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Long-term measurements (1979-1994) of meteorological parameters and of algal and crustacean biomass were used in conjunction with a comprehensive hydrodynamic model to evaluate the impact of weather conditions on plankton dynamics in a large, deep, temperate lake (Upper Lake Constance), and to identify potential causal mechanisms. The natural variability of weather conditions, including the exceptionally mild winters during the late eighties, allowed us the investigation of the covariation of meteorological parameters such as irradiance, air temperature, and wind with vernal algal and crustacean population growth. Crustacean zooplankton responded strongly to differences in surface water temperature, but not to mixing depth or algal biomass. Clear relationships between changes of algal biomass and meteorological factors were only found during the rare occasions when acted together to favour or hamper algal development. Otherwise, the impact of meterological conditions on the physical conditions which were most likely conducive to phytoplankton development, could not be followed by this simple approach. This problem was overcome with a one-dimensional hydrodynamic turbulent exchange model driven by the meteorological boundary conditions at the water surface. It was used to simulate the development of the vernal density stratification and to investigate the relationships between meteorological conditions and exchange rates from the euphotic to the aphotic zone. The beginning of the spring algal bloom was shown to depend on the stabilization of the upper part of the water column. As soon as mixing below 20 m was inhibited, confining the algae to the euphotic zone for prolonged periods of time, substantial increases in algal standing stock occurred consistently. In contrast, during periods when high vertical mixing rates were computed with the model no substantial increases of algal biomass were found. This tight coupling between the estimates of vertical mixing intensity and observed algal development, combined with knowledge about the impact of individual meteorological factors on mixing, enabled predictions about the response of algae to different weather conditions during spring.
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.
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.
Seasonal changes of the impact of daphnids on the plankton biomass size distribution, the biomass within individual size ranges, the average predator-prey weight ratios, and the efficiency to transfer matter and energy from small to large organisms are analyzed in large and deep Lake Constance based on comprehensive long-term observations. A comparison of daphnid biomass and production with those of other herbivorous groups (i. e. ciliates, rotifers, herbivorous crustaceans) reveals that in early spring daphnids play a minor role in relative and absolute values as compared to small fast growing ciliates. During this time, small algae and ciliates dominate which gives rise to a decreasing Sheldon-type size spectrum, low predator-prey weight ratios, and a low transfer efficiency along the size gradient. Around June, daphnids reach maximum abundances and become keystone species for the shape of the biomass size distribution, the food web structure, and the energy flow. They accumulate biomass in their size range one order of magnitude above the average. The slope of the normalized biomass size spectrum is less negative and positively correlated with daphnid biomass if the latter exceeds about 200 mg C/m2. This indicates a more efficient transfer along the size gradient with high predator-prey weight ratios and high trophic transfer efficiencies. The coefficients of determination of regression lines fitted to size distributions decrease with daphnid abundance, i. e. the size spectra become more irregular when daphnids dominate. In midsummer, daphnids lose their dominance and coexist with other herbivores (especially ciliates) in a highly diverse plankton community. The latter gives rise to a relatively smooth and almost flat Sheldon-type size distribution, lower predator-prey weight ratios, and a slightly reduced transfer efficiency along the size gradient. In late spring/early summer, negative relationships are found between daphnid biomass and the biomasses in the size ranges of autotrophic picoplankton, small phytoplankton, heterotrophic flagellates, and small and medium sized ciliates (0.06 - 32 pg C and 100-30,000 pg C). In mid- and late summer or on annual average, hardly any of these relationships existed. This cannot solely be attributed to lower daphnid abundance but points also to a more diverse control of small plankton organisms including nutrient limitation in summer. Ciliates influence the slope and shape of the size distribution much less than daphnids although they are at least of equal importance as daphnids in respect to herbivory and related fluxes in Lake Constance on annual average. The findings on the impact of daphnids on the energy flow within the plankton food web derived from size distributions are compared to, and are consistent with results obtained by mass-balanced carbon flow diagrams.
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.
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