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.     

The eutrophic Loosdrecht Lakes: Current ecological research and restoration perspectives

Features of the Loodsrecht Lakes, with emphasis on the main lake, are discussed with reference to restoration.Characteristics of the present situation are: (1) very low water transparency-Secchi-disc readings around 0.3 m occur in all seasons; (2) relatively small seasonal changes in sestonic matter; (3) important input of resuspended particles into the seston; (4) predominance of filamentous blue-green algae for most of the year; (5) relative scarcity of crustacean zooplankton, while rotifers are abundant; (6) poor development of littoral communities, and absence of benthic producers. The blue-green algae maintain high population density at very low growth rates: rates of loss are low. The zooplankton grazing rate is low due to inefficient filtering, but predation of larger crustaceans by fish may also be important. Studies on epipelon indicated that loss by deposition may be largely compensated by resuspension.Starting in 1984, the external phosphorus loading was markedly reduced. Results for 1984 and 1985 indicate that complementary measures are needed in order to improve water quality. Action should be directed towards increasing the phytoplankton turnover rates. Accelerated specific growth rate can be expected to accompany lower biomass, more successful competition by other algal groups, and enhancement of grazing pressure.Considering the shallowness of the system, promotion of littoral development and return of submerged vegetation may be important in establishing a new equilibrium of the system.     

Optimizing the complexity of phytoplankton functional group modeling: An allometric approach

Elucidating patterns and mechanisms that shape phytoplankton assemblages is a popular area of research for empirical and theoretical ecologists. Despite the daunting complexity of phytoplankton dynamics, much of our current understanding has been based on simple models describing food-web interactions with few differential equations. Skeptical views in the literature raise concerns about the increasing model complexity and advice to seek parsimony rather than simplicity. To address this controversy (simple versus complex models), we propose the introduction of an extra layer of causality into plankton models by connecting algal processes (maximum growth rates, nutrient kinetics, settling velocities, metabolic rates) with species-specific morphological features (cell volume, surface-to-volume ratio, shape). In this study, we demonstrate the capacity of a size-based plankton model to reproduce observed water quality patterns (phosphate, total phosphorus, nitrate, total ammonia, total nitrogen, chlorophyll a, and total zooplankton biomass) in the Hamilton Harbour, Ontario. Consistent with empirical evidence, our modeling analysis showed that small algal species have a distinct competitive advantage in summer epilimnetic environments across the range of cell volume and nutrient loading conditions examined; especially, when they are characterized by higher optimal temperature for growth. Strong top-down pressure mediated by high zooplankton abundance effectively controls the standing biomass of phytoplankton species that can otherwise realize high growth rates under the conditions typically prevailing in the end-of-summer epilimnetic environments (e.g., higher temperature optima, higher tolerance in low water clarity). Under high zooplankton control, the secondary variations of phytoplankton are modulated by the ambient phosphorus levels and the size-based strategies for resources procurement, such as the regulation of nutrient transport kinetics. By contrast, when the summer algal assemblage is released by the zooplankton grazing, the exceedance of critical phytoplankton biomass levels and the likelihood of harmful algal blooms are determined by the multitude of factors that shape inter-specific competition patterns (e.g., relative abundance of competing species, nutrient uptake kinetics). Our study evaluates the strengths and weaknesses of this approach and identifies future directions that would provide operational models founded upon concepts of allometry.     

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.     

Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.     

Size fractionation of algal chlorophyll, carbon fixation and phosphatase activity: relationships with species-specific size distributions and zooplankton community structure

Algal chlorophyll, carbon fixation and alkaline phosphataseactivity were net-fractionated using 22-µm, and 75-µmscreens in three lakes with contrasting zooplankton communities.Size distributions of algal biovolume were also determined throughmicroscopic examination. Relatively good correspondence wasfound between size distributions obtained through net fractionationand those determined by microscopic examination. Biovolume-specificcarbon fixation and chlorophyll decreased with increasing fractionsize but no differences were observed among fractions for chlorophyll-specificcarbon fixation. High algal standing stocks and low phosphorusdeficiency in Tuesday Lake were attributed to low grazing pressureby small, inefficient zooplankton and possible limitation bynutrients other than phosphorus. Algal standing stocks werelow and phosphorus deficiency was high in Peter and Paul Lakes,in which the zooplankton was dominated by large grazers. Differentalgal size fractions experienced differing degrees of phosphorusdeficiency. These size fraction differences in P-deficiencyin Peter and Paul Lakes were attributed to differences in algalspecies composition and to differing levels of zooplankton grazingpressure and nutrient regeneration. A unimodal relationshipbetween relative nanoplankton biovolume and zooplankton biomasswas found and reflects the positive (nutrient regeneration)and negative (grazing mortality) effects of zooplankton on thealgal community.     

Positive feedbacks between bottom-up and top-down controls promote the formation and toxicity of ecosystem disruptive algal blooms: A modeling study

Harmful algal blooms that disrupt and degrade ecosystems (ecosystem disruptive algal blooms, EDABs) are occurring with greater frequency and severity with eutrophication and other adverse anthropogenic alterations of coastal systems. EDAB events have been hypothesized to be caused by positive feedback interactions involving differential growth of competing algal species, low grazing mortality rates on EDAB species, and resulting decreases in nutrient inputs from grazer-mediated nutrient cycling as the EDAB event progresses. Here we develop a stoichiometric nutrient–phytoplankton–zooplankton (NPZ) model to test a conceptual positive feedback mechanism linked to increased cell toxicity and resultant decreases in grazing mortality rates in EDAB species under nutrient limitation of growth rate. As our model EDAB alga, we chose the slow-growing, toxic dinoflagellate Karenia brevis, whose toxin levels have been shown to increase with nutrient (nitrogen) limitation of specific growth rate. This species was competed with two high-nutrient adapted, faster-growing diatoms (Thalassiosira pseudonana and Thalassiosira weissflogii) using recently published data for relationships among nutrient (ammonium) concentration, carbon normalized ammonium uptake rates, cellular nitrogen:carbon (N:C) ratios, and specific growth rate. The model results support the proposed positive feedback mechanism for EDAB formation and toxicity. In all cases the toxic bloom was preceded by one or more pre-blooms of fast-growing diatoms, which drew dissolved nutrients to low growth rate-limiting levels, and stimulated the population growth of zooplankton grazers. Low specific grazing rates on the toxic, nutrient-limited EDAB species then promoted the population growth of this species, which further decreased grazing rates, grazing-linked nutrient recycling, nutrient concentrations, and algal specific growth rates. The nutrient limitation of growth rate further increased toxin concentrations in the EDAB algae, which further decreased grazing-linked nutrient recycling rates and nutrient concentrations, and caused an even greater nutrient limitation of growth rate and even higher toxin levels in the EDAB algae. This chain of interactions represented a positive feedback that resulted in the formation of a high-biomass toxic bloom, with low, nutrient-limited specific growth rates and associated high cellular C:N and toxin:C ratios. Together the elevated C:N and toxin:C ratios in the EDAB algae resulted in very high bloom toxicity. The positive feedbacks and resulting bloom formation and toxicity were increased by long water residence times, which increased the relative importance of grazing-linked nutrient recycling to the overall supply of limiting nutrient (N).     

A review: limnological management and biomanipulation in the London reservoirs

Low algal biomasses and high water transparencies are a feature of the storage reservoirs that supply most of London's treated water. This is a result of knowledgeable limnological management and biomanipulation and despite the eutrophic nature of the River Thames with its high nutrients (7 gN m⁻³; 1 gP m⁻³) and particulate organic carbon (2 gC m⁻³). Built-in possibilities of jetting input water are managed to prevent stratification, to ensure isothermy, to mix chemicals and plankton vertically and horizontally and to manipulate the mixed-depth of the algal populations such that their potential for biomass growth is reduced by light-energy limitation. Spring algal growth is delayed and the spring peak is reduced and curtailed by the grazing impact of considerable biomasses of large-bodied daphnid populations (Daphnia magna, pulicaria & hyalina) whose development is also supported by the continuous input of high riverine algal crops. The existence of a large-bodied daphnid zooplankton in the reservoirs is associated with low levels of fish predation since the late 1960s. Variations in the intensity and nature of this vertebrate predation during the subsequent twenty years (1968–88) are illustrated by the changes that have occurred in the relationship between the phytoplankton and zooplankton biomasses of the April-May-June quarter of the year. This example of the London reservoirs serves to illustrate biomanipulation in deep water bodies by bottom-up as well as top-down effects.     

Keystone predator effects and grazer control of planktonic primary production

If prey species exhibit trade-offs in their ability to utilize resources versus their ability to avoid predation, predators can facilitate prey turnover along gradients of productivity, shifting dominance from edible to inedible prey (the keystone predator effect). I tested this model under controlled, laboratory conditions, using a model aquatic system composed of zooplankton as the top consumer, a diverse community of algae as prey, and nutrients as basal resources. Nutrient manipulations (low and high) were crossed with presence–absence of zooplankton. Results supported theoretical predictions. Algal biomass increased in response to enrichment regardless of predator presence/absence. However, predators and nutrients had an interactive effect on algal biomass and size structure. At the low nutrient level, algal-prey were dominated by edible forms and attained similar biomass regardless of zooplankton presence/absence. At the high level of enrichment, presence of zooplankton favored higher levels of algal biomass and shifted dominance to large, inedible taxa. At the termination of the experiment, I performed a series of lab-based assays on the resultant algal community in order to quantify trade-offs among algal size classes in maximal population growth rates (as a measure of competitive ability for nutrients) and susceptibility to zooplankton grazing. Assays provided support for a size-based keystone trade-off. Small size classes of algae displayed higher maximal growth rates but were more susceptible to grazing effects. Large size classes were protected from grazing but showed low rates of population growth in response to enrichment.     

Relative growth rates and the grazing optimization hypothesis

Summary A mathematical analysis of the changes in plant relative growth rates necessary to increase aboveground production following grazing was conducted. The equation derived gives an isoline where production of a grazed and ungrazed plant will be the same. The equation has four variables (mean shoot relative growth rate, change in relative growth rate after grazing, grazing intensity, and recovery time) and may be analyzed graphically in a number of ways.Under certain conditions, small increases in shoot relative growth rate following grazing will lead to increased aboveground production. Under other conditions, very large increases in relative growth rate after grazing can occur without production being increased over that of ungrazed plants. Plants growing at nearly their maximum potential relative growth rate have little opportunity to respond positively to grazing and potentially can sustain less grazing than plants with growth rates far below maximum. Plants with high relative growth rates at the time of grazing require large increases in growth rate while slow growing plants require only small increases. High grazing intensities are least likely to increase production and high grazing frequencies require greater responses than infrequent grazing events.     

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.     

The spatial and temporal dynamics of deep chlorophyll layers in high-mountain lakes: effects of nutrients, grazing and herbivore nutrient recycling as growth determinants

Deep chlorophyll layers (DCL) are a common feature of oligotrophiclakes, yet the mechanisms that form and maintain them are notunderstood fully. These phytoplankton populations occur in themetalimnia of lakes where light levels are moderate to low,and where nutrient levels and zooplankton grazing pressure aredifferent than in the epilimnion. To test the importance ofnutrients and grazing pressure for algal growth in differentlake strata, microcosm experiments and monitoring were conductedin two oligotrophic lakes in the Rocky Mountains of North Americathat contain DCL. In situ microcosm experiments with naturalphytoplankton communities from three depth strata were conductedwith macronutrient additions and with and without the naturalzooplankton grazing communities. Alkaline phosphatase assaysand the in situ microcosm experiments indicated less nutrientlimitation in the metalimnia than in the epilimnia of both lakes.Zooplankton grazing in the experiments decreased algal populationgrowth rates by as much as 6% day^–1, with impacts shiftingto progressively deeper strata over the summer. Zooplanktongrazing losses, however, were partially offset by nutrient recyclingthat increased algal growth rates. Depth-differential nutrientdeficiency and zooplankton grazing and recycling interactedto maintain the DCL in these lakes.     

Modelling phosphorus fluxes in the hypertrophic Loosdrecht Lakes

A dynamic, deterministic model is presented to simulate the phosphorus cycle and plankton growth in the shallow, hypertrophic Loosdrecht Lakes (The Netherlands) before and after restoration measures. Besides inorganic phosphorus (SRP) in both the surface water and the interstitial water, the model comprises three algal groups, zooplankton, fish, detritus, zoobenthos and upper sediment (all modelled both in carbon and in phosphorus). Within the model system, the phosphorus cycle is completely closed. Carbon and phosphorus are described independently, so that the dynamics of the P/C ratios can be modelled. Sediment processes are described in a simplified form.Simulated values are largely within the range of observed ones. The detrital fraction of the seston (=phytoplankton+detritus) varies from 50–60% in summer to about 90% in winter. SRP in the surface water is very low during most of the year. Sensitivity for external phosphorus input is larger for algal and detrital P than for algal and detrital C and chlorophyll-a. So the P/C ratio of the seston decreases following restoration measures, as is observed in the lakes, while the much higher P/C ratios of zooplankton and fish remain constant. Phosphorus mobilisation from the sediment decreases with decreasing external input. Adaptation of the model system to the reduced loading takes place within about two years.Sources of uncertainty in the model include the limited knowledge on selective grazing as well as on mortality and mineralisation processes.     

Predictability and possible mechanisms of plankton response to reduction of planktivorous fish

The predictability of plankton response to reductions of planktivorous fish was investigated by comparing the plankton community in three biomanipulated lakes and ten unmanipulated lakes differing in intensity of fish predation. Data collected on total phosphorus, phytoplankton and zooplankton biomass and share of cyanobacteria and large grazers, as well as specific growth rate of phytoplankton, were further used to test some of the proposed underlying response-mechanisms. In the biomanipulated lakes the algal biomass and share of cyanobacteria decreased, specific growth rate of phytoplankton increased, and zooplankton biomass and share of large grazers increased or remained unchanged. This pattern was largely reflected in the differences in food-chain structure between the unmanipulated lakes with highversus those with low fish predation. The qualitative response to planktivorous fish reduction thus seems largely predictable. The biomanipulated lakes differed, however, in magnitude of response: the smallest hypertrophic, rotenone-treated lake (Helgetjern) showed the most dramatic response, whereas the large, deep mesotrophic lake (Gjersjøen), which was stocked with piscivorous fish, showed more moderate response, probably approaching a new steady state. These differences in response magnitude may be related to different perturbation intensity (rotenone-treatmentversus stocking with piscivores), food-chain complexity and trophic state. Both decreased phosphorus concentration and increased zooplankton grazing are probably important mechanisms underlying plankton response to biomanipulation in many lakes. The results provide tentative support to the hypothesis that under conditions of phosphorus limitation, increased zooplankton grazing can decrease algal biomassvia two separate mechanisms: reduction of the phosphorus pool in the phytoplankton, and reduction of the internal C:P-ratio in the phytoplankton cells.

     

The efficacy of Scenedesmus morphology as a defense mechanism against grazing by selected species of rotifers and cladocerans

Phytoplankton often develop various defense mechanisms in response to zooplankton grazing, such as spines and colonies. While it is now known that increased spine length and cells in a colony of members of the genus Scenedesmus, when zooplankton grazing is intense, helps in reducing zooplankton filtering rates, the effect of these defense mechanisms at the population level has been observed in few studies. Here we present data on the growth rates of four zooplankton species, Brachionus calyciflorus, B. patulus, Ceriodaphnia dubia and Daphnia pulex at two food levels using two species of colony-forming Scenedesmus spp.: S. acutus (cell length = 18.2 ± 0.4 µm; width = 4.2 ± 0.1 µm; average colony length = 90 µm; width: 21 µm) and S. quadricauda (cell length: 21 ± 0.5 width 7.5 ± 0.3 µm; average colony length: 84 µm; width: 30 µm). Whereas S. acutus had no spines, S. quadricauda had spines of 6–10 µm. Population growth experiments of the test rotifers and cladocerans were conducted in 100 ml containers with 50 ml of the medium with test algae. Algae concentrations used were: 13 and 52 mg dw l^–1 of each of the two algal species offered in colonial forms. We used an initial inoculation zooplankter density of 1 ind. ml^–1 for either of the rotifer species and 0.2 ind. ml^–1 for either of the cladoceran species. In all, we had 64 test containers (4 test species of zooplankton × 2 test species of algae × 2 algal densities × 4 replicates). We found a significant effect of algal size on the growth rates of all the four tested species of zooplankton. The population growth rates of zooplankton ranged from –0.58 to 0.66 and were significantly higher on diet of S. acutus than of S. quadricauda. Thus, our study confirms that the larger colony size and the formation of spines in S. quadricauda were effective defenses against grazing by both rotifers and smaller sized cladoceran Ceriodaphnia dubia but that larger-bodied Daphnia pulex could exploit both the algal populations equally.     

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.     

Pheopigment dynamics,zooplankton grazing rates and the autumnal ammonium peak in a Mediterranean lagoon

The dilution technique was used to estimate chlorophyll and pheopigment, net and gross production as well as zooplankton grazing over a 12-month period in a coastal lagoon in Southern France. Chlorophyll a (Cha) based gross growth rates of phytoplankton ranged from undetectable in February to 2.6 day⁻¹ in June, corresponding to 3.8 divisions per day. Cha-based grazing rates ranged from undetectable in February to 1.1 d⁻¹ in June. The seasonal growth pattern of picoplankton was similar to that of the whole community, with a peak in July, corresponding to four divisions per day. Grazing processes represented from 20 to 150% of the phytoplankton daily growth, and the grazing pressure was stronger on small phytoplankton cells than on larger cells. Gross growth rates of phytoplankton were related to zooplankton grazing rates, and both were related to water temperature. Mesozooplankton which escaped sampling or oysters had to be also invoked as additional sinks for the primary production. In the fall, pheopigment concentrations greater than chlorophyll concentrations coincided with high ammonium levels in the water column. Pheopigment a production rates were highly correlated to chlorophyll -based microzooplankton grazing rates. The pheopigment a to chlorophyll a ratio was correlated with ammonium concentrations and could be used an index of the balance between ammonium supply (degradation) and demand (uptake by phytoplankton). In addition, pheopigment degradation rates in absence of grazing could be related to irradiance, indicating photo-degradation of these compounds.     

The efficacy of Scenedesmus morphology as a defense mechanism against grazing by selected species of rotifers and cladocerans

Phytoplankton often develop various defense mechanisms in response to zooplankton grazing, such as spines and colonies. While it is now known that increased spine length and cells in a colony of members of the genus Scenedesmus, when zooplankton grazing is intense, helps in reducing zooplankton filtering rates, the effect of these defense mechanisms at the population level has been observed in few studies. Here we present data on the growth rates of four zooplankton species, Brachionus calyciflorus, B. patulus, Ceriodaphnia dubia and Daphnia pulex at two food levels using two species of colony-forming Scenedesmus spp.: S. acutus (cell length = 18.2 ± 0.4 µm; width = 4.2 ± 0.1 µm; average colony length = 90 µm; width: 21 µm) and S. quadricauda (cell length: 21 ± 0.5 width 7.5 ± 0.3 µm; average colony length: 84 µm; width: 30 µm). Whereas S. acutus had no spines, S. quadricauda had spines of 6–10 µm. Population growth experiments of the test rotifers and cladocerans were conducted in 100 ml containers with 50 ml of the medium with test algae. Algae concentrations used were: 13 and 52 mg dw l^–1 of each of the two algal species offered in colonial forms. We used an initial inoculation zooplankter density of 1 ind. ml^–1 for either of the rotifer species and 0.2 ind. ml^–1 for either of the cladoceran species. In all, we had 64 test containers (4 test species of zooplankton × 2 test species of algae × 2 algal densities × 4 replicates). We found a significant effect of algal size on the growth rates of all the four tested species of zooplankton. The population growth rates of zooplankton ranged from ?0.58 to 0.66 and were significantly higher on diet of S. acutus than of S. quadricauda. Thus, our study confirms that the larger colony size and the formation of spines in S. quadricauda were effective defenses against grazing by both rotifers and smaller sized cladoceran Ceriodaphnia dubia but that larger-bodied Daphnia pulex could exploit both the algal populations equally.     

Biomanipulation: a review of biological control measures in eutrophic waters and the potential for Murray cod Maccullochella peelii peelii to promote water quality in temperate Australia

Biomanipulation is a method of controlling algal blooms in eutrophic freshwater ecosystems. The most common approach has been to enhance herbivores through a reduction of planktivorous fish and introduction of piscivorous fish. The method was originally intended to reduce grazing pressure on zooplankton, thereby increasing grazing pressure on phytoplankton to increase water clarity and promote the growth of aquatic macrophytes. Biomanipulation has received considerable attention since it was proposed in 1975 where innovative approaches and explanations of the processes have been developed. Although many successful biomanipulation exercises have been conducted internationally, it has received comparatively little attention in the Southern Hemisphere and has not been trialled in the southern temperate climate of South Australia. This is a review to speculate upon the criteria for and against the application of biomanipulation in southern temperate Australia using the native species Murray cod (Maccullochella peelii peelii) and to suggest future research.     

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

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