Atmospheric nitrogen deposition is associated with elevated phosphorus limitation of lake zooplankton

Here, we present data that for the first time suggests that the effects of atmospheric nitrogen (N) deposition on nutrient limitation extend into the food web. We used a novel and sensitive assay for an enzyme that is over‐expressed in animals growing under dietary phosphorus (P) deficiency (alkaline phosphatase activity, APA) to assess the nutritional status of major crustacean zooplankton taxa in lakes across a gradient of atmospheric N deposition in Norway. Lakes receiving high N deposition had suspended organic matter (seston) with significantly elevated carbon:P and N:P ratios, indicative of amplified phytoplankton P limitation. This P limitation appeared to be transferred up the food chain, as the cosmopolitan seston‐feeding zooplankton taxa Daphnia and Holopedium had significantly increased APA. These results indicate that N deposition can impair the efficiency of trophic interactions by accentuating stoichiometric food quality constraints in lake food webs.     

UV radiation and phosphorus interact to influence the biochemical composition of phytoplankton

1. Numerous laboratory studies have shown that food quality is suboptimal for zooplankton growth. However, little is known about how food quality is affected by the interaction of potential global change factors in natural conditions. Using field enclosures in a high altitude Spanish lake, seston was exposed to increasing phosphorus (P) concentrations in the absence and presence of UV radiation (UVR) to test the hypothesis that interactions between these factors affected the biochemical and stoichiometric composition of seston in ways not easily predicted from studies of single factors. 2. Phosphorus enrichment increased the content of total fatty acids (TFA), ω3‐polyunsaturated fatty acids (ω3‐PUFA) and chlorophyll‐a : carbon (Chl‐a : C) and C : N ratios in seston. The pronounced increase in ω3‐PUFA was largely explained by the enhancement of 18:3n‐3 (α‐linolenic acid). In contrast, P‐enrichment lowered the content of highly unsaturated fatty acids (HUFA), the HUFA : PUFA ratio and, at high P loads, seston C : P ratio. Although phytoplankton assemblages dominated by Chlorophytes were not rich in HUFA, seston in the control had substantially higher 20:4n‐6 (arachidonic acid, ARA) content (79% of HUFA) than did P‐enriched enclosures. 3. The UVR increased the content of ω3‐PUFA and TFA in seston at the two ends of the trophic gradient generated at ambient and high concentrations of P, but decreased seston C : P and HUFA at all points on this gradient. ARA was not detected in the presence of UVR. 4. The interaction between P and UVR was significant for seston HUFA and C : P ratios, indicating that the effect of UVR in reducing HUFA (decreased food quality) and C : P ratios (enhanced food quality) was most pronounced at the low nutrient concentrations characteristic of oligotrophic conditions and disappeared as P increased. Therefore, any future increase in UVR fluxes will probably affect most strongly the food quality of algae inhabiting oligotrophic pristine waters although, at least in the Mediterranean region, these effects could be offset by greater deposition of P from the atmosphere.     

Nutritional quality of seston for the freshwater herbivore Daphnia galeata × hyalina: biochemical versus mineral limitations

The quality of natural seston as food for zooplankters can be highly variable. Thus far, experimental evidence on the factors affecting food quality under natural conditions is scarce. Hence, in this study, we set out to investigate how Daphnia galeata 2 hyalina responded to qualitative variation in natural seston. This was done in laboratory experiments where we supplement natural seston from a mesotrophic lake with dissolved phosphorus and emulsions of highly unsaturated "essential" fatty acids. The growth rate of juveniles increased upon the supply of both phosphorus and fatty acids. These results suggest that these phosphorus and highly unsaturated fatty acids are substitutable and thus challenge our existing interpretation/understanding of how herbivore growth is "limited".     

Glacier melting and stoichiometric implications for lake community structure: zooplankton species distributions across a natural light gradient

Glaciers around the globe are melting rapidly, threatening the receiving environments of the world's fresh water reservoirs with significant changes. The meltwater, carried by rivers, contains large amounts of suspended sediment particles, producing longitudinal gradients in the receiving lakes. These gradients may result in changes in the light : nutrient ratio that affect grazer performance by altering elemental food quality. Thus, glacial melting may induce a shift in the phytoplankton carbon : nutrient ratio and hence influence the dominance of herbivorous zooplankton through stoichiometric mechanisms. To test this hypothesis, we combined field and experimental data, taking advantage of a natural light intensity gradient caused by glacial clay input in a deep oligotrophic Patagonian lake. Across this gradient, we evaluated the abundances of two consumer taxa with different phosphorus requirements, the copepod Boeckella gracilipes and the cladoceran Daphnia commutata, using a six‐station transect along the lake. We found significant differences in light : nutrient ratio and stoichiometric food quality of the seston, together with a switch from dominance of P‐rich Daphnia in low carbon : nutrient stations to dominance of low‐P copepods in high carbon : nutrient stations. The laboratory experiments confirmed that the difference in the carbon : nutrient ratio across the gradient is sufficient to impair Daphnia growth. The overall patterns are consistent with our prediction that shifts in the environmental light : nutrient ratio as a result of glacial melting would contribute to shifts in the dominance of stoichiometrically contrasting taxa in consumer guilds.     

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.     

Threshold elemental ratios for carbon versus phosphorus limitation in Daphnia

1. The transition from carbon (C) to phosphorus (P) limited growth in Daphnia depends not only on the C : P ratio in seston, i.e. food quality, but also on food quantity. Carbon is commonly believed to be limiting at low food because of the energetic demands of basal metabolism. The critical C : P ratio in seston (otherwise known as the threshold elemental ratio, TER) above which P is limiting would then be high when food is scarce. 2. A new model that differentiates between the C : P requirements for growth and maintenance is presented that includes terms for both C and P in basal metabolism. At low food the calculated TERs for Daphnia of around 230 are only slightly higher than values of 200 or so at high intake. Seston C : P often exceeds 230, particularly in oligotrophic lakes where phytoplankton concentration is low and detritus dominates the diet, indicating the potential for limitation by P. 3. The analysis highlights the importance of P, as well as C, in maintenance metabolism and the overall metabolic budget, such that food quality is of importance even when intake is low. Further measurements of C and P metabolism at low food, in particular basal respiration and excretion rates, are needed in order to improve our understanding of the interacting roles of food quantity and quality in zooplankton nutrition.     

Trophic transfer and trophic modification of fatty acids in high Arctic lakes

1. A comparative study of fatty acid (FA) profiles in particulate matter (seston) and the key grazer Daphnia was performed in six high Arctic ponds (79°N, Svalbard). The ponds were all small and shallow, but followed a strong gradient with respect to nutrient content and optical properties. 2. A distinct locality‐specific pattern was detected by principal component analysis of FA profiles, where samples from each locality clustered both with regard to seston and Daphnia. Linear discriminant analysis using nine sestonic fatty acids as discriminant functions gave on average a correct prediction of the Daphnia lake membership in 47% of cases, with very high predictability in some lakes but poor predictability in others. 3. No significant correlation was detected between FA and nutrient concentration or levels of dissolved organic carbon. The major determinant of FA profiles as judged from a redundancy analysis was the taxonomic composition of phytoplankton communities, notably the biomass of Chlorophyceae. 4. The FA profiles of Daphnia were for some FAs strongly enriched relative to the seston, while diluted for others. Among the polyunsaturated fatty acids (PUFAs), a pronounced magnification of eicosapentaenoic acid (EPA, 20 : 5 n‐3), and to some extent 18 : 3 n‐3 and 20 : 4 n‐6 was found, while docosahexaenoic acid (DHA, 22 : 6 n‐3) contributed in general less to FAs in Daphnia than in seston and was hardly detectable in Daphnia from most localities. 5. The overall content of PUFAs in Daphnia was consistently high, close to 40% of total FA in all investigated localities, despite major differences in seston PUFA content. Daphnia thus acts as a regulator with regard to overall PUFAs, reflecting its physiological constraints and relatively fixed demands for PUFAs in general. The distinct locality‐specific profiles in Daphnia strongly suggest a kind of FA‐fingerprint, but our data do not allow strict statements on the use of specific FAs as trophic markers.     

Differential Impacts of Copepods and Cladocerans on Lake Seston,and Resulting Effects on Zooplankton Growth

In an enclosure study in Schöhsee, a small mesotrophic lake in Northern Germany, the impact of copepods and daphniids on the seston community was studied. In general, these two guilds differ in their feeding behaviour. Copepods actively select their food, with a preference for larger particles, whereas most cladocerans are unselective filter-feeders. In this study we investigate how the impact of the two different grazers affects zooplankton growth. We combine results obtained in the laboratory with results measured in situ in the enclosures. Copepods and cladocerans were cultured on seston from enclosures that were inhabited by density gradients of copepods or daphniids. We observed that Daphnia grew faster on seston that was pre-handled by copepods than on seston that was pre-handled by daphniids, and that somatic growth decreased with increasing densities of daphniids in the enclosures. In contrast, we observed no differences in development rates for copepods grown on the different media. The population growth rates of Daphnia in the Daphnia treatments were determined in the enclosures. Growth differences in both somatic- and population growth of Daphnia were correlated to food quality aspects of the seston. In the laboratory we found that Daphnia growth was correlated with several fatty acids. The strongest regression was with the concentration of 20:43 (r ²= 0.37). This particular fatty acid also showed the highest correlation with growth after normalisation of the fatty acids to the carbon content of the enclosures (r ²= 0.33). On the other hand, in the enclosure the population growth correlated most to the particulate nitrogen content (r ²= 0.78) and only to the N:C ratio, when normalised to carbon (r ²= 0.51).     

Field evidence for stoichiometric relationships between zooplankton and N and P availability in a shallow calcareous lake

1. According to stoichiometric theory, zooplankters have a species‐specific elemental composition. Daphniids have a relatively high phosphorus concentration in their tissues and copepods high nitrogen. Daphniids should, therefore, be more sensitive to phosphorus limitation and copepods more sensitive to nitrogen. A 2‐year study of a shallow marl lake in the west of Ireland investigated whether population fluctuations of the two dominant taxa, Daphnia spp. and the calanoid Eudiaptomus gracilis, were associated with the availability of phosphorus and nitrogen. 2. In accordance with stoichiometric predictions, Daphnia and Eudiaptomus reproduction had contrasting relationships with dietary phosphorus and nitrogen availability. Egg production by Daphnia was negatively associated with the ratio of dissolved inorganic nitrogen (DIN) : total phosphorus (TP) and the ratio of light to TP which was used as an indirect index for seston carbon (C) : phosphorus (P). Conversely calanoid egg production had a positive relationship with the DIN : TP ratio and was unrelated to the estimated C : P (light : TP) ratio. 3. Daphnia biomass was not, however, correlated with phosphorus availability, and neither was calanoid biomass correlated with nitrogen. The high ratio of DIN : TP when Daphnia dominated the zooplankton biomass and the low ratio when calanoids dominated, is consistent with Daphnia acting as a sink for phosphorus and calanoids as a sink for nitrogen and suggests consumer‐driven nutrient recycling.     

Seasonal shifts in the diet of lake zooplankton revealed by phospholipid fatty acid analysis

1. We analysed the phospholipid fatty acid (PLFA) profiles of seston and of the dominant zooplankter, Daphnia longispina, through the open water period in a small, dystrophic lake to investigate seasonal variation in the diet of Daphnia. Phytoplankton, heterotrophic bacteria, green sulphur bacteria and methane‐oxidizing bacteria (MOB) were all present in the water column of the lake, and previous studies have indicated that vertically migrating Daphnia can exploit all these potential food sources. 2. For adult Daphnia, although there was some correspondence between the PLFA profile of Daphnia and the concurrent seston PLFA profile, strongest correlations were between the Daphnia PLFA profile and those of potential food sources determined 7 days earlier. This interval presumably reflects the time it takes for adult Daphnia to turn over their fatty acid pool. 3. A correlation between the concentration of polyunsaturated fatty acids (PUFAs) in the epi‐ and metalimnion and measured primary production indicated that, within the total PLFA fraction, PUFAs can be useful biomarkers for phytoplankton in food‐web studies. Algal PUFAs contributed appreciably to total PLFAs in adult Daphnia during spring and summer, but less so in autumn. 4. Daphnia in the lake actually reached their highest biomass in autumn, when methanotrophic activity was also highest, and the highest magnitude of MOB‐specific PLFAs was recorded in both adult and juvenile Daphnia. A strong relationship existed between δ¹³C values of Daphnia reported previously and the proportion of MOB‐specific PLFAs in Daphnia. Autumnal mixing evidently stimulates bacterial oxidation of methane from the hypolimnion, and exploitation of the methanotrophic bacteria sustains a high Daphnia population late in the season. 5. Our results show that the PLFA composition of freshwater zooplankton like Daphnia corresponds rather well to that of their in situ diet of phytoplankton and bacteria, with a lag period of around 1 week in the case of adult animals. The PLFA profile of seston revealed the dominant available food sources, and relating these to the Daphnia PLFA profile provided insights into seasonal changes in Daphnia diet.     

Growth Responses of Tropical Cladocerans to Seston from Lake Monte Alegre (Brazil) Supplemented with Phosphorus, Fatty Acids, a Green Algae and a Cyanobacterium

The cladocerans Ceriodaphnia richardi, Daphnia ambigua, D. gessneri and Moina micrura were used to access food quality of Lake Monte Alegre’s seston. Experiments were carried out in summer and autumn as growth assays with lake seston only (control) and seston supplemented with phosphate, fatty acids or Synechococcus, and Scenedesmus. In summer, high C:P ratios in seston suggested strong phosphorus limitation, however, contrary to the expectations of stoichiometric theory, the addition of phosphate to seston did not improve cladoceran growth. Addition of PUFA increased growth rates and clutch size of D. gessneri, suggesting a possible deficiency in essential fatty acids in summer. Addition of Scenedesmus increased significantly growth rates of the cladocerans D. gessneri and C. cornuta, suggesting energy limitation in summer. In autumn, C:P ratios were lower than in summer, but still above the threshold ratio for Daphnia. At this time, addition of phosphate increased significantly growth rates of Daphnia suggesting strong P limitation, especially in D. gessneri. However, energy limitation was still important in autumn, as suggested by a further increase in growth rates in +Syn and +Sce treatments. Energy limitation was especially strong for Moina micrura, which is a fast-growing species, with high P content. Algal digestion resistance is a plausible hypothesis for energy limitation, since carbon concentrations in both seasons were above incipient limiting levels. These results show that the seston C:P ratio was not a consistent predictor of cladoceran P limitation and that factors other than P and energy limitation seem to be also important, such as PUFA or other biochemical factors. An erratum to this article is available at .     

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.     

<Emphasis Type="Italic">Daphnia</Emphasis> distribution in Andean Patagonian lakes: effect of low food quality and fish predation

Food quality in terms of carbon (C):phosphorus (P) ratio can constrain the success of highly demanding P herbivores as Daphnia. North Andean Patagonian lakes are ultraoligotrophic with low nutrient concentrations and well-developed euphotic zones. We investigated the distribution of the large Daphnia commutata in relation with food quality (sestonic C:P ratio) and predation risk in these lakes. The predation risk was estimated based on the fish species present and their relative eye diameter and transparency of the lake. The C:P ratios in the lakes were high, varying from 350 to >1,200. The lakes with D. commutata had significantly lower C:P ratio than those without these daphnids. On the other hand, those lakes where Daphnia is present have the lower predation risk than those were Daphnia is absent. In addition, we carried out growth experiments with neonates and natural seston of three lakes with different C:P ratio. The growth rates were inversely related with C:P of the food. Food quality and predation risk together determined the success or failure of large Daphnia populations in these Andean clear ultraoligotrophic lakes.     

Food quantity and quality regulation of trophic transfer between primary producers and a keystone grazer (Daphnia) in pelagic freshwater food webs

The transfer of energy and nutrients from plants to animals is a key process in all ecosystems. In lakes, inefficient transfer of primary producer derived energy can result in low animal growth rates, accumulation of nuisance phytoplankton blooms and dissipation of energy from the ecosystem. Most research on carbon transfer efficiency in pelagic food webs has focused on either food quantity or food quality, with the latter considered separately as either elemental stoichiometry or biochemical composition. The natural occurrence and magnitude of these types of growth limitations and their combined effects on Daphnia , a keystone grazer in pelagic freshwater ecosystems, are largely unknown. Our empirical models predict that the strength and nature of food quantity and quality limitation varies greatly with lake trophic state (total phosphorus, TP) and that Daphnia growth rates and thus energy and nutrient transfer efficiency are highest in lakes with intermediate trophic status (TP 10–25 μg l⁻¹). We predict that food availability place the greatest constraint on Daphnia growth in nutrient poor lakes (TP≤4 μg l⁻¹). Phosphorus limitation of Daphnia growth increased with decreasing TP, but the overall effect was never predicted to be the dominant constraining factor. Eicosapentaenoic acid (EPA, 20:5ω3) limitation was predicted to occur in both nutrient poor and nutrient rich lakes and placed the primary constraint on food quality in the most productive lakes. Two contrasting EPA-models gave different results on the magnitude of EPA-limitation, implying that additional food quality factors decrease Daphnia growth at high TP. In conclusion, the model predicts that Daphnia growth should peak in mesotrophic lakes, food quantity will place the greatest constraint on growth in oligotrophic lakes and EPA will primarily limit growth in eutrophic lakes.     

Resilience to changes in lake trophic state: Nutrient allocation into Daphnia resting eggs

During past decades, many lakes underwent drastic human‐caused changes in trophic state with strong implications for population dynamics and food web processes. We investigated the influence of trophic state on nutrient allocation into Daphnia resting eggs. The production of resting eggs is an important survival strategy, allowing Daphnia to cope with unfavorable environmental conditions. Allocation of essential nutrients into resting eggs may crucially influence embryonic development and offspring survival and thus is of great ecological and evolutionary interest. The capacity of Daphnia to adjust the allocation of nutrients into resting eggs may depend on the dietary nutrient supply, which may vary with trophic state‐related changes in the phytoplankton community composition. Resting eggs were isolated from sediment cores taken from Lake Constance, a large prealpine lake with a distinct eutrophication and reoligotrophication history, and analyzed for elemental (carbon, nitrogen, and phosphorus) and biochemical (sterols and fatty acids) nutrients. Carbon allocation into Daphnia resting eggs continuously decreased over time, irrespective of changes in trophic state. The allocation of nitrogen into Daphnia resting eggs followed the changes in trophic state, that is, nitrogen concentrations in resting eggs increased with eutrophication and decreased again with reoligotrophication. The allocation of phosphorus, sterols and long‐chain polyunsaturated fatty acids, such as eicosapentaenoic acid, into Daphnia resting eggs did not change significantly over time. Changes in trophic state strikingly influenced all trophic levels in Lake Constance. However, nutrient allocation into Daphnia resting eggs was mostly resilient to changes in lake trophic state.     

Relative importance of competition with Daphnia (Cladocera) and nutrient limitation on Anuraeopsis (Rotifera) population dynamics in a laboratory study

• 1 In the laboratory, the growth and reproduction of Anuraeopsis fissa were measured when fed on Scenedesmus species grown in nutrient‐sufficient, nitrogen‐limited and phosphorus‐limited media and in the presence or absence of one adult Daphnia longispina per vial.
• 2 Poor food quality may reduce the effect of competition on rotifers. Competition from Daphnia was stronger with nutrient sufficient algae than with nutrient limited algae. P‐limitation significantly reduced Anuraeopsis population growth rate and fecundity. The effect of nutrient limitation on Anuraeopsis was stronger than that of competition with Daphnia. The Anuraeopsis population declined with P‐limited food in both the presence and absence of Daphnia.
• 3 Exploitative competition by Daphnia on Anuraeopsis was stronger in the nutrient‐sufficient treatment than in the N‐limited one. Density, fecundity and population growth rate of Anuraeopsis were negatively affected by Daphnia in the nutrient‐sufficient treatment, while only fecundity was reduced by Daphnia in the N‐limited treatment. Consequently, in the N‐limited treatment, mortality should be lower in the presence of Daphnia. This result could suggest that Anuraeopsis lives longer when short of nitrogen.
• 4 Nutrient limitation may affect to the competitive interactions between zooplankton species. P‐limitation decreased the quality of algae as food for Anuraeopsis while N‐limitation decreased the susceptibility of this rotifer species to exploitative competition by Daphnia.

     

Potassium enrichment stimulates the growth and reproduction of a clone of Daphnia dentifera

Nutrient limitation commonly constrains organisms in natural ecosystems. Typically, ecologists focus on limitation by N and P. However, other nutrients can limit growth or reproduction. Here we focus on K limitation of invertebrate consumers (Daphnia dentifera) and phytoplankton in freshwater lakes. All organisms require K for several metabolic processes. In freshwater, K could limit growth because low external concentrations can increase the energetic costs of accumulating K. Furthermore, in a study linking K to disease, we previously found that K enrichment of water from one low-K lake stimulated the growth and reproduction of Daphnia. Here we test whether K could limit the production of Daphnia and phytoplankton across lakes and years. We repeated a life table experiment using water collected from a low-K lake during a different year. K again stimulated Daphnia reproduction. We also enriched water from 12 lakes with K or P and measured short-term growth of Daphnia and the resident algal community. Both nutrients increased Daphnia growth in five lakes. However, only P enhanced algal production. P stimulation of Daphnia positively correlated with algal quantity and the ratio of C to P in seston. However, K stimulation of Daphnia was not correlated with these factors or the background concentration of K. Thus, this study shows repeatable K-limited animal physiology in nature. Further, we can exclude the hypothesis that K stimulates Daphnia indirectly by enhancing algal production. These patterns call for future physiological studies to uncover the mechanistic basis of K limitation in natural systems.     

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.

     

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.     

Responses of algae, bacteria, Daphnia and natural parasite fauna of Daphnia to nutrient enrichment in mesocosms

Understanding responses of parasites to changes in nutrient regimes is necessary for prediction of their role in aquatic ecosystems under global change in nutrient loading. We studied the response of the natural parasite fauna of Daphnia longispina to nutrient enrichment in mesocosms in a small humic lake. We measured the concentrations of inorganic phosphorus and nitrogen in the water, total nutrients in the seston, algal and bacterial biomass, Daphnia population dynamics, Daphnia stoichiometry, Daphnia stable isotope values and the presence and abundance of parasites in treated mesocosms as compared to three control ones. Incorporation of the nutrient enrichment in the food web was seen as increased nutrient concentrations in the epilimnion and as a decrease in carbon:nutrient ratios and δ¹⁵N values in Daphnia. Nutrient enrichment did not significantly influence algal, bacterial or Daphnia biomass. One of the four parasite species observed, unidentified small gut parasite, had a higher prevalence (percentage of Daphnia infected) in treated mesocosms, but its intensity (number of parasites per infected host) remained the same among treatments. Our results suggest that the effect of nutrient enrichment on host–parasite dynamics is dependent on complex interactions within food webs and on the epidemiological traits of parasites.