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.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

淡水中型浮游动物群体与悬浮物碳氮磷比耦联研究

通过对湖北省13个湖库悬浮物和中型浮游动物群体碳氮磷比的分析, 探讨了浮游动物与食物之间的元素耦联和不平衡性是如何随食物质量的改变而变化, 以及这种不平衡性对生态系统浮游动植物间能量传递效率的影响。结果显示: 中型浮游动物C﹕N比和C﹕P比分别随悬浮物C﹕N比和C﹕P比的增加而变大。悬浮物-中型浮游动物之间的元素不平衡性不仅与悬浮物C﹕N比显著正相关(r=0.97, P<0.001), 且与悬浮物C﹕P比也显著正相关(r=0.81, P=0.001)。浮游动植物之间的能量传递效率与悬浮物-中型浮游动物C﹕P比差值呈显著负相关(r= –0.58, P=0.037), 但与其C﹕N比差值关系不显著(P>0.05)。此外, 中型浮游动物生物量也随其与悬浮物之间C﹕P比差值的增大而下降(r= –0.59, P=0.033)。这些结果表明, 悬浮物碳氮磷比能显著影响中型浮游动物群体的碳氮磷比组成, 两者之间的元素不平衡性随前者的增加而变大并进而降低能量向后者的传递。     

Landscape controls on seston stoichiometry in urban stormwater management ponds

1. Stormwater management ponds (SWMPs) are taking the place of natural ponds and wetlands in urban areas. SWMPs have the potential to serve as hotspots for nutrient cycling, yet little is known about how urban catchments affect nutrient chemistry and stoichiometry within these ponds. 2. We sampled 50 SWMPs in Southern Ontario, Canada, to characterise their seston stoichiometry and make comparisons with published lake and pond data and models of seston stoichiometry. We tested (i) whether C : N : P ratios were similar to natural ponds and small lakes, (ii) whether seston stoichiometry was scale dependent and (iii) whether variability in seston chemistry could be explained by landscape and pond characteristics, such as catchment imperviousness and hydrological condition (based on recently received rainfall). 3. Seston C : N and C : P ratios were significantly lower in SWMPs than published ratios for small lakes, likely because of high nitrogen and phosphorus concentrations in SWMPs. Our results also showed no dependency of stoichiometric ratios on pond size. Analyses of ratios versus landscape and pond characteristics revealed significant relationships only when ponds were grouped based on the hydrological condition of the catchments. 4. It is likely that SWMPs function very differently during wet and dry periods. When SWMPs are disconnected from the landscape after a lengthy dry period, internal processes become increasingly important for seston stoichiometry.     

Linking major nutrients (C,H, N,and P) to trace metals (Fe,Mn, and Cu) in lake seston in southern Brazil

We investigated the relationships between major nutrients (C, H, N, and P) and trace metals (Cu, Fe, and Mn) in seston samples from ten lake/lagoon systems in southern Brazil. The systems were characterized by a diverse set of limnological features, including surface areas from 10⁻¹ to 10² km², water color, a _CDOM(440), from 1.4 to 12.9 m⁻¹, and electrical conductivity from 50 to 100 000 μS cm⁻¹. Seston concentrations also varied a great deal, 32-fold. The elemental (C: N, C: P, and N: P) and C: Chl-a ratios in the seston samples indicated, however, common features; i.e., most of the lakes were N-and/or P-limited, and the seston organic fraction was composed of nonvascular plants (e.g., phytoplankton). Our intersystem comparison revealed that the relative content of organic matter in seston and seston concentrations in lake water tended to correlate positively and negatively, respectively, with trace metal concentrations across the seston samples. Possible influences of elemental and C: Chl-a ratios on the association of metals with seston matrices, although theoretically important, were only partially evidenced here; positive correlations were found between C: N and also Org-H: N ratios with trace metal concentrations. We speculate that such results could be circumstantial, as the nature of the seston matrices appeared to be very similar among them. This hypothesis should thus be the theme of further research. In short, our findings suggest that C: N and Org-H: N ratios as well as the relative content of major nutrients in seston and seston concentrations can be importantly related to trace metal concentrations in seston samples. In discussing the results, however, we consider that metal-seston relationships depend on a variety of physical, chemical, and biological factors and/or variables other than those measured in this study, which could also contribute for defining and explaining variations in metal-seston concentrations in lake ecosystems.     

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.     

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.     

Stoichiometric relationships in vernal pond plankton communities

1. The light-nutrient hypothesis (LNH) predicts that changes in light supply can alter the balance of nutrient and energy limitation in primary producers. We tested this prediction by examining temporal changes in vernal forest ponds, which are highly dynamic systems with respect to seasonal change in light and nutrient supply. In three vernal ponds that differ in productivity, we measured changes in light, total and seston nitrogen and phosphorus, and seston carbon and chlorophyll during the spring, before and after tree leaf-out. We also quantified changes in the population dynamics of the major zooplankton grazers in these systems.
2. In each pond, nutrient levels increased and light levels declined, creating a temporal shift in light-nutrient supply to the plankton. Results generally supported predictions of stoichiometric theory and the LNH, but there were notable exceptions.
3. Seston C : N : P ratios rapidly changed in response to dramatic increases in N and P supply rates. However, seston N : P was typically lower than values for total N : P in the water. Furthermore, as predicted, we observed a decline in seston C : P as the light : nutrient ratio declined, but seston C : N simultaneously increased. These results suggest an unexpected shift towards potential nitrogen limitation. Alternatively, this change in nutrient ratios may be driven by a seasonal change in phytoplankton composition or nutritional mode.
4. Seston carbon concentrations remained stable despite seasonal changes in grazing intensity associated with the phenology of large-bodied Daphnia grazers. However, chlorophyll concentrations declined dramatically as the season progressed, resulting in a simultaneous decline in the C : Chlorophyll ratio of seston. Both pond shading and increased grazing probably contributed to the decline in chlorophyll.     

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     

Effects of increased temperature and nutrient enrichment on the stoichiometry of primary producers and consumers in temperate shallow lakes

1. We studied the effects of increased water temperatures (0–4.5 °C) and nutrient enrichment on the stoichiometric composition of different primary producers (macrophytes, epiphytes, seston and sediment biofilm) and invertebrate consumers in 24 mesocosm ecosystems created to mimic shallow pond environments. The nutrient ratios of primary producers were used as indicative of relative nitrogen (N) or phosphorus (P) limitation. We further used carbon stable isotopic composition (δ¹³C) of the different primary producers to elucidate differences in the degree of CO₂ limitation. 2. Epiphytes were the only primary producer with significantly higher δ¹³C in the enriched mesocosms. No temperature effects were observed in δ¹³C composition of any primary producer. Independently of the treatment effects, the four primary producers had different δ¹³C signatures indicative of differences in CO₂ limitation. Seston had signatures indicating negligible or low CO₂ limitation, followed by epiphytes and sediment biofilm, with moderate CO₂ limitation, while macrophytes showed the strongest CO₂ limitation. CO₂ together with biomass of epiphytes were the key variables explaining between 50 and 70% of the variability in δ¹³C of the different primary producers, suggesting that epiphytes play an important role in carbon flow of temperate shallow lakes. 3. The ratio of carbon to chlorophyll a decreased with increasing temperature and enrichment in both epiphytes and seston. The effects of temperature were mainly attributed to changes in algal Chl a content, while the decrease with enrichment was probably a result of a higher proportion of algae in the seston and epiphytes. 4. Macrophytes, epiphytes and seston decreased their C : N with enrichment, probably as an adaptation to the different N availability levels. The C : N of epiphytes and Elodea canadensis decreased with increasing temperature in the control mesocosms. Sediment biofilm was the only primary producer with lower C : P and N : P with enrichment, probably as a result of higher P accumulation in the sediment. 5. Independently of nutrient level and increased temperature effects the four primary producers had significantly different stoichiometric compositions. Macrophytes had higher C : N and C : P and, together with epiphytes, also the highest N : P. Seston had no N or P limitation, while macrophytes and epiphytes may have been P limited in a few mesocosms. Sediment biofilm indicated strong N deficiency. 6. Consumers had strongly homeostatic stoichiometric compositions in comparison to primary producers, with weak or no significant treatment effects in any of the groups (insects, leeches, molluscs and crustaceans). Among consumers, predators had significantly higher N content and lower C : N than grazers.     

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.

     

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.     

Variation in the C:P ratio of suspended and settling seston and its significance for P uptake calculations

SUMMARY. 1. In 1981–84 limnocorral (LC) experiments were performed in Lake Lucerne. Switzerland, to manipulate the planktonic community by varying P fertilization and by removing large zooplankton (with a 95 μm screen).
2. The C:P ratios in both suspended and entrapped seston exceeded the 'ideal' C:P ratio of 106 proposed by Redfield, Ketchum & Richards (1963) when P was limiting algal growth.
3. P fertilization could decrease the sestonic C:P ratio to 106 only when P did not limit algal growth; P additions far exceeding the P loading of eutrophic lakes were necessary to obtain this situation.
4. Changes in epilimnetic C:P ratios were usually related to short- term changes in primary production, caused by variable in situ light conditions and turbulence, and subsequent variation in POC concentrations.
5. Entrapped seston in the 95 μm-filtered LCs showed C:P ratios slightly higher than those of suspended seston, indicating fast P release and slower C mineralization in the epilimnetic nutrient cycle.
6. Removing large crustacean zooplankton enhanced epilimnetic P mineralization, and C:P ratios of entrapped seston in the 95 μm-filtered LCs were increased.
7 Detritus formed a relatively high proportion of the seston and amounted to more than two-thirds of the measured POC concentration.
8. Calculations of algal P uptake using information on sestonic C:P ratios and ¹⁴C uptake rates are questionable, as long as detritus cannot be separated from algae and net carbon uptake cannot be accurately measured.     

Carbon, nitrogen and phosphorus stoichiometry of crustacean zooplankton in the Baltic Sea: implications for nutrient recycling

The carbon (C), nitrogen (N) and phosphorus (P) contents (%of dry weight) of some crustacean zooplankton were studied inthe Baltic Sea. The copepod Acartia sp. had a stable C and Ncontent (48.3 ± 0.8% C, 12.4 ± 0.2% N, C:N ratio4.5 ± 0.1). The P content was variable (1–2%),probably depending on developmental stage and season. Copepodsaccumulating fat, like Pseudocalanus minutus elongatus, hadhigher and more variable C content (50–60%), and lowerN and P content (7–12% N, 0.6–1.5% P). The highestC and lowest N and P contents were found in adult Limnocalanusmacrurus. However, the N:P ratio was apparently independentof fat content and between 14 and 27 for all copepods. The cladoceransBosmina longispina maritima and Evadne nordmanni had lower Ncontent (9.3–10.8%) and higher C:N ratio (5.1–5.7)than Acartia sp. The P content (1.2–1.4%) was similarto Acartia sp. and the N:P ratios (16–19) were in thelower range of that found for the copepods. The N:P ratio wasgenerally somewhat higher in the copepods than in seston, whichmost of the year had nearly Redfield C:N:P ratios. Potentially,nutrient recycling from crustacean zooplankton could enhanceN limitation of phytoplankton, but small stoichiometric differencessuggest that this effect is probably weak. The extent is dependenton the structure of the zooplankton community and the grossgrowth efficiencies. Acartia copepodites, which had nearly RedfieldN:P ratios, would have the opposite effect and enhance P limitationin late summer when seston N:P ratios increased.     

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     

Seasonal occurrence of planktic Crustacea in sediment trap samples at three depth horizons in the Greenland Sea

Zooplankton were quantitatively investigated in sediment trap samples deployed in the Greenland Sea at three locations (72°00.7'N, 7°02.5'W, 72°23.0'N, 7°42.7'W and 72°22.9'N, 7°42.7'W). Overall samples covered a period of three annual cycles (July 1989 to July 1992). The investigation of zooplankton, consisting almost exclusively of Crustacea (> 99%), shows that the majority of the species must be regarded as “swimmers”, which actively invaded the traps. Crustacea were composed of 45 species, mainly Copepoda, Amphipoda and Ostracoda. Each depth level showed a characteristic species composition. Diversity decreased with depth. The samples were dominted by Copepoda in terms of abundance and by Amphipoda with respect to biomass. The seasonal occurrence of Crustacea showed a maximum during winter and a minimum during summer. Carnivores were regularly found in the collected material, indicating that they had been actively attracted by the organic matter emanating from the traps. Omnivores showed a distinct seasonal occurrence. During winter, when primary production is low, omnivores change to a predatory way of life and they were consequently also found in increasing numbers. Herbivores were always rare. Due to the high number of “swimmers” found in winter, when particle flux is at its minimum, particle flux measurements may be severly biased at this time of the year. Received: 24 January 1996/Accepted: 28 June 1996     

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.     

Ecophysiological responses to warming events by two sympatric zooplankton species

Under current climate change scenarios, numerous lakes are predictedto experience increases in maximum temperatures and in the timing,frequency and duration of heat waves. To investigate the effectsof the magnitude and timing of thermal stress on zooplanktoncommunities we compared the thermal responses and seasonal phenologiesof two sympatric species, Epischura lacustris and Daphnia catawba.Thermal responses were measured in the laboratory as differencesin demography of field animals at temperatures characteristicof temperate lakes in spring, summer and autumn (15–30°C).Epischura lacustris had a significantly lower temperature tolerancethan D. catawba, which appears to explain differences in theirseasonal phenologies. Divergence in thermal tolerances and seasonalphenologies of these sympatric species strongly suggests thatthe response of the zooplankton community in this lake to warmingevents will differ greatly with the timing and severity of thethermal stress. Similar effects are to be expected in otherlakes containing these species and other zooplankton communities.     

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.     

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.