanka were studied. The temporal variation of zooplankton production wasstudied in Tissawewa on basis of fortnightly sampling on five fixed samplingstations for 2 years. Zooplankton production was relatively high, mainlybecause of high copepod production predominantly realised due to twocalanoid copepods, Phyllodiaptomus annae and Heliodiaptomus viduus. Abouthalf of the copepod production was contributed by the naupliar instarstages, whereas the contribution of the eggs was generally much smaller(<20%). In contrast, the cladoceran production consisted for ca50–70% of egg production. The results of this study werecompared with those from more than twenty other tropical and subtropicalwaterbodies reported in the literature by deriving empirical relationshipsbetween mean phytoplankton biomass and mean zooplankton biomass andproduction. Mean zooplankton biomass and annual zooplankton production werefound to be positively related to mean phytoplankton biomass, and meanphytoplankton biomass proved to be a good predictor of mean zooplanktonbiomass (r² = 0.58) and a moderate good predictor ofannual zooplankton production (r² = 0.43). However,the relationships between the mean phytoplankton biomass and zooplankton P/Bwere not significant neither for small-bodied and large-bodied cladoceransnor for copepods.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

The effect of a lake fertilization on the stability and material utilization of a limnetic ecosystem

The interaction between the phytoplankton, zooplankton and fish populations and certain abiotic environmental factors, was investigated in an oligotrophic Norwegian lake during a 5-yr period (1974–1978). The effects of adding artificial fertilizer in 1975 and 1976 were also studied. When cladoceran dominated, the zooplankton community was able to maintain a more or less constant phytoplankton biomass and a rather low phytoplankton production even when nutrient levels were raised. In years when rotifers were dominant, algal biomass and productivity increased, despite the amount of added nutrients being lower. The regression for the relationship between daily phytoplankton P/B and daily herbivore zooplankton P/B indicated that these trophic levels were highly interdependent. A change, from large-sized to smaller herbivorous zooplankton, due to fish predation, also led to an increase in phytoplankton turnover. The investigations show that planktivorous fish may be the key factor which determines the stability of limnetic systems and controls the material transfer from the algae to the higher trophic level.     

Phytoplankton responses to experimental enhancement of grazing pressure and nutrient recycling in a small Andean lake

1. Three series of field experiments with different zooplankton species composition and biomass were performed in a small lake in the south Andes. We attempted to measure the responses of phytoplankton species resulting from grazing mortality and stimulation of growth by nutrient recycling.
2. Nanoflagellates contributed substantially to total phytoplankton cell abundance. Chrysochromulina parva represented 93.4%, 92.2% and 95.9% of total phytoplankton density in December, January and February, respectively. This fraction was reduced in all treatments with increasing zooplankton biomass.
3. A negative relationship was obtained between C. parva cell numbers and increase in dissolved P. On the other hand, a significant positive relationship between the abundance of the diatom Aulacoseira granulata and P concentration was observed. These results indicate that the ungrazed diatom was able to capitalise on the increase in nutrient availability.
4. As a net result of the increase or decrease of algal species we observed a change in the nano:net phytoplankton relationship. The outcome of three‐day incubations with increased zooplankton biomass was an increasing importance of net phytoplankton.
5. The results indicate the importance of the indirect effects of zooplankton (through nutrient recycling) in the increase in diatoms, and the role of grazing as a growth‐limiting factor for the flagellate C. parva .     

Consistent trophic amplification of marine biomass declines under climate change

The impact of climate change on the marine food web is highly uncertain. Nonetheless, there is growing consensus that global marine primary production will decline in response to future climate change, largely due to increased stratification reducing the supply of nutrients to the upper ocean. Evidence to date suggests a potential amplification of this response throughout the trophic food web, with more dramatic responses at higher trophic levels. Here we show that trophic amplification of marine biomass declines is a consistent feature of the Coupled Model Intercomparison Project Phase 5 (CMIP5) Earth System Models, across different scenarios of future climate change. Under the business‐as‐usual Representative Concentration Pathway 8.5 (RCP8.5) global mean phytoplankton biomass is projected to decline by 6.1% ± 2.5% over the twenty‐first century, while zooplankton biomass declines by 13.6% ± 3.0%. All models project greater relative declines in zooplankton than phytoplankton, with annual zooplankton biomass anomalies 2.24 ± 1.03 times those of phytoplankton. The low latitude oceans drive the projected trophic amplification of biomass declines, with models exhibiting variable trophic interactions in the mid‐to‐high latitudes and similar relative changes in phytoplankton and zooplankton biomass. Under the assumption that zooplankton biomass is prey limited, an analytical explanation of the trophic amplification that occurs in the low latitudes can be derived from generic plankton differential equations. Using an ocean biogeochemical model, we show that the inclusion of variable C:N:P phytoplankton stoichiometry can substantially increase the trophic amplification of biomass declines in low latitude regions. This additional trophic amplification is driven by enhanced nutrient limitation decreasing phytoplankton N and P content relative to C, hence reducing zooplankton growth efficiency. Given that most current Earth System Models assume that phytoplankton C:N:P stoichiometry is constant, such models are likely to underestimate the extent of negative trophic amplification under projected climate change.     

<Emphasis Type="Italic">Daphnia</Emphasis> versus copepod impact on summer phytoplankton: functional compensation at both trophic levels

Here we report on a mesocom study performed to compare the top-down impact of microphagous and macrophagous zooplankton on phytoplankton. We exposed a species-rich, summer phytoplankton assemblage from the mesotrophic Lake Schöhsee (Germany) to logarithmically scaled abundance gradients of the microphagous cladoceran Daphnia hyalina×galeata and of a macrophagous copepod assemblage. Total phytoplankton biomass, chlorophyll a and primary production showed only a weak or even insignificant response to zooplankton density in both gradients. In contrast to the weak responses of bulk parameters, both zooplankton groups exerted a strong and contrasting influence on the phytoplankton species composition. The copepods suppressed large phytoplankton, while nanoplanktonic algae increased with increasing copepod density. Daphnia suppressed small algae, while larger species compensated in terms of biomass for the losses. Autotrophic picoplankton declined with zooplankton density in both gradients. Gelatinous, colonial algae were fostered by both zooplankton functional groups, while medium-sized (ca. 3,000 µm³), non-gelatinous algae were suppressed by both. The impact of a functionally mixed zooplankton assemblage became evident when Daphnia began to invade and grow in copepod mesocosms after ca. 10 days. Contrary to the impact of a single functional group, the combined impact of both zooplankton groups led to a substantial decline in total phytoplankton biomass.     

Negative effects of UVB-irradiated phytoplankton on life history traits and fitness of Daphnia magna

SUMMARY 1. We tested the effect of ultraviolet-B (UVB)-irradiated phytoplankton on life history characteristics of Daphnia magna . Two phytoplankton species were used, Chlamydomonas reinhardtii and Cryptomonas pyrenoidifera . The phytoplankton species were cultured under photosynthetically active radiation (PAR) conditions, and under PAR supplemented with ultraviolet-A and ultraviolet-B radiation, and fed to Daphnia .
2. Life history traits of Daphnia were negatively affected when fed on UVB-irradiated Cryptomonas . Size at maturity was depressed and fewer juveniles with lower fitness were produced in the UVB treatments. In the Chlamydomonas experiment, no significant effects were found.
3. The cause of the observed UVB effects is likely to be constraints in food quality. Ultraviolet-B radiation thus has the potential of inhibiting energy transfer from the first to the second trophic level.     

A model approach to planktonic stoichiometry and consumer-resource stability

1. The model explores stoichiometric feedback effects between an alga and a grazer ( Daphnia ) in a simplified chemostat-type system in stagnant conditions or with fixed dilution rate.
2. When running the model with fixed stoichiometry and P-sufficient food, the grazer with highest requirements for phosphorus (P) will exert the most efficient control of algal biomass owing to more P being allocated to zooplankton biomass and less P recycled.
3. When including potential P-limitation of the grazer, the grazer with high P requirements (high P : C ratio) will be the least efficient grazer in a system with fluctuating and temporarily low P : C ratio in algae ( Q _a).
4. Qualitatively deficient food will yield decreased growth efficiency in zooplankton. As Q _a decreases, the grazer isocline for zero net growth is shifted upwards, and the required algal biomass for positive growth increases. There may then be a critical level of Q _a below which the grazer with high P : C suffers negative population growth regardless of algal biomass. In cases with low minimum Q _a and a P-demanding grazer, this may cause the system to enter an irreversible stage with high biomass of P-deficient phytoplankton which do not support zooplankton growth.
5. Cumulative primary production for scenarios with continuous P input is, in general, higher the more Q _a is allowed to drop below saturation values, and highest when this is combined with a grazer with a high P : C ratio. The lower growth rate of P-deficient phytoplankton was compensated for by reduced success of the P-limited grazer, yielding low grazing pressure and resulting in larger phytoplankton biomass.     

Ecosystem Functions across Trophic Levels Are Linked to Functional and Phylogenetic Diversity

In experimental systems, it has been shown that biodiversity indices based on traits or phylogeny can outperform species richness as predictors of plant ecosystem function. However, it is unclear whether this pattern extends to the function of food webs in natural ecosystems. Here we tested whether zooplankton functional and phylogenetic diversity explains the functioning of 23 natural pond communities. We used two measures of ecosystem function: (1) zooplankton community biomass and (2) phytoplankton abundance (Chl a). We tested for diversity-ecosystem function relationships within and across trophic levels. We found a strong correlation between zooplankton diversity and ecosystem function, whereas local environmental conditions were less important. Further, the positive diversity-ecosystem function relationships were more pronounced for measures of functional and phylogenetic diversity than for species richness. Zooplankton and phytoplankton biomass were best predicted by different indices, suggesting that the two functions are dependent upon different aspects of diversity. Zooplankton community biomass was best predicted by zooplankton trait-based functional richness, while phytoplankton abundance was best predicted by zooplankton phylogenetic diversity. Our results suggest that the positive relationship between diversity and ecosystem function can extend across trophic levels in natural environments, and that greater insight into variation in ecosystem function can be gained by combining functional and phylogenetic diversity measures.     

Spatial insurance against a heatwave differs between trophic levels in experimental aquatic communities

Climate change-related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better-adapted taxa. Yet, experimental evidence for such predictions from multi-trophic communities and pulse-type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat-induced increase in metabolic costs, resulting in weaker top-down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer-lasting changes in ecosystem functioning.