OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Composition and Structure of Zooplankton Communities in Eighteen Arctic and Subarctic Lakes

The factors influencing the composition and structure of zooplankton communities in 18 lakes in the Canadian arctic and subarctic were determined during 1975 and 1976. Phytoplankton were consumed in very low numbers by all species (Diaptomus sicilis, Heterocope septentrionalis, Cyclops scutifer, Daphnia longiremis, Bosmina longirostris, Holopedium gibberum, Keratella cochlearis, Kellicottia longispina) and therefore differences in algal productivity among the lakes had little effect on the zooplankton. Variations in surface area, maximum water depth, pH and the ionic composition of the water were also unimportant in controlling the communities. However, low temperature exerted strong influence over the diversity, abundance, fecundity and vertical distribution of most species.     

Seasonal Cycles of Zooplankton and Related Phytoplankton Development in Three Shallow,Mesotrophic Lakes in Northern Canada

The primary goal of this study was to assess the importance of food in regulating densities of zooplankton in 3 northern Canadian lakes, where algal availability was low compared to temperate zone water bodies. Collections were made every 2 weeks during the summer and monthly during the winter from April 1978 to April 1979. Since the lakes were similar in most respects, including nannoplankton density, net phytoplankton density, temperature, depth, oxygen concentration and phosphorus levels, the seasonal cycles of the main species (Keratella cochlearis, Kellicottia longispina, Polyarthra remata, Polyarthra vulgaris, Cyclops spp., Diaptomus spp.) were generally similar throughout the study area. Changes in the densities of herbivorous species were poorly correlated with fluctuations in nannoplankton and net plankton availability, implying that food did not limit development. Although predatory copepods, particularly Cyclops spp., were abundant, they also had no measurable impact on the main species. It was therefore concluded that temperature controlled the seasonal cycles and the ultimate population size of most zooplankters.     

Phytoplankton and Zooplankton Associations in a Set of Alpine High Altitude Lakes: Geographic Distribution and Ecology

Species composition and interactions, biomass dominance, geographic distribution and driving variables were investigated for two key elements of the pelagic food web of Alpine lakes, the phytoplankton and the zooplankton, based on a single sampling campaign during summer 2000. Altogether, 70 lakes were surveyed, 49 of which located in three different lake districts of the west and eastern Italian Alps and 21 in the central Austrian Alps (within the uppermost Danube catchment). In addition to the analysis of environmental variables affecting distribution and species structure of the two planktonic compartments, a brief review of the main research lines and hypotheses adopted in the past for the study of phytoplankton and zooplankton in high Alpine lakes is given. The lakes, investigated partly within the European project EMERGE (EVK1-CT-1999-00032) and partly within a regional project in the eastern Alps, comprise a wide range of morphological, chemical and trophic conditions. The phytoplankton communities were found to be diverse and mostly dominated by flagellates (chrysophytes, cryptophytes and dinoflagellates), and only to a lesser extent by non-motile green algae, desmids and centric diatoms. The zooplankton communities were mainly dominated by Alpine cladocerans and copepod species, while rotifers were abundant within one group of Italian lakes (sampled in early summer). The multivariate statistical analyses (CCA) showed that catchment features (i.e. percentage of vegetation cover and geochemical composition) and nitrate concentration are essential drivers for the phytoplankton, whereas for zooplankton also trophic status of the lakes and phytoplankton structure are important. The combined variance analysis of the lake clusters outlined by the multivariate analyses on phytoplankton and zooplankton data, respectively, allowed the identification of four principal lake types (three located on siliceous and one on carbonaceous bedrock), each one characterised by a certain combination of habitat features, which in their turn influence trophic state, and phytoplankton and zooplankton species composition and functionality.     

Seasonal and Spatial Overlap between Cladocerans in Humic Lakes

Seasonal and vertical distribution, migratory patterns and reproductive effort in coexisting cladocerans were investigated in three humic lakes with different, but low phytoplankton abundances and varying fish predation pressure. Seasonal and vertical habitat or niche overlap varied, but were high within most pairs of species in all localities. Migration was conspicuous in presence of planktivorous fish, less so in the fish free lake. Despite algal densities below incipient limiting level (30–200 μg C I⁻¹) and a low ratio (3–10) of algal to cladoceran biomass, zooplankton distribution and reproductive parameters were not clearly related to algal biomass. Bacterial biomass equalled 10–50% of phytoplankton biomass, while detritus by far was the largest of the particulate compartements. It was concluded that with a possible exception of the early summer algal bloom, additional carbon sources (bacteria, detritus) are important to cladoceran nutrition in these humic lakes. A large share of N- and P-poor detritus in the diet would give zooplankton productivity limitation by food quality in terms of elemental composition rather than food quantity. This would permit coexistence even of species with rather high food overlap, but give low production rates for all species in agreement with the observations.     

Spatial patterns and relationships between phytoplankton, zooplankton and water quality in the Saimaa lake system, Finland

The interactions between phytoplankton and zooplankton were studied in two large lakes in the Saimaa lake system, Finland. Both are subjected to substantial waste water loading, and exhibit a clear trophic gradient between the loaded and unloaded areas. The phytoplankton and zooplankton were compared in terms of composition, abundance and biomass at 34–39 stations located in different parts of the lakes. At least four mechanisms were thought to affect the composition of plankton communities: (1) the amount of nutrients (trophic gradient), (2) grazing of algae by herbivores, (3) the effect of the algal species composition on feeding by zooplankters (large, colonial algae in the more loaded parts of the lakes) and (4) the regeneration and reorganization of nutrients.     

Temperature is the key factor explaining interannual variability of Daphnia development in spring: a modelling study

Plankton succession during spring/early summer in temperate lakes is characterised by a highly predictable pattern: a phytoplankton bloom is grazed down by zooplankton (Daphnia) inducing a clear-water phase. This sequence of events is commonly understood as a cycle of consumer-resource dynamics, i.e. zooplankton growth is driven by food availability. Here we suggest, using a modelling study based on a size-structured Daphnia population model, that temperature and not food is the dominant factor driving interannual variability of Daphnia population dynamics during spring. Simply forcing this model with a seasonal temperature regime typical for temperate lakes is sufficient for generating the distinctive seasonal trajectory of Daphnia abundances observed in meso-eutrophic temperate lakes. According to a scenario analysis, a forward shift of the vernal temperature increase by 60 days will advance the timing of the Daphnia maximum on average by 54 days, while a forward shift in the start of the spring bloom by 60 days will advance the Daphnia maximum only by less than a third (17 days). Hence, the timing of temperature increase was more important for the timing of Daphnia development than the timing of the onset of algal growth. The effect of temperature is also large compared to the effect of applying different Daphnia mortality rates (0.055 or 0.1 day(-1), 38 days), an almost tenfold variation in phytoplankton carrying capacity (25 days) and a tenfold variation in Daphnia overwintering abundance (3 days). However, the standing stock of Daphnia at its peak was almost exclusively controlled by the phytoplankton carrying capacity of the habitat and seems to be essentially independent of temperature. Hence, whereas food availability determines the standing stock of Daphnia at its spring maximum, temperature appears to be the most important factor driving the timing of the Daphnia maximum and the clear-water phase in spring.     

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.     

Long term study on the influence of eutrophication, restoration and biomanipulation on the structure and development of phytoplankton communities in Feldberger Haussee (Baltic Lake District, Germany)

Feldberger Haussee provides a classic example of eutrophication history of hardwater lakes in the Baltic Lake District (Germany) and of changes in their algal flora during the 20th century. The lake originally was regarded as slightly eutrophic. A process of drastic eutrophication from the 1950s until the end of the 1970s caused mass developments of blue-green and green algae. A restoration program was started in the 1980s to improve the water quality of the lake using both diversion of sewage outside the catchment area, and biomanipulation by altering the fish community. This restoration program led to positive changes in the lake ecosystem. Direct effects of biomanipulation resulted in an increase of herbivorous zooplankton, a decrease of phytoplankton biomass, and an increase of water transparency. The recovery of Feldberger Haussee also may have been indirectly enhanced by an increase in nutrient sedimentation as a consequence of intensified calcite precipitation, decrease in phosphorus remobilization due to a pH-decrease, increased NIP-ratio, and recolonization of the littoral zone by macrophytes. This paper concentrates on the long term development of the phytoplankton community as a response to changes in the food web structure as well as to alterations in the chemical environment of the algae. Both are reflected in four major stages passed by the algal assemblage between 1980 and 1994: (1) From 1980-summer 1985 dense green algal populations were found indicating similar conditions as in the 1970s during the period of maximum eutrophication. (2) A diverse phytoplankton community during summer 1985–1989 showed the first effects of a recovery. (3) From 1990–1992 the phytoplankton was characterized by ungrazeable filamentous blue-green algae first of all as a response to increased herbivory of zooplankton on edible species and to increasing N/P-ratios. (4) Finally, the algal species diversity increased in 1993 and 1994 whereas the phytoplankton biomass decreased showing the success of the combined restoration measures.     

Biotic interactions may overrule direct climate effects on spring phytoplankton dynamics

To improve our mechanistic understanding and predictive capacities with respect to climate change effects on the spring phytoplankton bloom in temperate marine systems, we used a process‐driven dynamical model to disentangle the impact of potentially relevant factors which are often correlated in the field. The model was based on comprehensive indoor mesocosm experiments run at four temperature and three light regimes. It was driven by time‐series of water temperature and irradiance, considered edible and less edible phytoplankton separately, and accounted for density‐dependent grazing losses. It successfully reproduced the observed dynamics of well edible phytoplankton in the different temperature and light treatments. Four major factors influenced spring phytoplankton dynamics: temperature, light (cloudiness), grazing, and the success of overwintering phyto‐ and zooplankton providing the starting biomasses for spring growth. Our study predicts that increasing cloudiness as anticipated for warmer winters for the Baltic Sea region will retard phytoplankton net growth and reduce peak heights. Light had a strong direct effect in contrast to temperature. However, edible phytoplankton was indirectly strongly temperature‐sensitive via grazing which was already important in early spring at moderately high algal biomasses and counter‐intuitively provoked lower and later algal peaks at higher temperatures. Initial phyto‐ and zooplankton composition and biomass also had a strong effect on spring algal dynamics indicating a memory effect via the broadly under‐sampled overwintering plankton community. Unexpectedly, increased initial phytoplankton biomass did not necessarily lead to earlier or higher spring blooms since the effect was counteracted by subsequently enhanced grazing. Increasing temperature will likely exhibit complex indirect effects via changes in overwintering phytoplankton and grazer biomasses and current grazing pressure. Additionally, effects on the phytoplankton composition due to the species‐specific susceptibility to grazing are expected. Hence, we need to consider not only direct but also indirect effects, e.g. biotic interactions, when addressing climate change impacts.     

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.     

Phytoplankton–zooplankton dynamics in periodic environments taking into account eutrophication

In this paper, we derive and analyze a mathematical model for the interactions between phytoplankton and zooplankton in a periodic environment, in which the growth rate and the intrinsic carrying-capacity of phytoplankton are changing with respect to time and nutrient concentration. A threshold value: “Predator’s average growth rate” is introduced and it is proved that the phytoplankton–zooplankton ecosystem is permanent (both populations survive cronically) and possesses a periodic solution if and only if the value is positive. We use TP (Total Phosphorus) concentration to mark the degree of eutrophication. Based on experimental data, we fit the growth rate function and the environmental carrying capacity function with temperature and nutrient concentration as independent variables. Using measured data of temperature on water bodies we fit a periodic temperature function of time, and this leads the growth rate and intrinsic carrying-capacity of phytoplankton to be periodic functions of time. Thus we establish a periodic system with TP concentration as parameter. The simulation results reveal a high diversity of population levels of the ecosystem that are mainly sensitive to TP concentration and the death-rate of zooplankton. It illustrates that the eruption of algal bloom is mainly resulted from the increasing of nutrient concentration while zooplankton only plays a role to alleviate the scale of algal bloom, which might be used to explain the mechanism of algal bloom occurrence in many natural waters. What is more, our results provide a better understanding of the traditional manipulation method.     

Tropical lakes — functional ecology and future development:. The need for a process-orientated approach

The major classes of tropical lakes include shallow, lowland lakes; deep, tertiary lakes; high altitudinal lakes; rainforests lakes; and man-made lakes at all latitudes and altitudes. Basic ecological processes are similar in temperate and tropical lakes, including grazing, competition, predation and abiotic adaptation. Small tropical lakes of intermediate age are probably not biotically more complicated than similar-sized temperate lakes. The structure of the areas of adaptative radiation and the dispersal ability of the species are important for the present distribution of taxa. Fish play a key role in the tropics since many species both consume zooplankton and compete with them for algal and pelagic sestonic food. This important co-evolution between fish and algae, leaving a fraction of the algal community with a predation refuge, may have decreased the ability of zooplankton to exploit algae. In addition, heavy predation from juvenile and adult fish may greatly simplify the zooplankton community, and have resulted in the scarcity of Cladocera, notably the efficient filter-feeder Daphnia. Little is known of possible physiological constraints to cladoceran distribution, however. Thus similar co-evolution as hypothesized between fish and algae seems not to have occurred to such a great extent between fish and zooplankton. Diurnal patterns in habitat selection of fish may also influence nutrient re-distribution in the tropics as in many temperate lakes. Serious environmental problems threaten tropical lakes, including eutrophication, clear-cutting of the rain forest, unwise introduction of new species not adapted to prevailing conditions, overfishing, extensive use of biocids, and probably acidic rain in areas with poorly buffered waters. Important processes in tropical lakes could be elucidated by concentrating research upon the fate of phytoplankton successional production, involving competition, grazing, sinking, fungi and bacterial attack. Co-evolution of fish and algae should be further investigated as it could in part explain the general scarcity and simplicity of the zooplankton community. Limnocorral experiments should also be used for further assessing processes in tropical lakes.     

Planktonic indices in the evaluation of the ecological status and the trophic state of the longest lake in Poland

Due to the intensive mixing polymictic lakes should be homogenous. However, morphometric diversity and high water dynamics contribute to the differentiation of many parameters in various areas of the lakes. This study analyzes both phytoplankton and zooplankton to assess differences in water quality along the north–south axis of the longest lake in Poland. New phytoplankton indicators were applied for determining the lake's ecological status: the Q index based on functional groups and the PMPL (Phytoplankton Metric for Polish Lakes) index based on phytoplankton biomass. TSI_ROT index (Rotifer Trophic State Index), which comprises the percentage of species indicating a high trophic state in the indicatory group and the percentage of bacteriovorus in the Rotifera population, was used for zooplankton analysis.TP content was different at different sites – we observed its gradual increase from the south to the north. Spatial variation of phosphorus did not considerably affect plankton diversity. The phytoplankton was dominated by Oscillatoriales, typical of shallow, well-mixed eutrophic lakes. The ecological status of the lake based on the EQR (Ecological Quality Ratio) was poor or moderate. The zooplankton was dominated by rotifers (at almost all sites), which indicates a eutrophic state of the lake. The values of phytoplankton indices at the studied sites did not differ considerably; the differences resulted more from local conditions such as the contaminant inflow and the macrophyte development than water dynamics.We have demonstrated that in the lake dominated by filamentous Cyanobacteria the ecological status should be determined according to the PMPL index or other indices dependent on the dominant Cyanobacteria species. Since the Q index does not include the functional group S1, the results can lead to the false conclusion that water quality improves with an increased amount of phytoplankton. The high abundance of Cyanobacteria in the lake may have contributed to the poor growth of rotifers.     

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 role of highly unsaturated fatty acids in aquatic foodweb processes

1. Polyunsaturated fatty acids (PUFA) are almost exclusively synthesized by plants. Animals can convert from one form of PUFA to another through elongation and desaturation, but very few can synthesize PUFA de novo. PUFA play an important role in regulating cell membrane properties, serve as precursors for important animal hormones and are essential for animals. 2. In aquaculture studies, highly unsaturated fatty acids (HUFA), a subset of PUFA, have been found to be critical for maintaining high growth, survival and reproductive rates and high food conversion efficiencies for a wide variety of marine and freshwater organisms. 3. The plankton literature suggests high food-quality algae species are rich in HUFA and low food-quality algae are poor in HUFA. Adding semi-pure emulsions of HUFA to algae monocultures can markedly increase the growth rates of zooplankton feeding on these mixtures. 4. A study measuring zooplankton biomass accrual when feeding on natural phytoplankton found a strong correlation between phytoplankton HUFA (specifically eicosapentaenoic acid) content and herbivorous zooplankton production. 5. The aquatic ecology literature suggests that planktonic foodwebs with high HUFA content phytoplankton have high zooplankton to phytoplankton biomass ratios, while systems with low HUFA phytoplankton have low zooplankton biomass. Also, the seasonal succession of plankton in many temperate lakes follows patterns tied to phytoplankton HUFA content, with intense zooplankton grazing and ‘clear-water-phases’ characteristic of periods when the phytoplankton is dominated by HUFA-rich species. 6. Herbivorous zooplankton production is constrained by the zooplankton’s ability to ingest and digest phytoplankton. It is becoming increasingly clear, however, that much of the phytoplankton which is assimilated may be nutritionally inadequate. HUFA may be key nutritional constituents of zooplankton diets, and may determine energetic efficiency across the plant–animal interface, secondary production and the strength of trophic coupling in aquatic pelagic foodwebs.     

Hydrobiological consequences of the addition of phosphate precipitants to inlet water of lakes

Summary In three model reservoirs (LUND, 1975) a method reducing bluegreen algal blooms in lakes was studied. Iron or aluminium were added to inlet waters for chemically binding the inflowing phosphorus.The research program, started in 1975, includes intensive monitoring of many chemical and hydrobiological variables, the determination of water and mass balances and since 1977 measurements or primary production rates with¹⁴C. In this paper only the results found in 1977 are discussed. An attempt is made to describe quantitatively how growth rates and changes in biomass are interconnected and how phosphorus precipitation changes these variables.In all reservoirs a large discrepancy was observed between the actual rate of increase in the algal population and the relative production rate. The latter appeared to be higher by one order of magnitude. The relative death rate due to grazing can account for the large difference between these growth rates only when selective grazing of zooplankton on phytoplankton is assumed.It can be concluded that treatment of inlet water with AVR, an aluminium salt, is unsuccessful in reducing algal development. Treatment with ironsulphate may be successful, but a reduction of the relative growth rates was not observed. The effects of grazing of zooplankton andDreissena polymorpha need further investigation.     

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.     

ANTARCTIC AQUATIC ECOSYSTEMS AS HABITATS FOR PHYTOPLANKTON

1. The Southern Ocean is a large-scale, relatively homogeneous upwelling ecosystem whose phytoplankton apparently grows suboptimally over much of its area. By contrast there is a wide variety of freshwater habitats in the Antarctic and in some of these phytoplankton growth efficiency is very high. The two habitats share similar temperature and irradiance regimes, but differ markedly in availability of inorganic nutrients, in grazing pressure and in the time- and space-scales on which various physical processes act. 2. Concentrations of inorganic nutrients in the marine ecosystem have been represented as being in excess of phytoplankton requirements, but the ionic composition of some nutrient pools may not conform to phytoplankton preferences. 3. Nutrient-limitation determines phytoplankton production in Antarctic lakes and gives rise to gross differences between lakes. 4. Irradiance in the water column varies greatly over the year in both marine and freshwater ecosystems. Most algae are shade-adapted, with the ability to utilize low irradiance but with sub-optimal response to high irradiance. However, local phytoplankton maxima may attain very high carbon fixation and growth rates. 5. Consistently low temperatures characterize both systems. Their effects on photo-synthetic carbon uptake mirror shade-adaptation. Division rates of marine phytoplankton may however be very much higher than predicted for ambient temperatures. 6. Vertical mixing is important in both ecosystems and influences the environment experienced by phytoplankton cells. This appears to have little effect on the average performance of phytoplankton in the strongly mixed surface water column of the Southern Ocean, where the mixed depth may exceed 100 m. This can be related partly to the shade-adapted photosynthetic response. Euphotic depths range from 20 to 100 m. 7. Strong vertical mixing under ice-free conditions in lakes may maximize photosynthetic efficiency, whilst distinct vertical stratification in permanently ice-covered lakes gives rise to segregation of nutrient uptake and regeneration. 8. Physical removal of phytoplankton biomass by grazing is locally important in the Southern Ocean, in contrast to the estimated mean mesoscale impact of grazing. Vertical sedimentation losses appear important in the context of mixing depth and generation time, and may be modified by vertical circulation of water. 9. Loss of phytoplankton biomass from lakes during the ice-free period is dominated by physical removal via the lake outflow. Grazing is generally unimportant, except where larvae of otherwise nektobenthic zooplankton hatch in synchrony with a phytoplankton maximum. Sedimentation is important under ice-cover.     

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.