The characterization of phosphorus excretion products of a natural population of limnetic zooplankton

The characteristics of phosphorus excretions of zooplankton collected in spring 1974 from Stonehouse Pond, New Hampshire were studied with gel filtration, ultraviolet spectroscopy and seston incorporation. The soluble phosphorus component released from a natural population has a molecular weight the same as orthophosphate and behaves similar to orthophosphate in seston incorporation studies. Approximately 15 percent of the total phosphorus released was organic, but could not be identified by ultraviolet spectroscopy. No evidence of nucleic acid excretion or any hydrolytic degradation product was detected in the ultraviolet spectrum.The experimental work was based on a thesis submitted to the Graduate School of the University of New Hampshire in partial fulfillment of the requirements for the degree of Doctor of Philosophy.     

Diel and vertical variability of seston food quality and quantity in a small subalpine oligomesotrophic lake

Vertical profiles of seston food quality and quantity were measuredin subalpine Castle Lake for particulate carbon, chlorophylla, fatty acids and phosphorus in addition to abiotic parametersincluding water temperature and dissolved oxygen levels. Fieldand laboratory incubation experiments were employed to manipulateDaphnia rosea growth environments. Sestonic eicosapentaenoicacid (EPA) content was much lower, whereas carbon to phosphorus(C:P) ratios were much higher in the epilimnion compared tothe deep-water chlorophyll maximum (16–20 m). In a temperature-controlledincubation, Daphnia grew faster when fed seston from the deep-waterchlorophyll maximum. In in situ cage incubations, D. rosea grewfastest in the epilimnion. Daphnia rosea in a migrating treatmentexhibited intermediate growth rates between the ones for epilimnionand hypolimnion. A projection of D. rosea growth rates by growthmodels without regard to water temperature showed much highergrowth potentials in the hypolimnion. However, with inclusionof water temperature, epilimnetic water always gave higher potentialgrowth rates. In this lake, warmer temperatures of epilimneticwater override the higher food quality and quantity providedby the deep-water chlorophyll maximum.     

Seasonal changes in the biochemistry of lake seston

1. The quantity of seston was measured and the elemental carbon, nitrogen and phosphorus (C, N, P) and biochemical composition (carbohydrate, protein, lipid) of the < 53 μm size fraction in three temperate lakes during one year was analysed. The lakes differed in nutrient concentration and were characterized as oligotrophic, mesotrophic and eutrophic. Linear regression analyses defined associations between seston composition and either lake trophic status, depth or season. 2. The concentration of particulate organic seston was greatest during spring and autumn and lowest during the clear water period in early summer. Seasonal patterns in seston elemental and biochemical percentage composition (quality) were observed to be independent of differences in seston quantity. 3. Concentrations of seston C, N and P were high in most cases in the spring and autumn and low in summer. Concentrations of P were particularly high during late summer and early autumn in the metalimnion, perhaps because of recovery of P from anaerobic sediments and hypolimnetic waters. Because seston C and N did not increase as markedly as P, C : P and N : P ratios both declined in the autumn. Primary production was thought to be co-limited by N and P in all three of these lakes; however, the data suggested that N might be more important as a major limiting nutrient in the eutrophic lake as the metalimnion increased in depth in late summer and autumn. 4. Concentrations of protein, carbohydrate, polar lipid and triglyceride generally increased with lake type as expected (greatest in the eutrophic lake), but showed no relationship with water depth. As the year progressed, no significant changes were measured in protein and carbohydrate concentrations; however, the concentration of polar lipid decreased and triglyceride increased significantly with time of year. 5. The biochemical composition of seston varied during the year and among lakes; for example, in Lake Waynewood the proportion of protein composing the seston (percentage protein by weight) varied from < 10% to > 40%. No statistically significant patterns in the percentage protein or carbohydrate were found. However, the proportion of seston comprised of triglyceride decreased with lake type and increased during the year; whereas the proportion of seston as polar lipid increased with lake type and decreased during the year. Triglyceride comprised most of the lipid. Both protein : lipid and protein : carbohydrate ratios tended to be greatest in summer and lowest in the spring and autumn. 6. Relationships between samples and biochemical composition analysed by Canonical Correspondence Analysis (Canoco) indicated similar patterns in seasonal changes in seston biochemistry for the three lakes, with samples separated primarily by vectors for lake type (oligotrophic to eutrophic) and the percentage polar lipid (proportion of total lipid) and secondarily by vectors for date and water depth (epilimnion or metalimnion). 7. These seasonal biochemical changes in the seston food base were compared with biochemical changes known to occur in algae grown under N-or P-limited conditions in the laboratory, and the resultant quality of this algal food for suspension-feeding consumers (zooplankton). It was concluded that zooplankton were likely to be physiologically challenged by these distinct seasonal shifts in the quality of lake seston.     

Phosphorus Losses from the Epilimnion in Římov Reservoir

The amount of settling phosphorus was measured in Římov Reservoir using sediment trap technique from April 1986 to April 1987. Sediment traps were placed at three depths near the dam of the reservoir and at the bottom along the reservoir. The highest amount of phosphorus in trapped material was found during the fall turnover in the epilimnion and near the bottom in both spring periods (1986,1997). During the growing season the changes in dry weight and total phosphorus in settling seston were related to changes of phytoplankton biomass in the trophogenic layer. The amount of trapped phosphorus was higher near the bottom than in the upper layers of the reservoir throughout the year.     

Regeneration of inorganic phosphorus and nitrogen from decomposition of seston in a freshwater sediment

The mineralization of phosphorus and nitrogen from seston was studied in consolidated sediment from the shallow Lake Arreskov (July and November) and in suspensions without sediment (July). In the suspension experiment, phosphorus and nitrogen were mineralized in the same proportions as they occurred in the seston. During the 30 days suspension experiment, 47 and 43% of the particulate phosphorus and nitrogen, respectively, was mineralized with constant rates.Addition of seston to the sediment had an immediate enhancing effect on oxygen uptake, phosphate and ammonia release, whereas nitrate release decreased due to denitrification. The enhanced rates lasted for 2–5 weeks, while the decrease in nitrate release persisted throughout the experiment. The increase in oxygen uptake (equivalent to 21% of the seston carbon) was, however, only observed in the July experiment. The release of phosphorus and nitrogen from seston decomposing on the sediment surface differed from the suspension experiments. Thus, between 91 and 111% of the phosphorus in the seston was released during the experiments. Due to opposite directed effects on ammonium and nitrate release, the resulting net release of nitrogen was relatively low.A comparison of C/N/P ratios in seston, sediment and flux rates indicated that nitrogen was mineralized faster than phosphorus and carbon. Some of this nitrogen was lost through denitrification and therefore not measurable in the flux of inorganic nitrogen ions. This investigation also suggests that decomposition of newly settled organic matter in sediments have indirect effects on sediment-water exchanges (e.g. by changing of redox potentials and stimulation of denitrification) that modifies the release of mineralized phosphate and nitrogen from the sediment.     

Temporal and vertical dynamics of phytoplankton net growth in Castle Lake, California

The impact of nutrient additions, zooplankton grazing and light intensity on phytoplankton net growth with depth and season was studied with five microcosm experiments in meso-oligotrophic, subalpine Castle Lake, California, during the period of summer stratification in June-September 1994. The incubations (4 day) were performed at 5 m intervals from the surface to the bottom using natural phytoplankton and zooplankton assemblages, with enrichments of phosphorus and nitrogen. The phytoplankton community was only limited by nutrients in the upper 5 m (epilimnion), as indicated by change in chlorophyll concentration. Nutrient enrichments had the greatest effect on the phytoplankton net growth in June and July. High light inhibited the phytoplankton net growth at the surface. Low light intensities limited phytoplankton at 20 m and below, and at the end of the growing season already around 10-15 m. A deep chlorophyll maximum in the hypolimnion in June-August was not limited by either light or nutrients. The results showed variation in grazers' impact on phytoplankton. These results suggest the importance of nutrient limitation only in the epilimnion with light inhibition at the surface, light limitation in the hypolimnion, and varying impact of zooplankton grazing in influencing the development of the phytoplankton in Castle Lake.      

Hypolimnetic phosphorus retrieval by diel vertical migrations of lake phytoplankton

SUMMARY. I. Movement of ³³P from hypolimnion to epilimnion in a small, dystrophic lake was investigated using small-diameter experimental tubes enclosing thermally stratified water columns. This approach was made possible by the extremely sharp stratification found in such lakes, in which the euphotic zone closely coincides with the epilimnion.
2. The vertical distribution of inorganic phosphorus in the lake showed a sharp increase across the thermocline so that enhanced concentrations were available to phytoplankton just below the thermocline. Inorganic nitrogen concentrations did not show such a marked relation to thermal stratification.
3. One abundant motile alga ( Cryptomonas marssonii ) showed striking and regular vertical migrations in the lake, moving below the thermocline at night and returning to the surface waters in early morning. These migrations took cells across a 10°C temperature gradient. Non-motile phytoplankton showed constant vertical distributions.
4. In the experimental tubes an upward movement of phosphorus took place from hypolimnion to epilimnion which was only attributable to transport by phytoplankton cells undertaking active vertical migrations. No equivalent movement of phosphorus occurred in control tubes from which algae were absent.
5. The possible significance of such nutrient retrieval is discussed with reference to plankton phosphorus budgets and competition between phytoplankton species.     

Bacterioplankton in a small polyhumic lake with an anoxic hypolimnion

Bacterioplankton biomass and dark fixation of inorganic carbon were measured in the highly humic (water colour up to 550 mg Pt l^?1) and acidic lake, Mekkojärvi. Strong thermal and chemical stratification developed in the water column early in spring and led rapidly to anoxia in the hypolimnion, which extended to less than 1.0 m from the surface. In the epilimnion only small bacteria were abundant. In the anoxic zone both the abundance and the mean size of bacteria were considerably higher than in the epilimnion. These differences are thought to be the result of different grazing pressure from zooplankton in the two zones. In late summer a high concentration of bacteriochlorophyll d in the upper hypolimnion indicated a high density of photosynthetic bacteria. Bacterial biomass was similar to that of phytoplankton in the epilimnion, but 23 times higher in the whole water column. In August, dark fixation of inorganic radiocarbon in the anaerobic zone was 51% of the total ¹⁴C-incorporation and the contribution of light fixation was only 5.4%. In the polyhumic Mekkojarvi, bacterioplankton was evidently a potentially significant carbon source for higher trophic levels, but bacterioplankton production could not be supported by phytoplankton alone. Allochthonous inputs of dissolved organic matter probably support most of the bacterial production.     

Zooplankton and bacteria contribution to phosphorus and nitrogen internal cycling in a tropical and eutrophic reservoir: Pampulha Lake,Brazil

Nutrient regeneration and respiration rates of natural zooplankton from a tropical reservoir were experimentally measured. Excretion rates of ammonia (E_a), orthophosphate (E_p) and community respiration rates (R) were estimated considering the variations in the concentrations of ammonia, orthophosphate and dissolved oxygen between control and experimental units. The ranges obtained for these rates from the 2 h assays were E_a = 1.95–4.95 μg N-NH₄ · mg · DW⁻¹ · h⁻¹; E_p = 0.12–0.76 μg P-PO₄ mg DW⁻¹ · h⁻¹. Respiratory rates were quite constant (R = 0.01–0.02 mg O₂ · mg DW⁻¹ · h⁻¹). The uptake of nutrients due to bacteria can affect the experimental determination of excretion rates of zooplankton. Orthophosphate release increased from 0.28 to 0.82 μg P-PO₄ · mg DW⁻¹ · h⁻¹ when bacterial activity was depleted by antibiotic addition in experimental vessels (Exp IV). This demonstrates that free living bacteria are able to consume promptly most phosphorus excreted by zooplankton. Ammonia excretion rates were lower in experimental units containing antibiotics. Lower excretion rates were also obtained with longer exposure times and higher biomass levels in the experimental units. Finally, this study also showed that zooplankton excretion can affect significantly turn over rates of total phosphorus in Pampulha Reservoir. In some periods, specially during the dry season when zooplankton biomass was very high, phosphorus release by zooplankton, during one single day, can be as high as 40% of the total phosphorus content in lake water (Turn over time = 2.5 days).     

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.     

Modelling phosphorus transfer rates in lake water

A modification of Lean's (1973a,b) four-compartment model of the transfer of phosphorus between soluble and particulate forms in the epilimnion of lakes is necessary to make the model consistent with certain observed transfer curves. By splitting the particulate compartment into two distinct fractions and incorporating appropriate transfer coefficients, differential rates of exchange with the particulate compartment can be simulated. At least two different rates of exchange with the particulate compartment appear to be necessary to make the model consistent with experimentally observed, diphasic phosphorus transfer curves.     

The Significance of Zooplankton in the Cycling of Phosphorus in Lakes of Different Trophic Categories

The species composition, number, biomass and phosphorus excretion rate (PER) of zooplankton (Crustacea and Rotatoria) in 8 mesotrophic and eutrophic lakes were studied. PER were calculated using specific expenditures on metabolism estimated from the oxygen uptake, phosphorus content in zooplankton and field observations of number and biomass of planktonic Rotatoria and Crustacea. The zooplankton biomass and PER increased from mesotrophic to eutrophic lakes. The values of PER were compared with primary production and spring phosphorus concentration. The relative significance of zooplankton in phosphorus cycling declined with increasing trophic category of lakes.     

Changes in bacterioplankton community structure and activity with depth in a eutrophic lake as revealed by 5S rRNA analysis

The community structure of bacterioplankton was studied at different depths (0 to 25 m) of a temperate eutrophic lake (Lake Plusssee in northern Germany) by using comparative 5S rRNA analysis. The relative amounts of taxonomic groups were estimated from 5S rRNA bands separated by high-resolution electrophoresis. Comparison of partial 5S rRNA sequences enabled detection of changes in single taxa over space and during seasons. Overall, the bacterioplankton community was dominated by 3 to 14 abundant (>4% of the total 5S rRNA) taxa. In general, the number of 5S rRNA bands (i.e., the number of bacterial taxa) decreased with depth. In the fall, when thermal stratification and chemical stratification were much more pronounced than they were in the spring, the correlation between the depth layers and the community structure was more pronounced. Therefore, in the fall each layer had its own community structure; i.e., there were different community structures in the epilimnion, the metalimnion, and the hypolimnion. Only three 5S rRNA bands were detected in the hypolimnion during the fall, and one band accounted for about 70% of the total 5S rRNA. The sequences of individual 5S rRNA bands from the spring and fall were different for all size classes analyzed except two bands, one of which was identified as Comamonas acidivorans. In the overall analysis of the depth profiles, the diversity in the epilimnion contrasted with the reduced diversity of the bacterioplankton communities in the hypolimnion, and large differences occurred in the composition of the epilimnion at different seasons except for generalists like C. acidivorans.     

Inter- and intra-annual variability in the phytoplankton community of a high mountain lake: the influence of external (atmospheric) and internal (recycled) sources of phosphorus

1. The inter‐ and intra‐annual changes in the biomass, elemental (carbon (C), nitrogen (N) and phosphorus (P)) and taxonomical composition of the phytoplankton in a high mountain lake in Spain were studied during 3 years with different physical (fluctuating hydrological regime) and chemical conditions. The importance of internal and external sources of P to the phytoplankton was estimated as the amount of P supplied via zooplankton recycling (internal) or through ice‐melting and atmospheric deposition (external). 2. Inter‐annual differences in phytoplankton biomass were associated with temperature and total dissolved phosphorus. In 1995, phytoplankton biomass was positively correlated with total dissolved phosphorus. In contrast, the negative relationship between zooplankton and seston biomass (direct predatory effects) and the positive relationship between zooplankton P excretion and phytoplankton biomass in 1997 (indirect P‐recycling effects), reinforces the primary role of zooplankton in regulating the total biomass of phytoplankton but, at the same time, encouraging its growth via P‐recycling. 3. Year‐to‐year variations in seston C : P and N : P ratios exceeded intra‐annual variations. The C : P and N : P ratios were high in 1995, indicating strong P limitation. In contrast, in 1996 and 1997, these ratios were low during ice‐out (C : P < 100 and N : P < 10) and increased markedly as the season progressed. Atmospheric P load to the lake was responsible for the decline in C : P and N : P ratios. 4. Intra‐annual variations in zooplankton stoichiometry were more pronounced than the overall differences between 1995 and 1996. Thus, the zooplankton N : P ratio ranged from 6.9 to 40.1 (mean 21.4) in 1995, and from 10.4 to 42.2 (mean 24.9) in 1996. The zooplankton N : P ratio tended to be low after ice‐out, when the zooplankton community was dominated by copepod nauplii, and high towards mid‐ and late‐season, when these were replaced by copepodites and adults. 5. In 1995, the minimum demands for P of phytoplankton were satisfied by ice‐melting, atmospheric loading and zooplankton recycling over 100%. In order of importance, atmospheric inputs (> 1000%), zooplankton recycling (9–542%), and ice‐melting processes (0.37–5.16%) satisfied the minimum demand for P of phytoplankton during 1996 and 1997. Although the effect of external forces was rather sporadic and unpredictable in comparison with biologically driven recycle processes, both may affect phytoplankton structure and elemental composition. 6. We identified three conceptual models representing the seasonal phosphorus flux among the major compartments of the pelagic zone. While ice‐melting processes dominated the nutrient flow at the thaw, biologically driven processes such as zooplankton recycling became relevant as the season and zooplankton ontogeny progressed. The stochastic nature of P inputs associated with atmospheric events can promote rapid transitional changes between a community limited by internal recycling and one regulated by external load. 7. The elemental composition of the zooplankton explains changes in phytoplankton taxonomic and elemental composition. The elemental negative balance (seston N : P < zooplankton N : P, low N : P recycled) during the thaw, would promote a community dominated by species with a high demand for P (Cryptophyceae). The shift to an elemental positive balance (seston N : P > zooplankton N : P, high N : P recycled) in mid‐season would skew the N : P ratio of the recycled nutrients, favouring dominance by chrysophytes. The return to negative balance, as a consequence of the ontogenetic increase in zooplankton N : P ratio and the external P inputs towards the end of the ice‐free season, could alleviate the limitation of P and account for the appearance of other phytoplankton classes (Chlorophyceae or Dinophyceae).     

Modelling phosphorus fluxes in the hypertrophic Loosdrecht Lakes

A dynamic, deterministic model is presented to simulate the phosphorus cycle and plankton growth in the shallow, hypertrophic Loosdrecht Lakes (The Netherlands) before and after restoration measures. Besides inorganic phosphorus (SRP) in both the surface water and the interstitial water, the model comprises three algal groups, zooplankton, fish, detritus, zoobenthos and upper sediment (all modelled both in carbon and in phosphorus). Within the model system, the phosphorus cycle is completely closed. Carbon and phosphorus are described independently, so that the dynamics of the P/C ratios can be modelled. Sediment processes are described in a simplified form.Simulated values are largely within the range of observed ones. The detrital fraction of the seston (=phytoplankton+detritus) varies from 50–60% in summer to about 90% in winter. SRP in the surface water is very low during most of the year. Sensitivity for external phosphorus input is larger for algal and detrital P than for algal and detrital C and chlorophyll-a. So the P/C ratio of the seston decreases following restoration measures, as is observed in the lakes, while the much higher P/C ratios of zooplankton and fish remain constant. Phosphorus mobilisation from the sediment decreases with decreasing external input. Adaptation of the model system to the reduced loading takes place within about two years.Sources of uncertainty in the model include the limited knowledge on selective grazing as well as on mortality and mineralisation processes.     

Effects of zooplankton on nutrient availability and seston C : N : P stoichiometry in inshore waters of Lake Biwa, Japan

Forty-eight-hour experimental manipulations of zooplankton biomass were performed to examine the potential effects of zooplankton on nutrient availability and phytoplankton biomass (as measured by seston concentration) and C : N : P stoichiometry in eutrophic nearshore waters of Lake Biwa, Japan. Increasing zooplankton, both mixed-species communities and Daphnia alone, consistently reduced seston concentration, indicating that nearshore phytoplankton were generally edible. The zooplankton clearance rates of inshore phytoplankton were similar to rates measured previously for offshore phytoplankton. Increased zooplankton biomass led to increased concentrations of nutrients (NH₄-N, soluble reactive phosphorus [SRP]). Net release rates were higher than those found in previous measurements made offshore, reflecting the nutrient-rich nature of inshore seston. Zooplankton nutrient recycling consistently decreased TIN : SRP ratios (TIN = NH₄ + NO₃ + NO₂). This effect probably resulted from the low N : P ratios of nearshore seston, which were lower than those commonly found in crustacean zooplankton and thus resulted in low retention efficiency of P (relative to N) by the zooplankton. Thus, zooplankton grazing inshore may ameliorate algal blooms due to direct consumption but tends to create nutrient supply conditions with low N : P, potentially favoring cyanobacteria. In comparison with previous findings for offshore, it appears that potential zooplankton effects on phytoplankton and nutrient dynamics differ qualitatively in inshore and offshore regions of Lake Biwa. Received: September 4, 2000 / Accepted: January 23, 2001     

Daphnia population growth but not moulting is a substantial phosphorus drain for phytoplankton

SUMMARY 1. Negative effects of zooplankton on the availability of phosphorus (P) for phytoplankton as a result of the retention of nutrients in zooplankton biomass and the sedimentation of exoskeletal remains after moulting, have been recently proposed. 2. In a mesocosm study, the relative importance of these mechanisms was tested for the freshwater cladoceran Daphnia hyalina×galeata. A total of 13 mesocosm bags was suspended in a mesotrophic German lake during summer 2000 and fertilised with inorganic P in order to obtain a total nitrogen to total P ratio closer to the Redfield ratio. D. hyalina×galeata was then added at a logarithmically scaled density gradient of up to 40 ind. L^?1. Zooplankton densities, dissolved inorganic, particulate organic (seston <100 μm), as well as total nutrient concentrations were monitored. Additionally, nutrient concentrations of sediment water removed from the bottom of the mesocosm bags via a manual pump were determined. 3. Seston carbon (C), seston P and total P were significantly negatively correlated with Daphnia densities. The amount of particulate P (～5–6 μg P L^?1) sequestered from the seston compartment by Daphnia corresponded roughly to the increase of zooplankton biomass (population growth). Soluble reactive phosphorous (SRP) was at all times high (～25–35 μg P L^?1) and possibly unavailable to phytoplankton as a result of P adsorption to calcite during a calcite precipitation event (whiting). P concentrations determined in sediment water were generally <60 μg P m^?2 and thus never exceeded 1% of the total amount of P bound in particulate matter of the overlying water column. 4. Seston C : P ratios followed a polynomial second‐order function: At Daphnia densities <40 ind. L^?1 a positive linear relationship was evident, which is explained by the stronger reduction of P compared with C in seston, and transfer of seston P to zooplankton. Highest seston C : P ratios of ～300 : 1 were observed at Daphnia densities of ～30–50 ind. L^?1, which is in agreement with proposed threshold values limiting Daphnia reproductive growth. At Daphnia densities >40–50 ind. L^?1 C : P ratios were decreased because of the strong reduction of seston C at close to constantly low seston P‐values of ～3–4 μg P L^?1. 5. At least for Daphnia, it may be concluded that – unlike population growth – the sedimentation of faecal pellets and carapaces after moulting seem negligible processes in pelagic phosphorus dynamics.     

On the cost of vertical migration: are feeding conditions really worse at greater depths?

SUMMARY 1. The ultimate explanation for diel vertical migration (DVM) of zooplankton is the avoidance of visual predation in surface waters. Studies on migrating zooplankton have shown that remaining in the cold and food-poor hypolimnion during the day, however, has demographic costs. Higher temperatures and greater food concentrations in the surface waters are thought to be the main reasons why Daphnia species move upwards at night.
2. In this study, we investigated the growth condition of daphniids raised on seston taken from different depths from a lake with and without a deep-water chlorophyll maximum.
3. Juvenile growth rates of Daphnia galeata x hyalina from the lake without a deep-water chlorophyll maximum were similar for all treatments. After temperature correction, however, growth rates were significantly higher on seston taken from the surface layers.
4. In contrast, in the lake with the deep-water chlorophyll maximum, D. galeata growth rates were higher in deeper strata, even after temperature correction. Although this lake had a weak temperature gradient, D. galeata left the food-rich strata at night and migrated into the surface food-poor environment. Invertebrate predation and oxygen depletion are probably not the reasons for the nocturnal upward migration into the surface strata. Therefore, we assume that D. galeata migrates upwards to take advantage of higher temperatures. Using several temperature–egg-development models, we could not, however, fully explain this behaviour.     

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.     

Long-term trends of epilimnetic and hypolimnetic bacteria and organic carbon in a deep holo-oligomictic lake

We analysed the long-term dynamics (1980–2007) of hypolimnetic and epilimnetic bacterial abundances and organic carbon concentrations, both dissolved (DOC) and particulate (POC), in the deep holo-oligomictic Lake Maggiore, included in the Southern Alpine Lakes Long-Term Ecological Research (LTER) site. During the 28 years of investigation, bacterial abundance and POC concentrations did not decrease with declining phosphorus concentrations, while DOC concentrations showed a pronounced decrease in the epi- and hypolimnion. We used the annual mean total lake heat content and total annual precipitation as climate-related variables, and in-lake total phosphorus as a proxy for trophic state. The model (forward stepwise regression, FSR) showed that reduced anthropogenic pressure was more significant than climate change in driving the trend in DOC concentrations. Bacterial dynamics in the hypolimnion mirrored the fluctuations observed in the epilimnion, but average cell abundance was three times lower. The FSR model indicates that bacterial number variability was dependent on POC in the epilimnion and DOC in the hypolimnion. In the hypolimnion, cell biovolumes for rod and coccal morphotypes were significantly larger than in the epilimnion.