Effects of nutrients and grazers on periphyton phosphorus in lake enclosures

SUMMARY. 1. Periphyton. measured as particulate phosphorus (PP) and expressed as periphyton PP, growing on vertically oriented substrata (polyvinyl impregnated nylon) under different nutrient loadings, light intensities (exposures), and grazer communities was examined in eight large enclosures (750 m3) where nutrients (N and P) and planktivorous fish (1+yellow perch) were added in a 2x2 factorial design.
2. During the first 3 weeks of the experiment (25 June to 15 July), there was a significantly higher accumulation of phosphorus into periphyton (periphyton PP) with fertilization, but fish addition had no effect. During the fourth to seventh weeks (16 July to 12 August), addition of fish was associated with lower abundance of amphipods and chironomids and higher concentration of periphyton PP. In the enclosures without fish, these invertebrates were over 25 times more abundant, and periphyton PP decreased substantially compared to the June-July period. Fertilization increased periphyton PP only at high exposures in the enclosures with fish.
3. Exposure had a significant effect on periphyton PP. In the enclosures with fish, high abundance of nanoplankton reduced water transparency, and periphyton PP was lower in the deeper waters which may have been due to limitation by low light. Lower periphyton PP was also observed at the surface on sunny sides of enclosures without fish, and therefore with high water transparency. This pattern may have been due to inhibitory effects of high light intensity.
4. Periphyton communities in the enclosures with fish had higher uptake rates for planktonic phosphorus, and lower rates of phosphorus release, suggesting that periphyton with high phosphorus demand may have high internal cycling of assimilated phosphorus.     

Temporal variation in body composition (C : N) helps explain seasonal patterns of zooplankton δ13C

1. The stable carbon isotope ratio δ¹³C is a useful tracer of energy flow in lake food webs, and the zooplankton signature is commonly used to establish a baseline for the pelagic habitat. However, sources of temporal variability in the δ¹³C of different zooplankton taxa are rarely considered. 2. Here, we investigate to what extent temporal variation in the δ¹³C of particulate organic matter (POM) (<41 μm) and the C : N of zooplankton can explain the temporal variability in δ¹³C of freshwater zooplankton. We compare temporal patterns of δ¹³C and C : N for Daphnia, Hesperodiaptomus franciscanus and Leptodiaptomus tyrelli over a 6‐month period at four sites in two oligotrophic lakes. 3. In all three taxa, seasonal variation in zooplankton C : N explained more of the variation in zooplankton δ¹³C than did the δ¹³C of POM. This suggests that variation in the lipid content of zooplankton can strongly influence temporal variation of δ¹³C in zooplankton. 4. Using these data, we evaluate procedures that estimate the δ¹³C of only the non‐lipid component of zooplankton. If zooplankton lipids are primarily dietary in origin, than extracting lipids or ‘normalising’δ¹³C based on C : N will exclude a major dietary source, and therefore may be inappropriate. 5. We conclude that temporal variation in body composition (C : N) of zooplankton can significantly influence the temporal variation of zooplankton δ¹³C signatures.     

Carbon pathways to zooplankton: insights from the combined use of stable isotope and fatty acid biomarkers

1. Numerous studies have quantified the relative contribution of terrestrial‐ and phytoplankton‐derived carbon sources to zooplankton secondary production in lakes. However, few investigated the pathways along which allochthonous and autochthonous carbon (C) was actually conveyed to consumers. 2. We suggest that the combined use of fatty acid and stable isotope biomarkers could solve this issue. We conducted a field study on two oligotrophic lakes, in which primary production increased significantly between 2002 and 2004. We used modelling to estimate the contribution of terrestrial‐ and phytoplankton‐derived C to particulate organic C (POC) and zooplankton production from their δ¹³C values in 2002 and 2004. 3. According to the isotope model, phytoplankton‐derived C accounted for a major part of the POC pool in both lakes and supported more Daphnia sp. production in 2004 than in 2002. Fatty acid data revealed that increased contribution of algal‐C to Daphnia production, although common between both lakes, was achieved through C pathways that were different. In one lake, Daphnia grazed more intensively on phytoplankton, whereas in the other there was greater grazing on bacteria. In the latter case, the increased primary production resulted in greater release of algal‐derived dissolved organic C (DOC), which may have supported extra bacterial and eventually Daphnia, production. 4. This is the first study illustrating that the combination of fatty acid and stable isotope biomarkers could further our understanding of the factors controlling the relative magnitude of food webs pathways conveying organic matter to zooplankton.     

Detecting trophic‐level variation in consumer assemblages

1. Stable isotopes of nitrogen are useful for quantifying the trophic structure of food webs, but only if the variation in trophic enrichment (Δ_N), which is the difference in δ¹⁵N between a consumer and its food, is small relative to the value of Δ_N itself. 2. We examined the sources of variation in zooplankton Δ_N by measuring the trophic enrichment (Δ_N) of seven species of freshwater cladocerans, and by testing for an effect of age and temperature on the Δ_N of Daphnia pulicaria. 3. We found that Δ_N was similar among Cladocera and was not correlated with body size. Overall, the Δ_N for D. pulicaria was 1.4‰ (SE = 0.69, n = 57), as was expected for the detritus diet that we used in our experiments. We found no effect of temperature (15–25 °C) on Δ_N, but found that Δ_N of D. pulicaria increased with increasing age (10–30 days). 4. We developed a new method to test for trophic‐level variation in a group of consumers that explicitly accounts for the uncertainty in Δ_N. Using this approach, we confirmed that natural assemblages of zooplankton feed at several trophic levels in lake food webs.     

Planktonic phosphorus pool sizes and cycling efficiency in coastal and interior British Columbia lakes

1. Limnologists have long acknowledged the importance of phosphorus (P) in determining the organism biomass and productivity of lake ecosystems. Despite a relatively large number of studies that have examined P cycling in lake ecosystems, there remain several substantial methodological issues that have impeded our understanding of P cycling in limnetic plankton communities. Two critical issues confronting ecologists are (1) a lack of precise measurements of the dissolved inorganic phosphorus (PO) and (2) accurate or complete measurements of dissolved P regeneration rates by plankton communities. 2. Here, we examine patterns of epilimnetic planktonic P pool sizes and turnover rates in eight lakes in British Columbia, Canada over a 2‐year period. We determine the concentrations and turnover times of P in various planktonic compartments (dissolved and various planktonic size fractions), using recently developed methods for estimating phosphate concentration and planktonic regeneration rates. 3. The pico‐ and nanoplankton size fraction (0.2–20 μm) played a central role in planktonic P cycling in lakes examined by this study. On average across lakes, pico‐ and nanoplankton contained >60% of the planktonic P, accounted for >90% PO uptake, and contributed 50% of the plankton community dissolved P regeneration rate. 4. PO concentrations determined by steady state bioassays (ssPO) were extremely low (87–611 pmol L⁻¹) and were 2–3 orders of magnitude less than simultaneously measured colorimetric soluble reactive phosphorus estimates. Lake ssPO concentrations increased linearly with total phosphorus (TP), and the slope of this relationship was approximately 1, indicating that PO remained a consistent proportion of the TP pool across a range of TP concentrations. 5. Turnover rates of the total planktonic P pool and the <20 μm pool became more rapid with increasing lake TP, indicating that, according to this metric, planktonic P cycling efficiency increased with TP concentrations. We also detected a significant relationship between particulate phosphorus (PP) <20 μm turnover time and seston N : P ratios, with PP <20 μm turnover times becoming slower with increasing seston N : P. These findings suggest that long‐standing conceptual models of nutrient cycling that predict slower cycling rates and decreasing cycling efficiency with increasing TP concentrations require further empirical examination. We postulate that patterns in lake P turnover and cycling efficiency are a result of complex interactions between plankton biomass and composition, and the ratios of multiple nutrients (C, N, P), rather than solely a function of the TP pool.