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Large (472 km2) and deep (zmean=101 m) Lake Constance is undergoing re-oligotrophication. Total phosphorus during winter mixing (TPmix) decreased from >80 during 1975-1981 to 22 ;g/l in 1996. Average summer values of secchi and euphotic depth increased significantly from 4.5 to 6.5 m and from 10.5 to 13 m, respectively. The algal species composition changed and, during summer, total algal biomass decreased by 50 % and primary production by 25 %. Standing stocks of well-edible algae, rotifers, and herbivorous and carnivorous crustaceans did not exhibit a trend with TPmix, whereas their species compositions or egg-ratios were partially altered. The age-at-capture of planktivorous whitefish increased slightly. I tested the hypotheses that (1) changes should first be observed at the level of individuals or within species (altering e. g. C:P or egg-ratios) prior to changes within communities (affecting e. g. the taxonomic composition) and at the community level (affecting e. g. total biomass or production). This would imply that it is more appropriate to conceptualize step-wise responses along a hierarchical gradient of increasing aggregation as suggested by hierarchy theory, rather than simultaneous changes at all hierarchical levels. (2) Responses become dampened along the food chain and with increasing body size, i. e. bottom-up control is most important for autotrophs. All communities studied (phytoplankton, crustaceans, fish) reacted at the individual level (e. g. by changes of (re)production rates), and/or within the community (e. g. altered taxonomic composition) whereas changes of bulk parameters of the entire community were restricted to phytoplankton. Hence, the first hypothesis is partially supported by the observed reactions and demands further testing. The second hypothesis is clearly supported by our data when comparing autotrophs and consumers, but not when comparing crustaceans and fish. The testing of these hypotheses is complicated by the large differences in size and, consequently, in reaction times of pelagic organisms on the one hand and the rather fixed time scale of limnological research on the other hand. The different time scales imply a selective perception of the various potential responses of the differently sized organisms as the time scales of the responses depend on body size and the level of aggregation. For example, we are more likely to establish physiological or behaviourial changes of fish, and taxonomical or biomass changes of phytoplankton. Acknowledging the scale dependence and level of aggregation is also crucial for cross-system comparisons.
The ecology of chroococcoid picocyanobacteria was studied from 1987 to 1997 in large, deep, mesotrophic Lake Constance in relation to various abiotic and biotic factors that may influence their population dynamics. Picocyanobacteria dominated the autotrophic picoplankton (APP) numerically in this lake at all depths and times. Their abundances did not respond unequivocally to the decline of winterly phosphorus concentrations by a factor of 2.5 during the decade of investigation. They showed a recurrent seasonal pattern with peaks in spring and late summer, interspersed by a pronounced minimum during and after the clear-water phase around June. The magnitude, timing, and number of peaks and troughs which varied interannually, could in part be related to weather conditions or the impact of other plankton groups. Larger phytoplankton and picocyanobacteria exhibited a distinct and predictable response to the vertical mixing intensity during early spring. Except for 1993, picocyanobacteria and larger phytoplankton decreased simultaneously during the mass development of daphnids in late May or June which gave rise to the clear-water phase. As the daphnid development depends more strongly on surface water temperature than on vertical mixing intensity an early onset of stratification may imply a longer spring development which contributed to a higher seasonal average of picocyanobacterial abundances in 1989-1991. The decline in picocyanobacteria around the clear-water phase was often more pronounced and lasted longer than did the decline in larger algae. The rate of decrease may be related to daphnid abundance, however, no such relationship existed in respect to its duration. Summer peaks of picocyanobacteria were recorded despite the presence of relatively high densities of daphnids. We conclude that with the exception of the clear- water phase, grazing control by nano- and microzooplankton may be more important for controlling picocyanobacterial numbers than is grazing by daphnids. Picocyanobacteria declined in autumn prior to or concomitant with larger algae without any obvious relationship to phytoplankton biovolume or the extent of vertical mixing within the uppermost 20 m. The as yet unexplained variation in the population dynamics of picocyanobacteria points to the significance of species- specific protist grazing and to shifts in picocyanobacterial species composition which should be tackled in future studies.
In Lake Constance, phytoplankton productivity, together with parameters relevant for the production process, was assessed year-round at about 500 dates between 1980 and 1995/1996. During this period, the concentration of total phosphorus during winter circulation decreased from more than 80 to 22 ;g/l as a consequence of sewage diversion and waste water treatment within the catchment area. By contrast, annual photosynthetic rates remained virtually unchanged for about 10 more years following phosphorus decline (mean value 288 " 21 g C m-2 a-1), and thereafter decreased only by about 25 % until 1996. The aim of this study is to analyze factors responsible for this pronounced resilience.