Impacts of nutrients and zooplankton on the microbial food web of an ultra-oligotrophic lake

In ultra-oligotrophic lakes and the sea, calanoid copepods arethe dominant mesozoo-plankton and cladocerans are generallysparse or absent. To determine the effects of predation andnutrient enrichment on the pelagic microbial food web of anultra-oligotrophic lake, we added copepods and cladocerans atlow biomasses (<60 µg l^–1 to in situ enclosuresin Lake Wakatipu, New Zealand, in the presence and absence ofadded nutrients (nitrogen and phosphorus). In response to nutrientfertilization, the concentrations of phototrophs >3 µmand heterotrophic bacteria increased by 50 and 15%, respectively,over 4 days, but those of cyanobacterial picoplankton decreasedby 68%. The presence of calanoid copepods (Boeckella dilatata)at ambient densities (1 and 4 l^–1) rapidly and severelysuppressed ciliate population growth over 4 days and also loweredthat of flagellates >3 µm, even when microbial growthwas enhanced by added nutrients. The presence of a small cladoceran,Ceriodaphnia dubia, at double the densities, but similar biomasses,to those of copepods, depressed the net growth rates of ciliatesand flagellates to a lesser degree. The net growth rate of heterotrophicbacteria after 4 days declined with flagellate abundance, consistentwith the possibility of regulation by flagellates. Althoughbacteria and algae increased in response to nutrient fertilization(bottom-up control), predation (top-down control) appeared toplay an important role in structuring the microbial food webof this ultra-oligotrophic lake in summer.     

Modelling nutrient cycles in relation to food web structure in a biomanipulated shallow lake

The modelPCLAKE describes the phosphorus and nitrogen cycles within a shallow lake ecosystem, including the sediment and a simplified biological food web. All components are modelled in a generalized way rather than a very detailed one. This model has been applied to Lake Zwemlust, a small biomanipulated lake in The Netherlands. Formerly, this highly eutrophic lake was dominated by cyanobacteria and devoid of macrophytes. Biomanipulation was carried out in 1987 by pumping-out of the water, removal of all fish, and refilling of the lake with seepage water. The lake was restocked with some rudd, pike, zooplankton and seedlings of macrophytes, and then monitored up to 1992. Macrophytes developed rather quickly and reached their maximum biomass during the six-years period in 1989. Despite the continuously high nutrient (N and P) loading, algal biomass remained low due to nitrogen limitation, caused by competition with the macrophytes. From 1990 onwards, the macrophytes declined again and a species shift occurred, following an increase of herbivorous birds on the lake and the development of herbivorous fishes.Model simulations grossly reproduced the observed developments in Lake Zwemlust before and after the biomanipulation measures. The existence of multiple steady states at the same trophic state and the possible shift between them could be simulated well. This study also demonstrates the interrelation between system structure and the distribution and cycling of nutrients. It is concluded, that within general boundary conditions set by the trophic state of the system, the food web structure determines the actual nutrient flows and the occurrence of nutrient limitations of the primary producers. It is shown that both aspects can be integrated in one mathematical model. The long-term stability of the macrophyte dominance in the lake is discussed.     

Responses to food web manipulation in a shallow waterfowl lake

The effects of fish stock reduction have been studies in 3 Dutch lakes (Lake Zwemlust, Lake Bleiswijkse Zoom and Lake Noorddiep) and 1 Danish lake (Lake Væng) during 4–5 years. A general response id described. The fish stock reduction led in general to a low fish stock, low chlorophyii-a, high transparency and high abunuance of macrophytes. Large Daphnia became abundant, but their density decreased, due to food limitation and predation by fish. The total nitrogen concentration became low due to N-uptake by macrophytes and enhanced denitrification. In Lake Bleiswijkse Zoom the water transparency deteriorated and the clear water state was not stable. The fish stock increased and the production of young fish in summer was high. lear water occurred only in spring. Large daphnids were absent in summer and the macrophytes decreased.In Lake Zwemlust, Lake Væng and Lake Noorddiep the water remained clear during the first five years. In summer of the sixth year (1992) transparency decreased in Lake Zwemlust (with high P-concentration of 1.0 mg P l^-1). Also in Lake Væng (with a low nutrient concentration of 0.15 mg P.^-1) a short term turbid stage (1.5 month) occurred in summer 1992 after a sudden collapse of the macrophytes. Deterioration of the water quality seems to start in summer and seems related to a collapse in macrophytes. At a low planktivorous fishstock (e.g. Lake Væng)thhe duration of the turbid state is shorter. than in presence of a high planktivorous fish biomass (e.g. Lake Zwemlust, and later years of Lake Bleiswijkse Zoom).     

Structure of planktonic microbial food web in a brackish stratified Siberian lake

The distribution of primary components of the microbial community (autotrophic pico- and nanoplankton, phototrophic bacteria, heterotrophic bacteria, microscopic fungi, heterotrophic flagellates, ciliates and heliozoa) in the water column of Lake Shira, a steppe brackish-water, stratified lake in Khakasia, Siberia (Russia), were assessed in midsummer. Bacterioplankton was the main component of the planktonic microbial community, accounting for 65.3 to 75.7% of the total microbial biomass. The maximum concentration of heterotrophic bacteria were recorded in the monimolimnion of the lake. Autotrophic microorganisms contributed more significantly to the total microbial biomass in the pelagic zone (20.2–26.5%) than in the littoral zone of the lake (8.7–14.9%). First of all, it is caused by development of phototrophic sulphur bacteria at the oxic-anoxic boundary. The concentrations of most aerobic phototrophic and heterotrophic microorganisms were maximal in the upper mixolimnion. Heterotrophic flagellates dominated the protozoan populations. Ciliates were minor component of the planktonic microbial community of the lake. Heterotrophic flagellates were the most diverse group of planktonic eucaryotes in the lake, which represented by 36 species. Facultative and obligate anaerobic flagellates were revealed in the monimolimnion. There were four species of Heliozoa and only three of ciliates in the lake.     

Evaluation of Microcystis as food for zooplankton in a eutrophic lake

Four experiments were conducted to evaluate Microcystis as food for zooplankton in Lake Kasumigaura, and the following results were obtained. (1) Moina micrura (Cladocera) showed little growth and no reproduction when the animal was reared with Microcystis cultured in the laboratory. The animal did not grow nor reproduce well when Chlorella was mixed with Microcystis as food. (2) Moina micrura assimilated Microcystis much less than Chlorella when the animal fed on single species of Microcystis or a mixture with Chlorella. (3) Microcystis collected from Lake Kasumigaura could not be utilized by Moina micrura even though the colonies were broken up into edible sizes. However, the alga turned into utilizable food when it was decomposed. (4) No inhibitors of Moina micrura population growth could be found in the non-filtered water of Lake Kasumigaura where Microcystis was blooming heavily. Decomposed Microcystis seemed to be utilized by zooplankton as an important food source in Lake Kasumigaura.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

Macroinvertebrate food web structure in a floodplain lake of the Bolivian Amazon

Two stable isotopes δ¹³C and δ¹⁵N were used to identify the energy sources and trophic relationships of the main freshwater macroinvertebrates in a floodplain lake of the Beni River (Bolivian Amazonia). Four energy sources (seston, bottom sediment, periphyton, and aquatic macrophytes) and macroinvertebrate communities were collected during three periods of the river hydrological cycle. Macroinvertebrates showed greater temporal variation in isotope values than their food sources. Six trophic chains were identified: four were based on seston, periphyton, C₃ macrophytes, and bottom sediments, and the last two chains on a combination of two carbon sources. One mixed seston and periphyton sources during the wet season while the other mixed periphyton and macrophytes sources during the wet and dry seasons. Periphyton was the most important energy source supporting the highest number of trophic levels and consumers. The macrophytic contribution was only significant during the dry season. Bottom sediments constituted a marginal energy source. As each season is associated with different physical and chemical conditions, processes organizing macroinvertebrate food web structure in the Beni floodplain seem strongly linked to hydrological seasonality.     

Experimental manipulations of microbial food web interactions in a humic lake: shifting biological drivers of bacterial community structure

A previous multiyear study observed correlations between bacterioplankton community composition (BCC) and abundance and the dynamics of phytoplankton populations and bacterivorous grazers in a humic lake. These observations generated hypotheses about the importance of trophic interactions (both top-down and bottom-up) for structuring bacterial communities in this lake, which were tested using two multifactorial food web manipulation experiments that separately manipulated the intensity of grazing and the composition of the phytoplankton community. Our results, combined with field observations, suggest that a hierarchy of drivers structures bacterial communities in this lake. While other studies have noted links between aggregate measures of phytoplankton and bacterioplankton communities, we demonstrate here correlations between succession of phytoplankton assemblages and BCC as assessed by automated ribosomal intergenic spacer analysis (ARISA). We used a novel approach linking community ARISA data to phylogenetic assignments from sequence analysis of 16S rRNA gene clone libraries to examine the responses of specific bacterial phylotypes to the experimental manipulations. The synchronous dynamics of these populations suggests that primary producers may mediate BCC and diversity through labile organic matter production, which evolves in quality and quantity during phytoplankton succession. Superimposed on this resource-mediated control of BCC are brief periods of intense bacterivory that impact bacterial abundance and composition.     

Relative impacts of copepods, cladocerans and nutrients on the microbial food web of a mesotrophic lake

Calanoid copepods, rather than cladocerans, frequently dominatethe zooplankton of lakes in New Zealand. The potential consequencesof this domination for the microbial community of mesotrophicLake Mahinerangi, New Zealand, were determined by field experimentsin which Boeckella and Daphnia were added to in situ enclosuresin the presence and absence of added nutrients. Boeckella hamataat ambient densities (2 and 81^–1) rapidly and severelysuppressed ciliate population growth over 4 days, even whenmicrobial growth was enhanced by added nutrients, but effectsof copepods on other components of the microbial community (bacteria,photosynthetic picoplankton, heterotrophic nano-flagellates,algae) were slight. In contrast, Daphnia carinata at the samedensities (but 3-fold higher biomasses per litre) had a relativelyweak effect on ciliates, suppressing ciliate abundance onlyafter 4 days at 8 Daphnia 1^–1 (330 µg 1^–1);this daphniid density also depressed abundances of large bacterialrods, some photosynthetic picoplankton and the dominant alga,Cyclotella. These results highlight the relative importanceof specific trophic linkages in a microbial food web; they alsosuggest that the dominance of Boeckella in many southern hemispherelakes may account for relatively low ciliate abundances in theselakes.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Spatial and temporal dynamics of the steady-state phytoplankton assemblages in a temperate shallow hypertrophic lake (Lake Manyas,Turkey)

Spatial and temporal dynamics of phytoplankton biomass and species composition in the shallow hypertrophic Lake Manyas, Turkey, were studied biweekly from January 2003 to December 2004 to determine steady-state phases in phytoplankton assemblages. Steady-state phases were defined when one, two or three coexisting species contributed to at least 80% of the standing biomass for at least 2 weeks and during that time the total biomass did not change significantly. Ten steady-state phases were identified throughout the study peiod. During those periods, Achnanthes microcephala (Kützing) Cleve twice dominated the phytoplankton biomass alone and contributed to more than 50% of the total biomass in seven phases. Microcystis aeruginosa (Kützing) Kützing, Anabaena spiroides Klebahn, Cyclotella stylorum Brightwell, Pediastrum boryanum (Turpin) Meneghini and Phacus pusillus Lemmermann were also represented once in steady-state phytoplankton assemblages. A. microcephala was dominant usually during cold periods of the year, while M. aeruginosa and A. spiroides were usually dominant in warm seasons. The total number of species showed a clear decrease during steady-state phases at all stations. All stations were significantly different in terms of the measured physical and chemical parameters (P < 0.05) and phytoplankton biomass (F = 117, P < 0.05).     

The structure of a plant-pollinator food web

The pollination biology literature is dominated by examples of specialization between plants and their pollinators. However, a recent review shows that it is generalization that prevails in the field, with most plants having a number of pollinators and most pollinators visiting a number of plants. Consequently, the vast majority of plant–pollinator interactions are embedded in a complex web of plant–pollinator interactions. These plant-pollinator webs can be studied in the manner of conventional food webs and the aim of this paper is to illustrate how contemporary methods of web construction and analysis can be applied to plant-pollinator communities.     

Trophic interactions within the microbial food web in a tropical floodplain lake (Laguna Bufeos, Bolivia)

Whether the primary role of bacterioplankton is to act as "remineralizers" of nutrients or as direct nutritional source for higher trophic levels will depend on factors controlling their production and abundance. In tropical lakes, low nutrient concentration is probably the main factor limiting bacterial growth, while grazing by microzooplankton is generally assumed to be the main loss factor for bacteria. Bottom-up and top-down regulation of microbial abundance was studied in six nutrient limitation and dilution gradient-size fractionation in situ experiments. Bacteria, heterotrophic nanoflagellates (HNF), ciliates and rotifers showed relatively low densities. Predation losses of HNF and ciliates accounted for a major part of their daily production, suggesting a top-down regulation of protistan populations by rotifers. Phosphorus was found to be strongly limiting for bacterial growth, whereas no response to enrichment with Nitrogen or DOC was detected. HNF were the major grazers on bacteria (g-0.43 d(-1)), the grazing coefficient increased when ciliates were added (g- 0.80 d(-1)) but decreased when rotifers were added (g- 0.23 d(-1)) probably due to nutrient recycling or top-down control of HNF and ciliates by rotifers.     

Stress and disturbance in the phytoplankton community of a shallow,hypertrophic lake

The validity of Connell's intermediate disturbance hypothesis in phytoplankton communities was tested on data from a hypertrophic, shallow lake, Hjarbæk Fjord, Denmark.The present data from Hjarbæk Fjord demonstrate the difficulties in distinguishing stress from disturbance in a phytoplankton community, and show that great changes in the phytoplankton community can take place within few days.A collapse of blue-green algae in late June 1986 caused remineralization of nutrients and resulted in a rapid increase of fast-growing small chlorococcal green algae and phytoplankton species diversity, without any external disturbances acting on the lake. External disturbances in the form of wind action and brackish water intrusion occurred several days after the onset of these events. Carbon depletion and pH 11.0 were severe stress factors on the phytoplankton community. They were induced by calm, warm weather, but eventually acted as a kind of disturbance to the normally well circulated lake.     

Grazing and food size selection by crustacean zooplankton compared to production of bacteria and phytoplankton in a shallow Norwegian mountain lake

Food size selection by suspension-feeding crustaceans in a shallowmountain lake was investigated with radiolabelled algae andbacteria, and with a series of monodisperse fluorescent latexbeads with diameters ranging from 0.25 to 5.18 µm. JuvenileDaphnia longispina was a more efficient grazer on bacteria-sizedparticles than adults. Bosmina longispina and Cyclops scutiferwere both able to ingest bacteria-sized food particles. Thecrustaceans harvested {small tilde} 100% of the primary productionduring the day investigated, but only 20% of the bacterial production.To keep the bacterial biomass in or near steady state, the microzooplanktonmust be harvesting most of the bacterial production.     

Phytoplankton primary production in a shallow,well- mixed,hypertrophic South African lake

This paper reports on a two-year analysis of the wind climateand its effect on phytoplankton primary production in ashallow (mean depth = 1.9 m), hypertrophic South Africancoastal lake, Zeekoevlei. The lake is subject to continuousmixing of the euphotic zone (Z _eu = 0.8 m), andcomplete mixing of the water column to the mean depth on adaily basis. Median annual wind speeds, prevailing fromeither the north or the south, were 6.4 m s^–1. There wasan almost total absence of calms, measured as hourly meanwind speeds of <1 m s^–1. Notwithstanding the highfrequency of mixing, the lake supports a dense population ofphytoplankton, dominated by Cyanophyte and Chlorophytespecies. Mean concentrations of chlorophyll-a were240 g l^–1. The attenuation of photosyntheticallyavailable radiation, PAR, was high, with mean K _dvalues of 6.4 m^–1 and water transparencies of <0.5 m.Levels of primary productivity, determined using the lightand dark bottle oxygen method, were very high, comparable toor exceeding that of the most productive systems yet studied.Maximum volumetric productivity ranged from 525 to 1524 mg Cm^–3 h^–1, and was confined to the upper 0.5 m of thewater column. Daily areal productivity, P _d,varied between 1.2 and 4.3 g C m^–2 d^–1, and that ofthe maximum chlorophyll-a specific photosynthetic rate,P _{B max}, between 1.6 and 7.9 mg C (mgChl-a)^–1 h^–1. Primary production was limited bywater temperature and the attenuation of PAR. The highfrequency of wind-induced mixing resulted in regular mixingof the phytoplankton through the euphotic zone, and reducedthe overall importance of P _max at a single layer inthe depth profile. Similarly, the regularity of mixing wasrecognized as a limitation of the incubation of bottle chainsto determine primary production levels.     

Spatial and temporal distribution of fish and zooplankton in a shallow lake

Summary 1. We performed both a large- and a small-scale echo sounding study on the spatial and temporal distribution of fish (mainly roach, Rutilus rutilus and perch, Perca fluviatlis ), as well as a small-scale study of zooplankton distribution in the small, shallow and eutrophic Lake Hanebjerg in Denmark. In the small-scale study, sampling was conducted in open water as well as in the edge zone immediately outside two different types of vegetation.
2. Fish daytime abundances differed between the northern and the southern parts of the lake and, on a small scale, small fish aggregated in the edge zones during day, preferably outside dense emergent vegetation. Copepods avoided emergent vegetation, while cladocerans showed no habitat preference. Both small fish and cladoceran numbers were found to be higher during night than day.
3. The relative abundance (number per sample) of cladocerans in the edge zone immediately outside vegetation was negatively correlated with the relative abundance of fish in that zone. There was no correlation between cladoceran and fish abundance in open water, or between the relative abundance of copepods and fish.
4. The presence of pelagic piscivores in combination with avoidance behaviour of both fish and zooplankton is a likely explanation for the observed distribution of small fish and cladocerans in Lake Hanebjerg. Both small- and large-scale distribution patterns may be dependent on the type and distribution of complex structure in the lake. Even in a small lake, large-scale patterns may affect the interpretation of small-scale data.     

Trophic web structure in a shallow eutrophic lake during a dominance shift from phytoplankton to submerged macrophytes

In Lake Krankesjön, southern Sweden, sago pondweed (Potamogeton pectinatus L.) and a stonewort (Chara tomentosa L.) expanded spatially during the second half of the 1980's after more than a decade of phytoplankton blooms and sparse submerged vegetation. During the expansion of submerged plants the number of resting and breeding waterfowl increased. The increase was significant for herbivorous birds such as coot (Fulica atra L.) and mute swan (Cygnus olor (Gmelin)), but also for omnivorous dabbling ducks. The shift from phytoplankton to submerged macrophytes caused structural changes on higher trophic levels, and an altered trophic web developed. The density of planktonic Cladocera decreased, which is suggested to be a result of decreased phytoplankton productivity and biomass as nutrient levels dropped. The benthic macroinvertebrate assemblage changed from low diversity and biomass dominated by Chironomidae and Oligochaeta on bare sediment, to high diversity and biomass characterized by plant-associated forms like snails and isopods in areas covered by macrovegetation. The mean size of perch (Perca fluviatilis L.) increased, probably as a result of higher availability of macroinvertebrates in the vegetation. The perch reached a mean size where the species is known to shift to a fish diet, permitting an increased top down effect on the ecosystem. The results support the idea that shallow eutrophic lakes can shift between two states, each one stabilized by feed-back mechanisms including both biotic and abiotic factors. Shifts between these states are suggested to be a possible explanation for observed drastic changes in abundance of waterfowl in shallow eutrophic lakes.