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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.
In large and deep Lake Constance, total phosphorus concentrations during winter mixing (TPmix) were reduced by a factor of three (> 80 to ca. 30 ;g/l) from about 1979 to 1993. This resulted in an amplification and lengthening of phosphorus (P) depleted conditions throughout the season and water column. The response of the phytoplankton community depended on the time of the year and the level of aggregation under consideration. Total phytoplankton biomass quantified in terms of algal biovolume or chlorophyll concentrations decreased in summer, i. e. during the period of most severe P depletion, to about half of the original values during the first decade. In subsequent years, summer chlorophyll concentrations remained at this lower level whereas total biovolume increased again despite further decreases of TPmix. Average algal biomass in spring and autumn fluctuated without a distinct relationship to TPmix although P was depleted below the detection level during parts of these time intervals in recent years. This moderate response by community level parameters is attributed to changes in the temporal and internal organization of the algal community. Population dynamics and the relative importance of various taxonomical and functional groups such as mixotrophs and less-edible forms clearly changed in spring and summer. The renewed increase in algal biovolume in summer is mostly caused by species which are able to exploit additional P sources. For example, Dinobryon is an evidently mixotrophic organism which ingests P rich bacteria, its strongest competitors for soluble reactive phosphorus (SRP). Ceratium hirundinella might be migrating between the euphotic zone and deeper, P enriched water layers under suitable hydrodynamical conditions. At the level of genera and higher taxa, consistent trends in respect to TPmix were observed in spring and summer mostly indicating an adaptation to more oligotrophic conditions. In contrast, the functional group of well-edible algae showed little interannual variability and did not change in absolute numbers. This suggests that, in contrast to less-edible algae, well-edible forms are more strongly under top-down than bottom-up control, and that the nutritional basis of most herbivores changed less than it would be expected from the decrease in total algal biomass.