5'-nucleotidase activity in a eutrophic lake and an oligotrophic lake

Differences in enzymatic hydrolysis of dissolved organic phosphorus and subsequent phosphorus uptake were compared by using dual-labeled (gamma-P and 2-H) ATP in oligotrophic Lake Michigan and a moderately eutrophic lake in southeastern Michigan. More than 50% of the phosphate that was hydrolyzed was immediately taken up into bacterium-sized particles in the eutrophic lake and at a near-shore site in Lake Michigan. Less than 50% of the hydrolyzed phosphate was taken up into bacterium-sized particles at an offshore site in Lake Michigan. It is hypothesized that differences in size-fractionated uptake were the result of greater phosphorus utilization capacity in bacteria in habitats where loading of organic carbon is greater. Substantial isotope dilution of labeled phosphate uptake by unlabeled phosphate occurred, which implied that the phosphate was hydrolyzed extracellularly in both systems. Comparable nucleotidase activities were measured in the eutrophic lake and Lake Michigan, but the significance of the phosphate regenerated relative to particulate phosphorus pools was an order of magnitude greater in Lake Michigan. Seventy percent of the nucleotidase activity was inhibited by 100 muM phosphate in the eutrophic lake, which suggests that most hydrolysis was by phosphatase. Therefore, nucleotidase activity may be more important to phosphorus regeneration in oligotrophic habitats than phosphatase activity.     

Are the banks a source of recolonization after disturbance: an experiment on aquatic vegetation in a former channel of the Rhône River

Hydrolysis of natural dissolved organic phosphorus (DOP) in three hardwater lakes of different trophic level was calculated from kinetic data of phosphatase activity (PA) in different size fractions. DOP as well as kinetics of PA were determined every fortnight in depth profiles during the year 1990. 60% of DOP was assumed to be suitable substrate for phosphatases. The rate of hydrolysis increased markedly with higher trophic level. Average hydrolysis rate of DOP in polytrophic lake Thaler See was 3.26 nM P min^–1 (6 µg P-PO₄ l^–1 h^–1). In oligotrophic Lake Herrensee, dissolved phosphatases were responsible for more than half of the total hydrolysis. In the other two lakes, bacterial and algal surface PA dominated hydrolysis in changing parts depending on kinetics and DOP concentration. The regeneration rate of phosphate by PA was compared to phosphorus (P) excretion rate of zooplankton. Excretion was calculated from zooplankton data and excretion equations from the literature. In oligotrophic Lake Herrensee, excretion by zooplankton recycled in average 18% of the phosphate amount which was hydrolysed from DOP by PA. With higher trophic level, relevance of P excretion from zooplankton decreased drastically.     

Biogeochemical silica mass balances in Lake Michigan and Lake Superior

Silica budgets for Lake Michigan and Lake Superior differ in several respects. Mass balance calculations for both lakes agree with previous studies in that permanent burial of biogenic silica in sediments may be only about 5% of the biogenic silica produced by diatoms. Because dissolution rates are large, good estimates of permanent burial of diatoms can not be obtained indirectly from the internal cycle of silica (silica uptake by diatoms and subsequent dissolution) but must be obtained from the sediment stratigraphy. The annual net production of biogenic silica in Lake Michigan requires 71% of the winter maximum silica reservoir which must be maintained primarily by internal cycling in this large lake whereas the comparable silica demand in Lake Superior is only 8.3%. The greater silica demand in Lake Michigan is the result of phosphorus enrichment which has increased diatom production. It is hypothesized that steady-state silica dynamics in Lake Michigan were disrupted by increased diatom production between 1955 and 1970 and that a new steady state based on silica-limited diatom production developed after 1970. Mass balance calculations for Lake Michigan show in contrast with previous work that the hypothesized water column silica depletion of 3.0 g · m^–3 could have occurred even though 90% or more of the biogenic silica production is recycled.     

Eutrophication of Lake Neuchâtel indicated by the oligochaete communities

Lake Neuchâtel (Switzerland), oligotrophic until 1950, was meso-eutrophic in 1980. The relative abundance in worm communities of Peloscolex velutinus and Stylodrilus heringianus was used to monitor the trophic state of the lake. In 1980, the median relative abundance of these oligotrophic species was 9% in the whole of Lake Neuchâtel compared with 70% in oligotrophic lakes, 35% in mesotrophic lakes, and 0% in eutrophic lakes. The scarcity of oligotrophic species in the deepest area (153 m) characterized better the meso-eutrophic state of Lake Neuchâtel than oxygen concentrations which never descended below 6 mg·1^-1. Location of the area within the lake from where worms were sampled was of critical importance to assess the trophic state: some areas reflected the past rather than the present state of the lake.     

Comparing Alkaline Phosphatase PhoX-Encoding Genes in Two Contrasting Habitats of the Large Eutrophic Lake Taihu,China

River mouth areas of eutrophic lakes have frequent cyanobacterial blooms and high organophosphate mineralization rates. To determine the genetic background of the high organophosphate mineralization rate in river mouth areas of eutrophic lakes, we compared the diversity and abundance of bacterial alkaline phosphatase PhoX-encoding genes in the river mouth areas and the lake center within the large eutrophic Lake Taihu, China. Results showed that the seasonal average values of phosphorus content, bacterial alkaline phosphatase activity, diversity, and abundance of bacterial phoX in the river mouths were significantly higher than those in the lake center, and 47% of all identified genotypes were present only in the river mouths. The genotypes most similar to alphaproteobacterial phoX dominated in both regions, with 72% and 64% of the total genotypes in the lake center and river mouths, respectively. More than 40% of the total phoX genotypes were only found in the river mouths, however, indicating the special microbial nature in this hypereutrophic habitat. The differences in contents of dissolved organic carbon and phosphorus were related to the divergences of bacterial phoX genotypes in these two contrasting habitats. Because some dominant genotypes most similar to phoX genes of alphaproteobacterial genus were observed only in the river mouth areas, such as Microvirga sp. WSM3557, and because the hydraulic retention time of Lake Taihu is insufficient to shape the special bacterial groups in the river mouths, allochthonous inputs of nutrients and bacterial species could make contributions to explain the different organophosphate mineralization rates in these two contrasting habitats of the large eutrophic lake.     

Effects of Nutrients on Specific Growth Rate of Bacterioplankton in Oligotrophic Lake Water Cultures

The effects of organic and inorganic nutrient additions on the specific growth rates of bacterioplankton in oligotrophic lake water cultures were investigated. Lake water was first passed through 0.8-μm-pore-size filters (prescreening) to remove bacterivores and to minimize confounding effects of algae. Specific growth rates were calculated from changes in both bacterial cell numbers and biovolumes over 36 h. Gross specific growth rates in unmanipulated control samples were estimated through separate measurements of grazing losses by use of penicillin. The addition of mixed organic substrates alone to prescreened water did not significantly increase bacterioplankton specific growth rates. The addition of inorganic phosphorus alone significantly increased one or both specific growth rates in three of four experiments, and one experiment showed a secondary stimulation by organic substrates. The stimulatory effects of phosphorus addition were greatest concurrently with the highest alkaline phosphatase activity in the lake water. Because bacteria have been shown to dominate inorganic phosphorus uptake in other P-deficient systems, the demonstration that phosphorus, rather than organic carbon, can limit bacterioplankton growth suggests direct competition between phytoplankton and bacterioplankton for inorganic phosphorus.     

Isolation and Characterization of Organic Phosphorus-Mineralizing Bacteria in Sediment of a Chinese Large Shallow Eutrophic Lake (Lake Taihu)

The organic phosphorus mineralizing bacteria (OPB) play an important role in phosphorus cycling in lake sediment, to which less attention has been paid. Monthly sediment samples in 2009 ending in October, together with the samples from different seasons (May, July, October, and December) in 2008, were collected at 6 sites in a Chinese large shallow eutrophic lake (Lake Taihu). The sediment OPB numbers ranged from 2.2 × 10⁶ to 1.79 × 10⁸ cells g^?1 (dry weight) at different sampling sites and in different seasons, with the average being 3.88 × 10⁷ cells g^?1 (dry weight). Its number was highest at the most polluted site and peaked in spring and summer, which can be explained by the enrichment of organic matter in sediment. Furthermore, there existed a significant positive relationship between the OPB numbers and alkaline phosphatase activities in the sediment. The 6 OPB strains isolated from the sediment were distinct in terms of their colony morphology on the yolk agar, biochemical characteristics and phosphorus release abilities. According to the 16S rDNA sequences, these OPB belong to Bacillus cereus, Stenotrophomonas maltophilia, Stenotrophomonas sp., Bacillus cereus, Xanthomonas sp., Pseudomonas sp. They were distinguished from the OPB species recorded in a Chinese small shallow eutrophic lake whose sediment organic content was remarkably higher. Taken together, organic matter shaped the OPB community not only quantitatively but also qualitatively, which in concert facilitated the enzymatic hydrolysis of organic phosphorus in lake sediment.     

Methanogenesis from Methanol and Methylamines and Acetogenesis from Hydrogen and Carbon Dioxide in the Sediments of a Eutrophic Lake

¹⁴C-tracer techniques were used to examine the metabolism of methanol and methylamines and acetogenesis from hydrogen and carbon dioxide in sediments from the profundal and littoral zones of eutrophic Wintergreen Lake, Michigan. Methanogens were primarily responsible for the metabolism of methanol, monomethylamine, and trimethylamine and maintained the pool size of these substrates below 10 μM in both sediment types. Methanol and methylamines were the precursors for less than 5 and 1%, respectively, of the total methane produced. Methanol and methylamines continued to be metabolized to methane when the sulfate concentration in the sediment was increased to 20 mM. Less than 2% of the total acetate production was derived from carbon dioxide reduction. Hydrogen consumption by hydrogen-oxidizing acetogens was 5% or less of the total hydrogen uptake by acetogens and methanogens. These results, in conjunction with previous studies, emphasize that acetate and hydrogen are the major methane precursors and that methanogens are the predominant hydrogen consumers in the sediments of this eutrophic lake.     

Phosphatase activity in relation to phytoplankton composition and pH in Swedish lakes

SUMMARY. 1. Potential phosphatase activity and phytoplankton from several lakes of different character were compared in order to evaluate the importance of lake water pH and phytoplankton composition for the activity and pH optimum of lake water phosphatases.
2. In oligotrophic lakes, in which phytoplankton biomass was most often dominated by Ochromonadaceae spp., optimum phosphate activity was found at pH values <6. In eutrophic lakes, where species of Cyanophyceae and Bacillariophyceae dominated the phytoplankton biomass, optimum phosphatase activity was found at pH 7.5 or 8.5.
3. The pH optimum of phosphatase activity often differed from the corresponding lake water pH.
4. Experimental variation in phosphorus availability resulted in predictable changes in phosphatase activity. However, specific phosphatase activity, calculated per biomass of phytoplankton, was dependent on plankton species composition.     

Spatial and temporal variability of internal and external phosphorus loads in Mona Lake,Michigan

A study was conducted in Mona Lake, a small eutrophic lake located in western Michigan (USA) to address the temporal and spatial variability of external and internal phosphorus loading. External P load varied among subbasins, which was mostly related to discharge, but also to land use. Black Creek, which drains lands with natural cover and agriculture, accounted for the majority of flow, and total phosphorus (TP) and soluble reactive phosphorus (SRP) load, to Mona Lake. However, the relative contribution of SRP load was greater in Little Black Creek, which flows through a mostly urbanized subbasin, than in Black Creek. The relative importance of internal loading was strongly related to season, as internal TP loads contributed only ∼9% of the overall P load in April 2005, but ∼68–82% of the overall P load in the summer and early fall seasons. Internal TP and SRP loading was greater under anaerobic than aerobic conditions. Mean anaerobic TP release rates ranged from 0.80 to 15.56 mg P m⁻² d⁻¹, varying with site and season. Spatial variability in both internal phosphorus loading and sediment P concentration was also evident. By taking into account the spatial and temporal variability of different loading sources, management practices can be targeted to optimize nutrient source control strategies.     

Phosphorus fractions and alkaline phosphatase activity in sediments of a large eutrophic Chinese lake (Lake Taihu)

Spatial, vertical, and seasonal variations in phosphorus fractions and in alkaline phosphatase activity (APA) were investigated in sediments in a large-shallow eutrophic Chinese lake (Lake Taihu) in 2003–2004. The phosphorus content was highest in the most seriously polluted lake area. Iron-bound phosphorus (Fe(OOH)～P) dominated (47% on average) among the phosphorus fractions determined according to Golterman (Hydrobiologia 335:87–95, 1996). Notably, organically-bound P comprised a further significant additional portion (acid-soluble + hot NaOH-extractable organic P = 25%), which was highest at the most polluted sites. The Fe(OOH)～P content was the lowest in spring (April, 2004), suggesting that degradation of organic matter led to the release of iron-bound phosphates. Sediment APA showed a significant positive relationship with both organically-bound P and Fe(OOH)～P. Consequently, organically-bound P is an important portion of the sediment phosphorus in Lake Taihu. It is mainly derived from freshly-settled autochthonous particles and from external discharges. Organically-bound P induces APA and may lead to the release of bioavailable phosphates from the organic sediments, thereby accelerating lake eutrophication.     

Seasonal Variations in Chlorophyll a Concentrations in Relation to Potentials of Sediment Phosphate Release by Different Mechanisms in a Large Chinese Shallow Eutrophic Lake (Lake Taihu)

The relationship between chlorophyll a and fractionation of sediment phosphorus, inorganic phosphate-solubilizing bacteria (IPB), and organic phosphate-mineralizing bacteria (OPB) was evaluated in a large Chinese shallow eutrophic lake (Lake Taihu) and its embayment (Wuli Bay). At the three study sites, the increase of chlorophyll a concentrations in April paralleled those of the iron bound phosphate accounting for major portion of sediment inorganic phosphate, and in June significantly higher OPB and IPB numbers (especially OPB) in sediment were main contributors to the peaks of chlorophyll a concentration. Even though IPB peaked from February to June, it should serve as an unimportant P source due to the irrelevancy with chlorophyll a and soluble reactive phosphorus (SRP). By contrast, at the other site in the embayment, the calcium-bound phosphate was predominant and solid, which was difficult to be released, and neither IPB nor OPB were detectable in the sediment, indicating weak potential for phosphorus release from the sediment, which was reflected in the small seasonal variation in SRP concentration in water column. Hence, the extents to which the three general mechanisms behind phosphate release from sediment (desorption of iron bound phosphate, solubilization by IPB and enzymatic hydrolysis by OPB) operated were different depending on seasons and sites in Lake Taihu, they may jointly drive phosphate release and accelerate the eutrophication processes.     

Eutrophication of Lake Geneva indicated by the oligochaete communities of the profundal

Primary production rates and total phosphorus concentrations indicated that Lake Geneva (Switzerland) was meso-eutrophic from 1970 to 1983. Worm communities of the profundal (50–309 m deep) were very similar in 1978 and 1983. Species numerically dominant in eutrophic lakes — such as Potamothrix hammoniensis, P. heuscheri and Tubifex tubifex — constituted the bulk (75%) of the communities. Species numerically dominant in mesotrophic lakes (mostly Potamothrix vejdovskyi) or in oligotrophic lakes (mostly Stylodrilus heringianus) constituted respectively 18% and 7% of the worm communities. The dominance of eutrophic species increased with depth in the whole lake; it increased also in the eastern region of the lake which is directly exposed to the heavy organic inputs of the Rhône River. Oligotrophic and mesotrophic species decreased along the same gradients. Species dominant in oligotrophic lakes were absent in 1978 and 1983 from the deepest area of Lake Geneva (300–309 m) whereas they constituted therein 25% of worm communities in 1967. Data based on worm species groups — i.e. species with similar resistance to eutrophication pooled together — were more easy to analyse statistically than those based on the isolated species. Thus, the relative abundance of three species groups, expressed in several ways, can indicate precisely the trophic state of a lake.     

Variable rates of phosphate uptake by shallow marine carbonate sediments: Mechanisms and ecological significance

We determined phosphate uptake by calcareous sediments at two locations within a shallow lagoon in Bermuda that varied in trophic status, with one site being mesotrophic and the other being more eutrophic. Phosphate adsorption over a six hour period was significantly faster in sediments from the mesotrophic site. Uptake at both sites was significantly less than that reported for a similar experiment on calcareous sediments in an oligotrophic lagoon in the Bahamas. The difference in phosphorus adsorption between our sites did not appear to be related to sediment characteristics often cited as important, such as differences in surface area (as inferred from grain size distributions), total organic matter content, or iron content. However, the sediment total phosphorus contents were inversely related to phosphorus uptake at our sites in Bermuda, and at the previously studied Bahamas site.We hypothesize that phosphate uptake in these calcareous sediments is a multi-step process, as previously described for fluvial sediments or pure calcium carbonate solids, with rapid initial surface chemisorption followed by a slower incorporation into the carbonate solid-phase matrix. Accordingly, sediments already richer in solid phase phosphorus take up additional phosphate more slowly since the slower incorporation of surface-adsorbed phosphate into the carbonate matrix limits the rate of renewal of surface-reactive adsorption sites.Although carbonate sediments are a sink for phosphate, and thereby reduce the availability of phosphorus for benthic macrophytes and phytoplankton in the shallow overlying water, phosphate uptake by these sediments appears to decrease along a gradient from oligotrophic to eutrophic sites. If our result is general, it implies a positive feedback in phosphorus availability, with a proportionately greater percentage of phosphorus loading being biologically available longer as phosphorus loading increases. This pattern is supported by the significantly higher tissue phosphorus content of the seagrass,Thalassia testudinum, collected from the eutrophic inner bay site. Over time, this effect may tend to cause a shift from phosphorus to nitrogen limitation in some calcareous marine environments.     

Budding and prosthecate bacteria from freshwater habitats of various trophic states

Budding and prosthecate bacteria were enumerated in spring and summer by viable counting procedures in several freshwater habitats in Australia including oligotrophic lakes, a mesotrophic lake, and eutrophic ponds.Caulobacter spp. were the most numerous type encountered. They were present in the highest concentrations (exceeding 1000/ml) in the mesotrophic lake during the summer. Their proportion to total viable heterotrophic bacteria was also highest (35.1 to 37.7) in this habitat. From 17 to 330/mlCaulobacter spp. were counted in the eutrophic habitats where their proportion to total viable numbers was less than 1.0%. In the oligotrophic lakes they varied from 5 to 23/ml and comprised greater than 5% of the total viable count.Hyphomicrobium- like bacteria were also numerous in the mesotrophic lake and in one oligotrophic lake during the summer sampling period.Ancalomicrobium spp. occurred in high concentrations (130/ml) in the mesotrophic lake. Budding bacteria of thePlanctomyces-Pasteuria group were most numerous in the eutrophic habitats where as many as 240/ml were counted; their proportion to total heterotrophs remained relatively constant regardless of trophic state, however. A similar pattern was observed withProsthecobacter spp.     

Determining trophic state in Lake Whatcom,Washington (USA), a soft water lake exhibiting seasonal nitrogen limitation

We evaluated an eleven year data set to assess trophic state and nutrient limitation in Lake Whatcom, an oligotrophic, soft water, chain lake located in the Puget Sound lowlands of Washington (U.S.A.). Although total phosphorus (TP) and soluble reactive phosphate (SRP) concentrations were relatively low throughout the lake, there were significant differences between the northern basin (Site 1) and the other sampling sites (Sites 2–4). Nonparametric correlation coefficients (Kendall's ) were highest between chlorophyll (CHL), Secchi depth (SD), total nitrogen (TN), and dissolved inorganic nitrogen (DIN). Late summer algal biomass correlated best with DIN and TP. Trophic State Indices based on TP, TN, CHL and SD revealed that although algal growth was most likely phosphorus limited throughout the year, the northern basin of the lake may have developed nitrogen co-limitation during late summer and fall. During this period, N/P ratios were often less than 20, and in 1998 the epilimnetic DIN concentrations dropped below 20 g l^–1 while DIN/TP ratios fell below 4. Reviews of the literature suggest that while co-limitation by phosphorus and nitrogen is fairly common in unproductive lakes, the patterns seen in Lake Whatcom were more similar to those reported for eutrophic lakes experiencing secondary nitrogen limitation resulting from excess phosphorus loading.     

Reversal of eutrophication in Lake Geneva: evidence from the oligochaete communities

1. In Lake Geneva (Switzerland), total phosphorus concentrations have decreased from 90 mg m^?3 in 1979 to 55 mg m^?3 in 1990. 2. To assess the effects of this improvement, tubificid and lumbriculid communities were sampled in 1982 and in 1991 in the same areas, at a depth of 40 m. 3. Abundance of mesotrophic species (mostly Potamothrix vejdovskyi) and of eutrophic species (mostly Potamothrix hammoniensis) was lower in 1991 than in 1982; in contrast, oligotrophic species (mostly Peloscolex velutinus) were more abundant in 1991 than in 1982. 4. The changes recorded in 1991 were the same as those associated with a decrease of organic sedimentation. 5. Mean relative abundance of oligotrophic species increased from 17% in 1982 to 41% in 1991. According to these values, Lake Geneva was mesoeutrophic in 1982, but mesotrophic in 1991.     

Sediment diatom assemblages and composition of pore-water dissolved organic matter reflect recent eutrophication history of Lake Peipsi (Estonia/Russia)

A paleolimnological approach was used for the assessment of the recent eutrophication history and identification of possible reference conditions in the large, shallow, eutrophic Lake Peipsi. Lake Peipsi is the fourth largest lake by area, and the largest transboundary lake in Europe, being shared between Estonia and Russia. Lake Peipsi has been anthropogenically impacted over a longer time-scale than that covered by instrumental limnological monitoring. The ²¹⁰Pb record and down-core distribution of fly-ash particles in the 40-cm core from the middle part of the lake suggest 130 years of sediment accumulation. Diatom assemblages indicate alkaline mesotrophic conditions and a well-illuminated water column, sediment pore-water fluorescence index values suggest low autochthonous productivity and a stable aquatic ecosystem similar to natural reference conditions during the second half of 19th and early 20th century. Near-synchronous stratigraphic changes including the expansion of the eutrophic planktonic diatom Stephanodiscus parvus, the appearance of new species associated with eutrophic lakes and the decrease in the relative abundance of littoral diatoms, together with changes in the fluorescence properties of sediment pore-water dissolved organic matter, imply increased nutrient availability, enlarged phytoplankton crops, reduced water-column transparency and the onset of human-induced disturbances in the lake since the mid-20th century. The most conspicuous expansion of eutrophic planktonic diatoms and maximum concentration of siliceous microfossils occur simultaneously with changes in the fluorescence indexes of pore-water dissolved organic matter, indicating a pronounced increase in the contribution of autochthonous organic matter to the lake sediment. This implies that nutrient loading and anthropogenic impact was at a maximum during the 1970s and 1980s. Sedimentary diatom flora may reflect a reduction of phosphorus loading since the 1990s. However, the absolute abundance of planktonic diatoms and sediment pore-water fluorescence index values vary greatly implying that the lake ecosystem is still rather unstable.     

Zooplankton and environmental factors of a recovering eutrophic lake (Lysimachia Lake, Western Greece)

The present study investigates the zooplankton community dynamics and the abiotic environment in the eutrophic Lake Lysimachia (western Greece). The lake is considered to be recovering from eutrophication after the termination of an urban sewage inflow in 2000, and its waters are replenished constantly from the nearby oligotrophic Lake Trichonis. The results show that, although a decrease in nutrient concentrations was observed compared to the past, the lake still has eutrophic characteristics. This was reflected in the zooplankton community which is typical of those found in eutrophic lakes where rotifers prevail. Similarities among this lake and other nearby lakes were found considering the zooplankton community composition and seasonal variation. However, Lake Lysimachia is inhabited also by a number of different and even unique species (e.g., Moina micrura), suggesting that this ecosystem may be an important biodiversity refuge. Most of the zooplankton species were correlated with water temperature and, to a lesser extent, eutrophication key-water quality variables. Although there are few available data on the zooplankton of the lake, the abundance and composition of the community presenting characteristics indicative of intermediate trophic conditions and suggesting that the lake is probably under a kind of “biological” recovery.     

Phosphorus inputs to Lake Naivasha,Kenya, from its catchment and the trophic state of the lake

The main river supplying Lake Naivasha, Kenya, the Malewa, drains a catchment given over to largely subsistence cultivation and animal husbandry. The lake itself is the focus for an intensive horticultural industry based upon irrigation from the lake. The Malewa, however, is relatively independent of the impact of industry, and so its contribution to eutrophication of the lake was evaluated. Two periods of study, a very wet-dry and a `normal' wet-dry season showed that the river contribution of phosphorus led to a total phosphorus loading of 1.4 g m<sup>–2</sup> lake surface ann<sup>–1</sup> in the very wet period compared to 0.2 in the `normal'. Chlorophyll `a' in the open water of the lake was significantly related to soluble reactive phosphorus. The lake is now eutrophic by normal limnological criteria.