Determinants of seston C : P-ratio in lakes

1. The ratio of carbon to phosphorus (C : P) in seston is a major determinant of energy transfer in aquatic food webs and may vary more than an order of magnitude owing to various extrinsic and intrinsic factors. In this study, the determinants of C : P‐ratios in lake particulate matter (seston) was assessed in 112 Norwegian lakes, covering a C : P (atomic ratio) from 24 to 1842 (mean 250). 2. No overall effects of lake area, season or latitude on C : P was detected. Particulate P, but not particulate C, correlated with C : P. Multivariate analysis including a range of lake properties revealed total dissolved P, as the major determinant of sestonic C : P, with the fraction of detritus in total seston, chlorophyll or Secchi depth and lake colour as significant contributors. Together these parameters explained 30% of observed variance if using dissolved P and 81% if using total P as input variable to the multivariate model. 3. Chlorophyll and Secchi depth were highly correlated and substitutable in the analysis. Phytoplankton community composition did not affect seston C : P, probably reflecting the fact that live phytoplankton generally contributed <25% of the seston pool. 4. Total P correlated positively with C : P and is the key determinant of phytoplankton biomass and thus Secchi depth; the latter parameters contributed negatively to seston C : P, probably owing to increased light attenuation. These lake data thus support the light : nutrient ratio hypothesis, i.e. that high light and low P cause skewed uptake ratios of C to P. 5. Zooplankton biomass in general and Daphnia biomass in particular, was negatively correlated with C : P, probably reflecting a negative impact of poor seston quality at high C : P. Zooplankton grazing and nutrient recycling may also have contributed to a negative correlation between zooplankton biomass and sestonic C : P.     

Seasonal changes of C:P ratios of seston, bacteria, phytoplankton and zooplankton in a deep, mesotrophic lake

• 1 The C:P ratios of seston, bacteria, phytoplankton and zooplankton were measured twice a week in situ in mesotrophic, large and deep Lake Constance from April to December 1995. Except for zooplankton, a strong seasonality was exhibited with low C:P ratios during P‐enriched early spring conditions and high values during P‐depleted summer conditions.
• 2 Molar C:P ratios of seston varied between 180:1 and 460:1 demonstrating moderate phosphorus limitation in spring and during the clear‐water phase, and strong limitation for the rest of the season. The sestonic C:P ratio increased significantly during two decades of re‐oligotrophication of Lake Constance, reflecting an enhanced phosphorus limitation of the plankton community in summer. Molar C:P ratios of bacteria and phytoplankton varied seasonally between 50:1 and 130:1 and 180:1 and 500:1, respectively, and indicate carbon or light limitation in winter and phosphorus limitation in summer. Zooplankton had a molar C:P ratio of about 124:115 which was nearly constant throughout the seasons.
• 3 These differences in the C:P ratios of planktonic organisms have direct implications for phosphorus recycling within the food web as C:P ratios of excreta should be highly variable.

     

Can results from limnocorral experiments be transferred to in situ conditions?

In 1982–1984 eight limnocorral (LC) experiments, each lasting for two weeks, were performed in mesotrophic Lake Lucerne, Switzerland, to study the effects of biomanipulation (removal of crustaceans by 95 µm filter nets) on zooplankton — phytoplankton relationships and epilimnetic carbon and phosphorus fluxes. Seston concentrations and to a lesser extent primary production rates were reduced in control LCs through zooplankton grazing, and settling flux increased through fecal pellet production. But C and P regeneration were not significantly affected.We found several indications that the LCs, despite of their large size ( 70 m³, 3 m in diameter and 11 m long) were artificial systems when compared to the surrounding lake: The eddy diffusion was diminished by about one order of magnitude, nutrients depleted (but phosphorus was supplied to the LCs), the phyto- and zooplankton showed lower standing crops in the control LCs than in the lake, the phytoplankton showed a shift from nannoplankton to netplankton, the crustacean zooplankton was mostly limited in their vertical migration, and the POC and PP sedimentation rates were increased. Moreover, in our set of experiments we always found outliers, which may have been caused by the different in situ conditions at the beginning of the experiments, and further enhanced by the complexity of the enclosed system.The problems of the LC-technique, such as replicability, scaling (size and time) and data extrapolation are discussed. The impact of crustacean zooplankton on particulate matter, i.e. seston reduction and sedimentation enhancement, can be extrapolated on a qualitative rather than on a quantitative basis for the Lake Lucerne ecosystem.     

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     

Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

The Influence of Crustacean Zooplankton on the Size Structure of Algal Biomass and Suspended and Settling Seston (Biomanipulation in Limnocorrals II)

In a limnocorral (LC) experiment performed in mesotrophic Lake Lucerne, Switzerland, a control LC containing a planktonic community similar to that in the lake was compared with a LC where the large zooplankton had been removed by filtration at the beginning of the experiment. The herbivorous zooplankton (mainly Daphnia galeata and D. hyalina) reduced algal biomass and primary production, however did not influence the size structure of the phytoplankton and the relative amount of different size classes contributing to the total primary production. Likewise, the seston (POC and PP) concentrations were diminished without changes in particle size composition. Since herbivorous zooplankton transform smaller particles into larger particles (fecal aggregates) through their grazing activity and thus enhanced particle formation owing to coagulation and microbial activity, sedimentation losses of POC and PP were increased in the LC with zooplankton. The importance of planktonic and sestonic particle size structure in aquatic ecosystems is stressed, as the smallest algae (< 12 μm) contributed 80 % of the primary production, which is in contrast to the 33 % contributed to the total POC sedimentation by this particle size class.     

Seasonal correlations of elemental and ω3 PUFA composition of seston and dominant phytoplankton species in a eutrophic Siberian Reservoir

The elemental and fatty acid composition of seston was studied for 3 years, from May to October, in a small Reservoir. Under comparatively low C:P ratio, multivariate canonical analysis revealed no straightforward simple correlations between phosphorus and single ω3 PUFA species, but complex significant interaction between elemental composition (stoichiometry) of seston and total sestonic ω3 PUFA as a whole. Since sestonic C, P and N were found to originate mostly from phytoplankton, the contents of particulate elements and PUFA were attributed to single species in periods of their pronounced dominance. Phytoplankton species of genera of Stephanodiscus, Peridinium, Gomphosphaeria, Planktothrix and Anabaena in periods of their pronounced dominance had relatively constant species-specific elemental and PUFA composition. Phytoplankton species significantly differed in their elemental and PUFA composition, as well as in ratios of C:N, N:P, PUFA:P and partly C:P that indicate food quality for zooplankton. Hence, there were no phytoplankton species of clearly high or low nutritional value. All of phytoplankters, or at least detritus, that originated from them, may meet specific elemental and biochemical requirements of specific groups of zooplankton. Dividing phytoplankton on basis of their elemental and biochemical composition, i.e., nutrition quality, into large taxa (cyanobacteria, diatoms, etc.) appeared to be too coarse for assessing nutritional value for zooplankton.     

Seasonal seston stoichiometry: effects on zooplankton in cyanobacteria-dominated lakes

Seasonal dynamics in elemental composition [carbon (C), nitrogenand phosphorus (P)] of seston and zooplankton were studied overseveral years in three hypereutrophic Dutch lakes with persistentdominance and high biomass of cyanobacteria. In all three lakes,there was a strong pattern with decreased P-content and increasedC:P ratio in seston (<150 µm) coinciding with the increasein water temperature. The seston C:P ratios (at:at) were morethan doubled with the rising temperature, i.e. from 200 (at:at)in winter to 500 in summer. Sestonic C:P ratios increased overthe growing season, suggesting that seasonal dynamics amongautotrophs with high P-uptake in winter and support of subsequentphytoplankton growth by consumption of internal cellular P (P-quota)was the main cause of low sestonic P contents in late summer.This could, however, occur in concert with a physiologicallydriven decrease in cell-specific P at higher temperatures insummer. In contrast, the annual variation of C:P ratios of thezooplankton fraction was only 10% of that of seston. The variationsof C:P ratios of the zooplankton were, nevertheless, stronglycorrelated with those of seston. For most of the summer, sestonC:P ratios were far above the threshold ratio for P-limitationin Daphnia and other P-demanding species. This will pose furtherconstraints on growth performance of Daphnia in these lakes,thus adding to the fish predation pressure and the poor foodquality of cyanobacteria per se. The low grazing pressure causesa high biomass of low-quality autotrophs, promoting a stablestate with low trophic transfer efficiency.     

Food quality for Daphnia in humic and clear water lakes

1. Growth and reproduction of Daphnia fed lake seston were measured in two categories of meso‐ to eutrophic lakes differing with respect to terrestrial organic matter influence (humic and clear water lakes). The content of highly unsaturated fatty acids (HUFA), P and N, as well as the taxonomical composition of seston were analysed. 2. Seston HUFA and C : P ratios were similar between lake categories, whereas C : N ratios were lower in the clear water lakes in both spring and summer. Despite the similarity in HUFA and P content of seston, Daphnia growth rate, clutch size and the proportion of gravid females were, respectively, about 1.5, 3 and 6 times higher in the clear water lakes. 3. Differences in growth and reproduction were related to a combination of higher N content and good fatty acid quality of the seston in the clear water lakes. Relatively high biomass of edible algae, such as Rhodomonas sp. and Cryptomonas sp., in the clear water lakes, and differences in water pH likely contributed to the observed differences in Daphnia growth and reproduction between lake categories. Additionally, it is possible that Daphnia was energy limited in the humic lakes despite high particulate organic carbon (POC) concentrations, as the contribution of non‐algal and detrital C to the POC pool was high. 4. Our results suggest that dietary HUFA content has the potential to improve herbivore growth and reproduction if N and P are not limiting. N merits more attention in studies of zooplankton nutrition.     

Seasonal and vertical variation in the C : P ratio of suspended and settling seston of lakes

In the absence of field data it is often assumed that the C : P ratio of the seston in lakes corresponds to the ideal ratio of 106 : 1, reported by Redfield et al. (1963) for marine phytoplankton. However, we found out that in nearly all lakes studied C : P ratios deviate systematically from this value especially during periods, when phosphorus is growth limiting. Thus, during summer the sestonic C : P ratio generally exceeds 106 in the epilimnion, but decreases with depth in the hypolimnion. Possible mechanisms which may lead to these seasonal and spatial changes in the C : P ratio are discussed.     

Influence of biochemical,mineral and morphological features of natural food on tropical cladocerans

Five laboratory experiments were performed to evaluate the effect of supplements of fatty acids and a green alga on the individual growth and reproduction of three species of tropical cladocerans Ceriodaphnia richardi, Daphnia ambigua, and D. gessneri feeding on natural seston from the Brazilian Lake Monte Alegre. Cohorts of newborns from cultivated females were submitted to one of the following treatments: (1) Natural seston, (2) Natural seston + microcapsules of EPA and DHA or linoleic and linolenic fatty acids, (3) Natural seston + oil-free microcapsules, and (4) Natural seston + green alga Scenedesmus spinosus (1 mg C l⁻¹). Particulate organic carbon, algal carbon, C:P ratios of seston and green alga, polyunsaturated fatty acids (PUFA) content of seston and cladocerans, as well as phytoplankton composition, size, and shape were measured. The addition of fatty acids to seston did not significantly enhance growth and reproduction of the cladocerans, suggesting that sestonic PUFA content is sufficient for promoting cladoceran development, even in the cool–dry season when the fatty acids used in the experiments were 5–10 times lower in the seston than in the warm–wet season. Despite high C:P molar ratios in most experiments, there was only one indication of growth limitation by P. Reproduction was more affected than individual growth on some occasions by food quantity (energy) caused apparently by algal size, morphology, and digestion resistance. Energy availability, which is affected by algae morphological characteristics, seems to prevail over PUFA and P in controlling growth and reproduction of cladocerans in tropical Lake Monte Alegre.     

Elemental stoichiometry of lower food web components in arctic and temperate lakes

Lakes were surveyed to assess the potential patterns of latitudinalvariation in carbon:nitrogen:phosphorus (C:N:P) stoichiometryof lower food web components. Thirty-four lakes were surveyedat an arctic latitude (68°38'N, 149°38'W) and 10 lakesat a temperate latitude (46°13'N, 89°32'W) during 1997.The temperate data set was augmented with earlier survey resultsemploying similar methods. It was hypothesized that differencesin environmental variables across latitude would cause differencesin community C:N:P ratios, leading to differences in trophicinteractions. Physical measurements (light, temperature), sestonand zooplankton were collected from each lake. Seston and zooplanktonwere analyzed for C, N and P content, and zooplankton were countedand measured for biomass estimates. The degree of trophic interactionbetween seston and zooplankton was assessed by (i) measuringelemental imbalances between seston and zooplankton and (ii)calculating the potential recycling ratio by the zooplanktoncommunity available for seston. Seston C:nutrient, but not N:P,ratios were higher in temperate than arctic lakes. Conversely,arctic zooplankton had higher C:nutrient, but not N:P, ratiosthan zooplankton in temperate lakes. Elemental imbalances weregreater in temperate than in arctic lakes, but N:P stoichiometryof potential zooplankton recycling was nearly identical betweenthe two latitudes. Zooplankton community C:N:P ratios were notrelated to either latitude or seston C:N:P. In accordance withstoichiometric theory, relative abundances of calanoid copepodswere positively correlated with seston C:N in temperate lakes.Additionally, relative abundances of Daphnia were negativelycorrelated with seston C:N ratios in temperate and arctic lakes,and positively correlated with N:P ratios in the arctic. Ingeneral, these results suggest that seston and zooplankton communitystoichiometry differ across latitude, and these differenceshave the potential to affect trophic interactions.     

The stoichiometry of N and P in the pelagic zone of Castle Lake, California

We measured the concentrations, as well as lake-wide amounts,of nitrogen (N) and phosphorus (P) in dissolved, seston andzooplankton pools throughout the water column of Castle Lake,California, during summer, 1991. This allowed us to determinethe stoichiometric ratios of important elements in each pool(C:N, C:P, N:P) as well as for the entire lake. Dissolved andseston pools were the predominant storage compartments for bothN and P; zooplankton never contained >5% of N or 10% of Plake wide. However, by late summer, the concentrations of Pin seston and in zooplankton were similar in the upper portionsof the water column, suggesting that changes in food web structurethat alter zooplankton biomass and community composition (andhence elemental storage in the zooplankton) may produce significantshifts in nutrient storage among pelagic pools. Lake-wide levelsof dissolved N were largely constant over the study period;however, lake-wide dissolved P increased. These dynamics suggestedthat the majority of nutrients stored in dissolved pools wereunavailable for phytoplankton growth. N:P and C:P ratios indicatedthat Castle Lake phytoplankton became severely deficient inP during the course of our observations. These ratios also greatlyexceeded recently reported threshold values for elemental constraintson growth and reproduction for several species of zooplankton.The ratio of N to P in the zooplankton pool was relatively constantand consistently lower than that in the sestion. As a result,the predicted N:P ratio of zooplankton-regenerated nutrientsexceeded the N:P ratio of the seston, implying that zooplanktonnutrient regeneration further skewed N and P supply ratios,and potentially enhanced P limitation of phytoplankton in CastleLake. ¹Present address: Department of Biology, Box 19498, Universityof Texas at Arlington, Arlington, TX 76019, USA     

Effects of nitrogen enrichment on zooplankton biomass and N:P recycling ratios across a DOC gradient in northern-latitude lakes

We used data from whole-lake studies to assess how changes in food quantity (phytoplankton biomass) and quality (phytoplankton community composition, seston C:P and N:P) with N fertilization affect zooplankton biomass, community composition and C:N:P stoichiometry, and their N:P recycling ratio along a gradient in lake DOC concentrations. We found that despite major differences in phytoplankton biomass with DOC (unimodal distributions, especially with N fertilization), no major differences in zooplankton biomass were detectable. Instead, phytoplankton to zooplankton biomass ratios were high, especially at intermediate DOC and after N fertilization, implying low trophic transfer efficiencies. An explanation for the observed low phytoplankton resource use, and biomass responses in zooplankton, was dominance of colony forming chlorophytes of reduced edibility at intermediate lake DOC, combined with reduced phytoplankton mineral quality (enhanced seston N:P) with N fertilization. N fertilization, however, increased zooplankton N:P recycling ratios, with largest impact at low DOC where phytoplankton benefitted from light sufficiently to cause enhanced seston N:P. Our results suggest that although N enrichment and increased phytoplankton biomass do not necessarily increase zooplankton biomass, bottom-up effects may still impact zooplankton and their N:P recycling ratio through promotion of phytoplankton species of low edibility and altered mineral quality.

     

Inter-and intraspecific differences in fatty acid composition of freshwater crustacean zooplankton

1. The composition of fatty acids (FA) from C14 to C18 was measured for edible seston and for individual Daphnia galeata , Diaphanosoma brachyurum and Eodiaptomus japonicus from Lake Biwa using a pyrolysis–gas chromatography (Py–GC) method. Based on the relative abundance of the FA, inter-and intraspecific differences in composition were examined.
2. Among the FA, C16 : 0 was the most abundant, both in the three zooplankton species and in edible seston smaller than 20 μm, suggesting that the composition of the zooplankton was roughly reflected by their diet. However, the abundance of C18 : 1 relative to C16 : 1 and C18 : 0 was much higher in each zooplankton species than in the diet.
3. Comparison of the relative abundance of FA among the three zooplankton species revealed that intraspecific differences in FA composition are greater than interspecific differences. These results indicate that variability in FA composition is not necessarily species-specific, and can be obscured by variation in composition between individuals.     

Pelagic C:N:P Stoichiometry in a Eutrophied Lake: Responses to a Whole-Lake Food-Web Manipulation

Changes in the ecological stoichiometry of C, N, and P in the pelagic zone are reported from a whole-lake manipulation of the food web of Lake 227, an experimentally eutrophied lake at the Experimental Lakes Area, Canada. Addition of northern pike eliminated populations of planktivorous minnows by the third year (1995) after pike introduction, and in the fourth year after pike addition (1996), a massive increase in the abundance of the large-bodied cladoceran Daphnia pulicaria occurred. Accompanying this increase in Daphnia abundance, zooplankton community N:P declined, seston concentration and C:P ratio decreased, and dissolved N and P pools increased. During peak abundance, zooplankton biomass comprised a significant proportion of total epilimnetic phosphorus (greater than 30%). During the period of increased Daphnia abundance, concentrations of dissolved inorganic nitrogen (TIN) increased more strongly than dissolved phosphorus (TDP), and thus TIN:TDP ratios were elevated. Sedimentation data indicated that increased grazing led to greatly reduced residence times of C, N, and especially P in the water column during 1996. Finally, previously dominant N-fixing cyanobacteria were absent during 1996. Our results show that strong effects of food-web structure can occur in eutrophic lakes and that stoichiometric mechanisms play a potentially important role in generating these effects.     

Zebra mussels mediate benthic–pelagic coupling by biodeposition and changing detrital stoichiometry

1. The zebra mussel ( Dreissena polymorpha ) is one of the most successful invasive species; it has colonised many aquatic systems in Europe and North America with strong impacts on various ecosystem processes. The effect of D. polymorpha filtration on pelagic seston concentrations has been quantified in several studies, but the magnitude and stoichiometry of the transfer of sestonic biomass into benthic detritus by D. polymorpha and the accompanying enrichment of the benthic habitat is still under-investigated.
2. We studied biodeposition by zebra mussels in two series of laboratory experiments with the food algae Cryptomonas erosa and Scenedesmus obliquus . We also measured the year-round biodeposition rate under natural conditions in the oligotrophic Lake Constance.
3. In all experiments, zebra mussel biodeposition was linearly related to seston concentration. In the field, the relationship changed with a seasonal shift in algal composition and lower biodeposition rates during the spring algal bloom.
4. For both algal species in laboratory experiments, biodeposited material was depleted in phosphorous at an algal concentration ≤0.6 mg ash-free dry mass L⁻¹, but not at higher concentrations. This effect was not observed in the field, probably because of high variation in C : N : P stoichiometry.
5. By mediating the transfer of pelagic resources into the benthos zebra mussels provide a sufficient amount of detritus for benthic invertebrates, especially during summer. Thus, material biodeposited by the mussels might increase benthic secondary production from pelagic resources, and zebra mussels are important mediators of this flux of organic matter from the pelagic zone into the benthos.     

The relationship between nutrients and trophic‐level biomass in turbid tropical ponds

1. The total phosphorus–algal biomass relationship from a set of turbid tropical ponds in Kenya was compared with predictions derived from surveys of temperate and subtropical lakes. Despite high concentrations of total phosphorus (TP) (up to 797 μg L ^–1) and inorganic turbidity (up to 800 mg L^–1), the log–log relationship between algal biomass and TP was steeper than expected.
2. No evidence of nitrogen limitation was found at high TP, and total nitrogen (TN):TP ratios were higher than in lakes with similar TP levels studied previously. High TN:TP ratios may be a consequence of excretion by cattle into the ponds, a nutrient source characterized by a high ratio of available N to available P.
3. Despite extremely high turbidity, the ratio of mixed layer depth to euphotic depth was generally low because these ponds are shallow (≤ 2 m), and was not related to algal yield. A positive relationship was also found between TP and zooplankton biomass, and between TP and the density of the zooplanktivorous bug, Anisops . In contrast, no relationship was found between fish biomass and TP, algal biomass or zooplankton biomass.     

Evidence of phosphorus-limited individual and population growth of Daphnia in a Canadian Shield lake

We performed bag experiments in a Canadian Shield lake with generally high seston (suspended food particles mainly composed of algae) carbon (C):phosphorus (P) ratios, and investigated the responses of individual and population growth of herbivorous Daphnia dentifera on their abundance with (+P) and without (−P) a phosphorus enrichment to lake water. In both treatments, increased abundance of D. dentifera reduced seston C concentration and was accompanied by decreases in population and individual growth rates. However, P-enrichment increased seston P concentration and then reduced seston C:P ratio from 400–700 to ca 100 (by atoms). As a result, both individual and population growth rates were significantly higher in the +P treatment at all animal abundances even though seston C concentrations were similar between the treatments. The magnitude of the growth enhancement by the P-enrichment was independent of animal abundance. Stepwise regression analyses revealed that 71 and 90% of the variance in the population and individual growth rates, respectively, were explained by seston C and P concentrations, and that the contribution of the seston P concentration was roughly the same as that of seston C. Such joint effects of seston C and P indicate that food quality (P content) as well as food quantity (C concentration) can influence Daphnia not only at the level of individual growth but also at the level of population dynamics in P-limited lakes. Our results thus strongly corroborate the hypothesis that the population development of a key herbivore Daphnia in P-limited Canadian Shield lakes is inhibited by the direct effects of P-limited food on individual growth, which weaken the strength of trophic cascading interactions starting from piscivorous fish through planktivorous fish and zooplankton to algae.     

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.