Priit Zingel

Contribution of different zooplankton groups in grazing on phytoplankton in shallow eutrophic Lake Võrtsjärv (Estonia)

The grazing impact of different sized zooplankton on ‘edible’ and total phytoplankton biomass and primary production was measured in L. Vortsjarv during a seasonal study in 1998 and 2000. The organisms of 48–100 μm size class, composed of ciliates and rotifers, contributed significantly to the total grazing of zooplankton community throughout the study period (average 68%). The average daily filtering and grazing rate of the whole zooplankton community (micro- and macro-zooplankton) remained low, corresponding to a filtration of 44% of the water volume, 4% of the total phytoplankton biomass and 29% of primary production. However, a strong grazing pressure on small-sized phytoplankton (<30 μm) was estimated in most of the study period (average 44% d−1). Among size classes of ‘edible’ phytoplankton, the size range 5–15 μm was the most important algal food for the dominant zooplankton grazers (herbivorous ciliates, Polyarthra spp., Chydorus sphaericus and Daphnia cucullata) in L. Vortsjarv.

Seasonal and annual population dynamics of ciliates in a shallow eutrophic lake

Seasonal population dynamics and community composition of planktonic ciliates were studied for over a decade in a naturally eutrophic shallow lake, Lake Vortsjarv, which has a large open surface area (270 km 2 on average) and highly turbid water (Secchi depth 300 µm) gymnostomes. There was a positive correlation between ciliates and phyto- and bacterioplankton, implying that the ciliates were clearly bottom-up or food-controlled. This was further confi rmed by the positive correlation between ciliates and metazooplankton. As both bacteria and phytoplankton are coupled with resuspension of lake sediments, the large fl uctuations in water level infl uenced the ciliate community biomass annually.

Vertical and Seasonal Dynamics of Planktonic Ciliates in a Strongly Stratified Hypertrophic Lake

Seasonal population dynamics and the vertical distribution of planktonic ciliates in a hypertrophic and strongly stratified temperate lake were studied from April to October in 2000 and from April to June in 2001. In the epi- and metalimnion the ciliate abundance peaked in spring and late summer, reaching maximum values in the metalimnion (86 cells ml−1) on 7th August 2000. In the epilimnion, the highest biomass content (414 µg C l−1) was observed on 8th May 2000. In the hypolimnion only a late summer peak occurred and the ciliate numbers were always lower than in the epi- and metalimnion. Five groups dominated the community of ciliates: Oligotrichida, Gymnostomatea, Prostomatida, Hymenostomata and Peritrichia, and the community composition varied greatly with depth. In the epilimnion the ciliate numbers were dominated by oligotrichs but small algivorous prostomatids, peritrichs and gymnostomes were also numerous. In the metalimnion these groups were gradually replaced by scuticociliates and mixotrophic Coleps spp. In the hypolimnion scuticociliates and species known as benthic migrants dominated. In the epilimnion and upper metalimnion in spring large herbivores and in summer small bacterivores were more numerous.

From bacteria to piscivorous fish: estimates of whole-lake and component-specific metabolism with an ecosystem approach.

The influence of functional group specific production and respiration patterns on a lakes metabolic balance remains poorly investigated to date compared to whole-system estimates of metabolism. We employed a summed component ecosystem approach for assessing lake-wide and functional group-specific metabolism (gross primary production (GPP) and respiration (R)) in shallow and eutrophic Lake Võrtsjärv in central Estonia during three years. Eleven functional groups were considered: piscivorous and benthivorous fish; phyto-, bacterio-, proto- and metazooplankton; benthic macroinvertebrates, bacteria and ciliates; macrophytes and their associated epiphytes. Metabolism of these groups was assessed by allometric equations coupled with daily records of temperature and hydrology of the lake and measurements of food web functional groups biomass. Results revealed that heterotrophy dominated most of the year, with a short autotrophic period observed in late spring. Most of the metabolism of the lake could be attributed to planktonic functional groups, with phytoplankton contributing the highest share (90% of GPP and 43% of R). A surge of protozooplankton and bacterioplankton populations forming the microbial loop caused the shift from auto- to heterotrophy in midsummer. Conversely, the benthic functional groups had overall a very small contribution to lake metabolism. We validated our ecosystem approach by comparing the GPP and R with those calculated from O2 measurements in the lake. Our findings are also in line with earlier productivity studies made with 14C or chlorophyll a (chl-a) based equations. Ideally, the ecosystem approach should be combined with diel O2 approach for investigating critical periods of metabolism shifts caused by dynamics in food-web processes.

Protozoan grazing in shallow macrophyte-and plankton lakes

The aim of the present study was to describe the role of ciliates and heterotrophic nanoflagellates in macrophyte and phytoplankton dominated shallow lakes and to compare the grazing on pico- and nano-sized particles by these protozoans. We sampled three shallow lakes, seasonally, one dominated by macrophytes, one by phytoplankton, and one with both conditions (two basins of this lake were sampled separately). The role of ciliates as consumers of bacteria and small algae was important in both lake types. In plankton dominated lakes the grazing rate of ciliates on bacteria was higher than in macrophyte dominated ones. But when ciliates were grazing on small algae, the highest rates were found in macrophyte lakes. Our results suggest that the microbial loop is weaker in macrophyte dominated lakes and grows stronger, when a lake becomes more turbid. The dominance of herbivorous ciliates in macrophyte lakes in summer and their high grazing rates on phytoplankton may have contributed to the decrease in phytoplankton abundance in these lakes. Heterotrophic nanoflagellates were relatively less abundant and, due to their low specific filtering rates, grazed only a minor fraction of the bacterioplankton.

The effects of predation by planktivorous juvenile fish on the microbial food web.

The feeding impact of planktivorous fish on microbial organisms is still poorly understood. We followed the seasonal dynamics of the food web in two natural fishponds for two years: one was stocked with planktivorous whitefish while the other had no planktivorous fish. The aim of the study was the simultaneous assessment of the feeding behaviours of planktivorous fish and of bacterivorous meta-/protozooplankters. We hypothesized that in the presence of planktivorous fish there would be fewer metazooplankton, more protozoans and decreased numbers of bacteria. Our results showed that the amount of metazooplankton eaten by the fish was indeed negatively correlated with metazooplankton biomass. The feeding impact of planktivorous fish in shaping the microbial loop was remarkable. The main grazers of bacteria in the fishpond were ciliates, whereas in the pond without fish these were heterotrophic nanoflagellates. In the fishless pond the role of the top predator shifted to the predaceous metazooplankter Leptodora kindtii which controlled the abundance of herbivorous metazooplankters. We found a negative relationship between the number of bacteria and flagellates in the fishless pond, while the number of bacterivorous ciliates was suppressed by predaceous ciliates. Therefore the bacteria-grazing activity was higher in the absence of planktivorous fish.

Ciliate communities of a large shallow lake: association with macrophyte beds.

We investigated the influence of macrophyte composition on ciliate community structure in a large, shallow, eutrophic Lake Võrtsjärv. We hypothesized that macrophyte composition must have strong influence on the dispersal of ecologically different ciliate groups in a shallow lake and that more diverse macrophyte stands cause also a greater diversity in the ciliate community. In Võrtsjärv macrophyte distribution is spatially strongly polarized both in east-west and north-south directions in relation to abiotic factors. Phragmites australis and Myriophyllum spicatum were the most widespread species occurring in most parts of the lake. Correlation of environmental, macrophyte and planktonic ciliate variables confirmed the suggested spatial gradients. More diverse macrophyte stands supported a high species richness and abundance of epiplanktonic community but showed negative influence on the number and abundance of euplanktonic ciliate taxa. Opposite trends were found relative to the abundance of P. australis. Benthic ciliates showed a similar distribution pattern to euplanktonic taxa being most abundant in sites were the Shannon-Weaver index for macrophytes was low. Strong polarizing effect of the lakes vegetation on planktonic ciliate diversity was reflected in correlations of the number of ciliate taxa as well as the numbers of eu- and epiplanktonic taxa with geographic co-ordinates.

The influence of zooplankton enrichment on the microbial loop in a shallow, eutrophic lake.

With increasing primary productivity, ciliates may become the most important members of the microbial loop and form a central linkage in the transformation of microbial production to upper trophic levels. How metazooplankters, especially copepods, regulate ciliate community structure in shallow eutrophic waters is not completely clear. We carried out mesocosm experiments with different cyclopoid copepod enrichments in a shallow eutrophic lake to examine the responses of ciliate community structure and abundance to changes in cyclopoid copepod biomass and to detect any cascading effects on bacterioplankton and edible phytoplankton. Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by the decline of predaceous ciliate abundance, which would otherwise graze effectively on the small-sized ciliates. The inverse relationship between crustacean zooplankton and large predaceous ciliates is an important feature adjusting not only the structure of the ciliate community but also the energy transfer between meta- and protozooplankton. Still we could not detect any cascading effects on bacterio- or phytoplankton that would be caused by the structural changes in the ciliate community.

Quick springtime changes in community structure of pelagic ciliated protozoa

Quick changes in community composition and population dynamics of planktonic ciliates were studied in the ponds of two fish farms obtaining water from different sources (from a fast flowing river or a stillwater reservoir). Samples were taken at two-day intervals during a short period after springtime filling of the ponds. At the beginning of the investigation period the community in the lotic ponds was dominated by species that favour periphyton or benthos as their habitat. They were gradually replaced by euplanktonic species. In the lentic ponds euplanktonic species dominated throughout our study. The similarity index ISj between lotic ponds and their inflow was 0.67 at the start of our study but decreased to 0.2 during 14 days. If the inflow was from a stillwater reservoir, the similarity index decreased only moderately. In the lotic ponds ciliate community shifted from bacterivory to algivory while in the case of lentic ponds the situation was the opposite - algivorous species were gradually replaced by bacterivores. Abiotic and biotic factors that determine ciliate community structure were separated. The abiotic features are ruling when the system goes through major changes (in our study the lotic pond system where water flow ceased dramatically). The biotic factors are crucial in more stable systems (in our study the lentic pond system) and can change the community structure in same degree as the abiotic ones.

Effects of nutrient and water level changes on the composition and size structure of zooplankton communities in shallow lakes under different climatic conditions: a pan-European mesocosm experiment

Lentic ecosystems act as sentinels of climate change, and evidence exists that their sensitivity to warming varies along a latitudinal gradient. We assessed the effects of nutrient and water level variability on zooplankton community composition, taxonomic diversity and size structure in different climate zones by running a standardised controlled 6-months (May to November) experiment in six countries along a European north–south latitudinal temperature gradient. The mesocosms were established with two different depths and nutrient levels. We took monthly zooplankton samples during the study period and pooled a subsample from each sampling to obtain one composite sample per mesocosm. We found a significant effect of temperature on the community composition and size structure of the zooplankton, whereas no effects of water depth or nutrient availability could be traced. The normalised size spectrum became flatter with increasing temperature reflecting higher zooplankton size diversity due to higher abundance of calanoid copepods, but did not differ among depths or nutrient levels. Large-bodied cladocerans such as Daphnia decreased with temperature. Taxonomic diversity was positively related to size diversity, but neither of the two diversity measures demonstrated a clear pattern along the temperature gradient nor with nutrient and water levels. However, genus richness decreased at the warm side of the temperature gradient. Our experiment generally supports recent empirically based findings that a continuing temperature increase may result in lower genus richness and lower abundance of large-sized zooplankton grazers, the latter likely resulting in reduced control of phytoplankton.