Diversity of catechol 2,3-dioxygenase genes of bacteria responding to dissolved organic matter derived from different sources in a eutrophic lake

Catechol 2,3-dioxygenase (C23O) is an extradiol dioxygenase that plays an important role in degrading aromatic compounds such as those found at polluted sites. However, little is known about the diversity of C23O genes in unpolluted environments. In such environments, various factors, including the quality and quantity of dissolved organic matter (DOM), could influence the composition and behaviour of bacterial community possessing C230 genes. We investigated C23O genes in bacteria responding to DOM from various sources in a eutrophic lake by PCR and cloning. Six microcosms filled with lake water containing indigenous bacteria and DOM from different sources were incubated for 10 days. After 1 or 2 days of incubation, C23O genes were detected in the microcosms enriched with DOM recovered from inflow river water and humus from reed grass. The sequences were very diverse but had features conserved in extradiol dioxygenases. The clone libraries generated on day 2 showed distinctive compositions among microcosms, indicating that bacteria possessing a variety of C23O genes responded differently to DOM from different sources. After 10 days of incubation, C23O genes in a previously unidentified gene cluster, 'Cluster X', became dominant in the libraries.     

Metabolic rate estimates for a eutrophic lake from diel dissolved oxygen signals

Estimates of net primary production, community respiration (R'), and gross primary production (P_g) are developed and presented for the productive layers of eutrophic Onondaga Lake, NY, U.S.A., for time scales ranging from diel to several months, based on 4 months of robotic diel profiles of dissolved oxygen (DO) and temperature. Metabolic rate calculations are made through application of a DO mass balance framework that also accommodates inputs and losses of DO mediated by exchange across the air–water interface and across the lower boundary of the productive layers. It is demonstrated that the dynamics of the flux across the air–water interface are important to the metabolic rate estimates, while vertical mixing-based losses to the underlying layers can be ignored. Study average estimates of R' (1.49 g O₂ m^–3 d^–1) and P_g (1.60 g O₂ m^–3 d^–1) obtained by this non-isolated community approach are consistent with levels reported in the literature for similar chlorophyll a concentrations, based on isolated community (bottle experiment) protocols to measure these metabolic rates. The non-isolated community approach is shown to have limited utility for quantifying day-to-day changes in these rates in this lake, apparently because of horizontal exchange with waters of different DO concentrations. However, this approach may support reliable estimates of metabolic rates at intermediate time scales; e.g., several days to a week. The DO mass balance framework is demonstrated to be valuable in resolving the relative roles of various physical and biological processes in regulating the DO pool of the productive layers.     

Uptake and transport of phosphorus by Agrostis capillaris seedlings from rapidly hydrolysed organic sources extracted from32 P-labelled bacterial cultures

Cultures of the soil bacterium Serratia liquifaciens grimesii were grown with³² P labelled phosphate, to produce a uniformly³² P labelled source of microbial P. Extracts of the bacteria were prepared by sonication, dialysis and filtration to provide a clear sterile solution which was characterised in terms of dissolved organic and condensed P (DOP and DCP) and molecular weight range. The extract was used as a source of P to Agrostis capillaris L. seedlings in nutrient solution from which orthophosphate was omitted. In a time course experiment, root surface phosphatase activity increased as soon as extract was added to the root medium, DOP was rapidly hydrolysed and orthophosphate concentration increased rapidly. These processes were complete within about 8 h, after which phosphatase activity fell to its original level, and the plants absorbed molybdate reactive P from the nutrient solution so that it reached its original concentration over 48 h. DCP concentrations did not change significantly throughout the experiment. This work clearly demonstrated that DOP but not DCP, as a component of a bacterial extract produced by a relatively straightforward method, was quickly hydrolysed and the P made available for plant uptake.     

Characterization of orthophosphate release from dissolved organic phosphorus by gel filtration and several hydrolytic enzymes

The molecular weight distribution of dissolved organic phosphorus (DOP) and the possible mechanisms of orthophosphate (Pi) release were examined by gel filtration and incubation with some hydrolytic enzymes. Sixty five percent of the DOP appeared to have apparent molecular weights between 300 to 10000 daltons. Less than 10% of the DOP estimated higher molecules greater than 10000 daltons. Alkaline phosphatase released Pi more easily from low molecular weight (< 1500 daltons) DOP than from high molecular weight fractions. While, addition of nucleases or phosphodiesterase alone did not appear Pi release from high molecular weight DOP compounds. Pi release from those DOP compounds increased markedly (more than 30%) when alkaline phosphatase was incubated with nucleases or phosphodiesterase. However, 60% of DOP did not release Pi when alkaline phosphatase was incubated with either enzymes.     

The winter and summer partitioning of dissolved nitrogen and phosphorus. Observations across the Irish Sea during 1997 and 1998

Winter and summer water column profiles from a transect co-inciding with the 54° N parallel across the Irish Sea were examined for relationships between the fractions of total dissolved nitrogen and phosphorus. Inorganic nitrogen and phosphorus gave the expected seasonal variation. As inorganic nitrogen concentrations decreased during the summer, dissolved organic nitrogen concentrations increased. Dissolved organic phosphorus showed little seasonal variation and formed a smaller fraction of the total dissolved pool compared to nitrogen. Winter inorganic nitrogen and phosphorus salinity relationships were conservative: winter organic nitrogen and phosphorus salinity relationships were not conservative. In the winter, dissolved organic nitrogen and phosphorus formed a significant fraction of the total dissolved pool.     

Iron:phosphorus ratio in surface sediment as an indicator of phosphate release from aerobic sediments in shallow lakes

Analysis of Danish lakes showed that both mean winter and mean summer concentrations of lake water total phosphorus in the trophogenic zone correlated negatively with the total iron to total phosphorus ratio (Fe:P) in surface sediments. No correlation was found between the water total phosphorus concentration and either the sediment phosphorus concentration alone or with sediment calcium concentration. The increase in total phosphorus from winter to summer, which is partly a function of net internal P-loading, was lowest in lakes with high Fe:P ratios in the surface sediment.A study of aerobic sediments from fifteen lakes, selected as representative of Danish lakes with respect to the sediment Fe and phosphorus content, showed that the release of soluble reactive phosphorus was negatively correlated with the surface sediment Fe:P ratio. Analysis of phosphate adsorption properties of surface sediment from 12 lakes revealed that the capability of aerobic sediments to buffer phosphate concentration correlated with the Fe:P ratio while the maximum adsorption capacity correlated with total iron. Thus, the Fe:P ratio may provide a measure of free sorption sites for orthophosphate ions on iron hydroxyoxide surfaces.The results indicate that provided the Fe:P ratio is above 15 (by weight) it may be possible to control internal P-loading by keeping the surface sediment oxidized. Since the Fe:P ratio is easy to measure, it may be a useful tool in the management of shallow lakes.     

The distribution of dissolved DNA in an oligotrophic and a eutrophic river of Southwest Florida

The distribution of dissolved DNA concentrations and some microbial variables were compared in an oligo-mesotrophic river (the Crystal River) and a phosphate-rich eutrophic river (the Alafia River) in Southwest Florida over a 15 month period. Concentrations of phosphate and nitrate in the Alafia River averaged 135 and 18.2 times the respective phosphate and nitrate concentrations of the oligo-mesotrophic Crystal River. The seasonal average dissolved DNA concentration for the Alafia River exceeded that of the Crystal River by a factor of 1.8 (8.2 g 1^–1 compared to 4.6 g 1^–1, respectively). The greatest concentrations of dissolved DNA in the Alafia River were found in areas that contained the largest populations of phytoplankton and bacteria (a reservoir formed from an abandoned phosphate mining pit and two downstream stations near the mouth of the river). Differences in dissolved DNA concentrations between these environments and more pristine environments (i.e. all Crystal River Stations and upstream Alafia River stations) were of the same order of magnitude (1.8 to 2.2-fold) as the differences in bacterial abundance and activity, but considerably less than differences in phytoplankton abundance and activity between such environments. Seasonal variations in dissolved DNA concentrations in the Crystal River corresponded to seasonal variations in microbial populations, with minimal values in January and greater values in July. In the Alafia River, lowest concentrations for dissolved DNA occurred in July during the wet season, when seasonal flooding of area of leaf litter yielded high levels of dissolved organic carbon (DOC) which were low in dissolved DNA. These results suggest that: 1) in situ planktonic activity is a greater source of dissolved DNA than allochthonous or terrestrial sources of DOC; 2) factors that control the magnitude of heterotrophic bacterial populations are more likely to control dissolved DNA levels than factors regulating autotrophic population activity and abundance; 3) differences in dissolved DNA between eutrophic and oligo-mesotrophic environments are often much smaller than the differences in nutrient concentration between such environments.     

A comparison of phosphorus and DOC leachates from different types of leaf litter in an urban environment

1. The leaching rates of filterable reactive phosphorus (FRP) and dissolved organic carbon (DOC) from five leaf litter types commonly occurring in urban environments in Mediterranean regions of Southern Australia were compared. The relative composition, bioavailability and oxygen demand of this DOC were also assessed. Four tree species were assessed, including the native river red gum (Eucalyptus camaldulensis) and three introduced deciduous species, the English elm (Ulmus procera), London plane (Platanus acerifolia) and white poplar (Populus alba). Grass cuttings (mixed species) were selected as a common garden waste. 2. Except for English elm, the majority of FRP and DOC was released within the first 48 h. Grass cuttings released the highest amount of FRP with white poplar releasing the most DOC. Species that released relatively high amounts of DOC (white poplar, English elm, river red gum) released relatively low amounts of FRP. Conversely, species that released relatively low amounts of DOC (grass cuttings and London plane) tended to release relatively high amounts of FRP. 3. Analysis of DOC composition, combined with the differing oxygen demand and DOC depletion curves, demonstrated that there were substantial differences in the DOC leached from the leaf litter of the different species. Biochemical oxygen demand and the biodegradability of the DOC was positively correlated with the proportion of hydrophilic and hydrophobic acids present in the leachate. 4. These results demonstrate that simple measurements of nutrient release per gram of leaf litter would be insufficient to predict the ecological impact on receiving waters resulting from changes in dominant vegetation. Furthermore, the use of traps to prevent particulate leaf material from entering streams may have limited potential for reducing the load of dissolved nutrients. We conclude that any changes to vegetation type which substantially alter the timing of leaf fall or the composition of leaf leachates should be avoided.     

Chromophoric dissolved organic matter (CDOM) absorption characteristics in relation to fluorescence in Lake Taihu,China, a large shallow subtropical lake

Absorption measurements from chromophoric dissolved organic matter (CDOM) and their relationships with dissolved organic carbon (DOC) and fluorescence were studied in Lake Taihu, a large, shallow, subtropical lake in China. Absorption spectra of lake water samples were measured from 240 nm to 800 nm. Highest values of a(λ), DOC and F _n (355) occurred near the river inflow to Meiliang Bay and decreased towards the central lake basin. A significant spatial difference was found between Meiliang Bay and the central lake basin in absorption coefficient, DOC-specific absorption coefficient, exponential slope coefficient, DOC concentration and fluorescence value. The spatial distribution of CDOM suggested that a major part of CDOM in the lake was from river input. CDOM absorption coefficients were correlated with DOC over the wavelength range 280–500 nm, and a(355) was also correlated with F _n (355), which showed that CDOM absorption could be inferred from DOC and fluorescence measurement. The coefficient of variation between a(λ) and DOC concentration decreased with increase in wavelength from 240 nm to 800 nm. Furthermore, a significant negative linear relationship was recorded between S value and CDOM absorption coefficient, as well as DOC-specific absorption coefficient. S value and DOC-specific absorption coefficient were used as a proxy for CDOM composition and source. Accurate CDOM absorption measurements are very useful in explaining UV attenuation and in developing, validating remote sensing model of water quality in Lake Taihu.     

Release and bioavailability of dissolved organic matter from floodplain litter: influence of origin and oxygen levels

1. We determined the rate of release and microbial uptake of dissolved organic carbon (DOC) leached from three components (leaves, bark and twigs) of river red gum ( Eucalyptus camaldulensis ) forest litter originating from different parts of a floodplain and under different oxygen levels.
2. Preliminary experiments showed that substantially more DOC was released from leaves than from bark or twigs; there was relatively little DOC release from coarse particulate matter or soil.
3. Both the amount of DOC released from each litter component and the amount metabolized by the microbial community were independent of position on the flood-plain or amount of oxygen available to microbes.
4. Although the bioavailability of DOC was independent of oxygen concentration, the microbial utilization of DOC under aerobic and anaerobic conditions differed. Under aerobic conditions, leaves were colonized by fungi, while bacteria were dominant under anoxic conditions.
5. Phospholipid fatty acid profiles of the microbial communities growing on leaf extracts showed that different microbial communities developed in each oxygen concentration treatment suggesting that, irrespective of flood conditions, a microbial community will develop to utilize a significant proportion of the DOC leached from litter.     

Release rates and potential fates of nitrogen and phosphorus from sediments in a eutrophic reservoir

1. Nutrients released from lake sediments can influence water column nutrient concentrations and planktonic productivity. We examined sediment nutrient release [soluble reactive phosphorus (SRP) and ammonia (NH)] at two sites in a eutrophic reservoir (Acton Lake, OH, U.S.A.) that differed in physical mixing conditions (a thermally stratified and an unstratified site). 2. Sediment nutrient release rates were estimated with three methods: sediment core incubations, seasonal in situ hypolimnetic accumulation and a published regression model that predicted sediment phosphorous (P) release rate from sediment P concentration. All three methods were applied to the deeper stratified site in the reservoir; however, we used only sediment core incubations to estimate SRP and NH release rates at the shallow unstratified site because of the lack of thermal stratification. We also compared the total P concentration (TP_S) of sediments and the concentration of P in various sediment fractions at both sites. 3. Anoxic sediments at the stratified site released SRP at rates more than an order of magnitude greater than oxic sediments at the shallow unstratified site. However, P accumulated in the hypolimnion at much lower rates than predicted by sediment core incubations. In contrast, NH was released at similar rates at both sites and accumulated in the hypolimnion at close to the expected rate, indicating that P was ‘lost’ from the hypolimnion through biogeochemical pathways for P, such as precipitation with inorganic material or biological uptake and sedimentation. 4. TP_S was significantly greater at the deeper stratified site and organically bound P accounted for >50% of TP_S at both sites. 5. We examined the magnitude of SRP fluxes into the study reservoir in 1996 by comparing the mean summer daily SRP fluxes from anaerobic sediments, aerobic sediments, stream inflows and gizzard shad excretion. While the SRP release from anaerobic sediments was high, we hypothesise that little of this SRP gained access to the epilimnion in mid‐summer. SRP flux to the reservoir from aerobic sediments was less than from gizzard shad excretion and streams. Large interannual variability in thermocline stability, gizzard shad biomass and stream discharge volumes, will affect SRP loading rates from different sources in different years. Therefore, construction of P budgets for different years should account for interannual variation in these parameters.     

Response of a eutrophic, shallow subtropical lake to reduced nutrient loading

1. Lake Apopka (FL, U.S.A.) was subjected to decades of high nutrient loading from farms developed in the 1940s on converted riparian wetlands. Consequences included perennially high densities of cyanobacteria, low water transparency, elimination of submerged vegetation, modified fish community, and deposition of nutrient‐rich, flocculent sediments. 2. Initial steps were taken to reduce phosphorus (P) loading. Through strengthened regulation and purchase of farms for restoration, external P loading was reduced on average from 0.56 to 0.25 g P m^?2 year^?1 (55%) starting in 1993. The P loading target for the lake is 0.13 g P m^?2 year^?1. 3. For the first 6 years of P loading reduction the annual sedimentation coefficient (σ) averaged 13% less than the prior long‐term value (0.97 versus 1.11 year^?1). The sedimentation coefficient, σ, was lower in the last 3 years of the study, but this period included extreme low‐water conditions and may not be representative. Annual σ was negative (net P flux to the water column) only 1 year. 4. Wind velocity explained 43% of the variation in σ during the period before reductions in total phosphorus (TP) concentration of lake water, but this proportion dropped to 6% after TP reductions. 5. Annual mean TP concentrations differed considerably from values predicted from external loading and hydraulic retention time using the Vollenweider–Organization for Economic Co‐operation and Development relationship. Reductions in lake water TP concentration fit model predictions better when multiyear (3‐year) mean values were used. 6. Evidence available to date indicates that this shallow, eutrophic lake responded to the decrease in external P loading. Neither recycling of sediment P nor wind‐driven resuspension of sediments prevented improvements in water quality. Reductions in TP concentration were evident about two TP‐resident times (2 × 0.9 year) after programmes began to reduce P loading. Improvements in concentrations of chlorophyll a and total suspended solids as well as in Secchi transparency lagged changes in lake‐water TP concentration but reached similar magnitudes during the study.     

Relationship of trophic and chemical conditions to photobleaching of dissolved organic matter in lake ecosystems

Dissolved organic matter (DOM) is a major light-absorbing substance, responsible for much of the color in water bodies. When sunlight energy is absorbed by DOM, some color can be lost by the process of photobleaching. We measured rates of DOM photobleaching in thirty lakes that varied greatly in color, trophic status and ionic composition. Loss of color (measured as absorbance at 440 nm and expressed as absorption coefficients) was a first order function of sunlight dose, and rates were nearly identical for 0.2m- and GF/F-filtered samples suggesting that the process was predominantly abiotic. Photobleaching rates were rapid (color loss of 1–19% d^–1) and varied about seven-fold among lakes. Our method under-estimated the actual rate by 15–20% based on comparisons between the glass bottles we used in the survey and quartz containers. The large variation in photobleaching rates was examined in relation to lake trophy and chemical conditions. The best predictor of this variability was acid-neutralizing capacity (ANC) (r ²=0.94;p<0.001) such that photobleaching was most rapid in the most alkaline lakes. The relationship between ANC and photobleaching suggests that differences in ionic conditions among lakes may influence the solubility and configuration of humic and fulvic acids and hence their susceptibility to photobleaching.     

Alternative explanations for rising dissolved organic carbon export from organic soils

Since 1988, there has been, on average, a 91% increase in dissolved organic carbon (DOC) concentrations of UK lakes and streams in the Acid Waters Monitoring Network (AWMN). Similar DOC increases have been observed in surface waters across much of Europe and North America. Much of the debate about the causes of rising DOC has, as in other studies relating to the carbon cycle, focused on factors related to climate change. Data from our peat‐core experiments support an influence of climate on DOC, notably an increase in production with temperature under aerobic, and to a lesser extent anaerobic, conditions. However, we argue that climatic factors may not be the dominant drivers of DOC change. DOC solubility is suppressed by high soil water acidity and ionic strength, both of which have decreased as a result of declining sulphur deposition since the 1980s, augmented during the 1990s in the United Kingdom by a cyclical decline in sea‐salt deposition. Our observational and experimental data demonstrate a clear, inverse and quantitatively important link between DOC and sulphate concentrations in soil solution. Statistical analysis of 11 AWMN lakes suggests that rising temperature, declining sulphur deposition and changing sea‐salt loading can account for the majority of the observed DOC trend. This combination of evidence points to the changing chemical composition of atmospheric deposition, particularly the substantial reduction in anthropogenic sulphur emissions during the last 20 years, as a key cause of rising DOC. The implications of rising DOC export for the carbon cycle will be very different if linked primarily to decreasing acid deposition, rather than to changes in climate, suggesting that these systems may be recovering rather than destabilising.     

Aluminum sulfate treatment: Short-term effect on complex phosphorus compounds in a eutrophic lake

Application of aluminum sulfate to the hypolimnion of West Twin Lake, a hardwater eutrophic lake in northeastern Ohio, did not efficiently remove complex phosphorus compounds from the water column. The majority of the complex phosphorus compounds remaining after alum treatment released P0₄ following brief treatment with alkaline phosphatase.     

Effects of different molecular weight fractions of dissolved organic matter on the growth of bacteria,algae and protozoa from a highly humic lake

Effects of different molecular size fractions (< 1000 MW, < 10 000 MW, < 100 000 MW and <0.1 μm) of dissolved organic matter (DOM) on the growth of bacteria, algae and protozoa from a highly humic lake were investigated. DOM from catchment drainage water as well as from the lake consisted mostly (59–63%) of high molecular weight (HMW) compounds (> 10 000 MW). With excess inorganic nutrients, the growth rate and yield of bacteria were almost identical in all size fractions. However, in < 1000 MW fractions and with glucose added, a longer lag phase occurred. Without added nutrients both the growth rates and biomasses of bacteria decreased towards the smaller size fractions and the percentage of dissolved organic carbon (DOC) used during the experiment and the growth efficiency of bacteria were lower than with excess nutrients. The growth efficiency of bacteria was estimated to vary between 3–66% in different MW fractions, largely depending on the nutrient concentrations, but the highest growth efficiencies were observed in HMW fractions and with glucose. The growth of algae was clearly lowest in the < 1000 MW fraction. In dim light no net growth of algae could be found. In contrast, added nutrients substantially enhanced algal growth and in deionized water with glucose, algae achieved almost the same growth rate and biomass as in higher MW fractions of DOM. The results suggested that bacteria and some algae were favoured by DOM, but protozoans seemed to benefit only indirectly, through bacterial grazing. The utilization of DOM by bacteria and algae was strongly affected by the availability of phosphorus and nitrogen.     

Problems with the application of diatom-total phosphorus transfer functions: examples from a shallow English lake

1. A diatom‐total phosphorus transfer function has been applied to a sedimentary diatom sequence from Groby Pool, a small shallow lake in Leicestershire, U.K. 2. Extensive aquatic plant records exist for Groby Pool dating back over two centuries. These records, in conjunction with selected aquatic pollen and herbarium diatom data, provide independent, qualitative evidence for the progression of eutrophication and its effects on aquatic plant communities and habitat structure. 3. Before 1800, Groby Pool was probably mesotrophic with clear water and a diverse submerged macrophyte community, but subsequently it experienced considerable nutrient enrichment. Key evidence for this includes: (i) historical plant records indicating the loss of species associated with mesotrophic waters and their replacement by others typical of eutrophic conditions, (ii) a significant increase in the percentage of planktonic diatoms in the lake sediment record (particularly Cyclostephanos dubius) after 1890, and (iii) increases in percentages of Stephanodiscus parvus and Cocconeis placentula in the second half of the twentieth century. 4. Diatom‐inferred total phosphorus (DI‐TP) estimates were inconsistent with the qualitative evidence for eutrophication at Groby Pool. In particular the DI‐TP profile was thought to overestimate phosphorus during the period of dominance by small Fragilaria spp. before 1890, and to misjudge the timing and direction of subsequent changes in nutrient loading. 5. This study highlights some of the problems associated with the application of diatom‐TP transfer functions to sedimentary diatom sequences from shallow lakes. The major problem relates to the frequent dominance of non‐planktonic diatoms in the sediments of these systems, many species of which (particularly small Fragilaria spp.) appear to be more sensitive to changes in habitat availability than to phosphorus. Potential ways of improving diatom‐TP models via altered approaches to sampling are suggested.     

The distributions and effects of nutrients in the sediments of a shallow eutrophic Chinese lake

The temporal and spatial distribution of total nitrogen and total phosphate in the sediments of a shallow eutrophic Chinese lake (Lake Chao), and their relationships with the physical and chemical features of sediments, and their effects on the lake water quality and trophic state, are presented in this paper. The following results were obtained: (1) higher concentration of Tot-N and Tot-P in the sediments occurred in the summer and the autumn seasons; (2) higher annual average Tot-N and Tot-P concentrations were observed in the sediments near the various river mouths and in the western part of the lake; (3) Tot-N and Tot-P concentration in the lake sediments generally increased with increase in lake sediment Eh, pH, and Al₂O₃, and declined with decrease in lake sediment size diameters and SiO₂; and, (4) correlations were observed in both the temporal and spatial distributions between the trophic state, Tot-N and Tot-P concentration in the lake water, and the Tot-N and Tot-P concentration found in the lake sediments.     

Coupling between benthic biomass of Microcystis and phosphorus release from the sediments of a highly eutrophic lake

Variations in microbial biomass and activity in the sediments of hypereutrophic Lake Vallentunasjön were followed during a period of 5 years. The data were compared to the calculated release of phosphorus from the sediments during the same period. A strong co-variation was found between biomass of Microcystis, heterotrophic bacterial activity in the sediments and internal phosphorus loading. These parameters exhibited mainly a declining trend during the investigation period. A pronounced stability of the sediment chemistry, including the fractional composition of the sediment phosphorus, during the studied period indicates that microbial activity affected the phosphorus release from the sediments. Calculations of the percentage of sediment bacteria that was associated to the mucilage of Microcystis colonies imply, together with the specific bacterial production, that Microcystis in the sediment stimulates bacterial production. In the highly phosphorus-saturated sediments of Lake Vallentunasjön this would ultimately lead to an increased release of phosphorus from the sediment. Lake Vallentunasjön does not follow the common pattern of recovery after reduction of external phosphorus loading. The large biomasses and long survival of Microcystis in the sediment are probably important reasons for the delayed recovery of the lake.