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.

     

Variable Production by Different Pelagic Energy Mobilizers in Boreal Lakes

We studied production by three key pelagic energy mobilizer communities, phytoplankton (PP), heterotrophic bacteria (HB), and methanotrophic bacteria (MOB), in five boreal lakes of varying size and concentration of dissolved organic carbon (DOC). Production by PP was responsible for most (>55%) of the total pelagic energy mobilization in all five lakes. Production by HB and PP estimated for the whole water column during the ice-free period were positively correlated, but with the exception of the clearest and most eutrophic lake PP apparently could not support the total carbon demand of bacteria. However, the DOC concentration did not explain the variability of heterotrophic bacterial production (HBP) within or between the lakes. Thus, our results provide circumstantial evidence for the “priming effect” whereby labile organic matter from autochthonous production enhances decomposition of allochthonous DOC. However, HBP was only 10–23% of the total pelagic energy mobilization in the lakes, suggesting that only a minor fraction of allochthonous DOC became available for higher trophic levels. High MOB activity was detected in the water columns of the stratified lakes when the molar ratio of CH₄:O₂ varied between 0.5 and 12. In the small stratified lakes (area < 0.01 km²), MOB production contributed 13–52% of the total pelagic energy mobilization, being greatest during the autumn mixing period. Our results indicate that in small stratified lakes (area < 0.01 km²) bacteria, especially MOB, are potentially quantitatively important supplementary food resources for zooplankton. However, in larger lakes primary producers are the most important (>70%) potential food source for zooplankton.     

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.     

Browning affects pelagic productivity in northern lakes by surface water warming and carbon fertilization

Global change impacts important environmental drivers for pelagic gross primary production (GPP) in northern lakes, such as temperature, light, nutrient, and inorganic carbon availability. Separate and/or synergistic impacts of these environmental drivers on pelagic GPP remain largely unresolved. Here, we assess key drivers of pelagic GPP by combining detailed depth profiles of summer pelagic GPP with environmental and climatic data across 45 small and shallow lakes across northern Sweden (20 boreal, 6 subarctic, and 19 arctic lakes). We found that across lakes summer pelagic GPP was strongest associated with lake water temperatures, lake carbon dioxide (CO₂) concentrations impacted by lake water pH, and further moderated by dissolved organic carbon (DOC) concentrations influencing light and nutrient conditions. We further used this dataset to assess the extent of additional DOC-induced warming of epilimnia (here named internal warming), which was especially pronounced in shallow lakes (decreasing 0.96°C for every decreasing m in average lake depth) and increased with higher concentrations of DOC. Additionally, the total pools and relative proportion of dissolved inorganic carbon and DOC, further influenced pelagic GPP with drivers differing slightly among the boreal, subarctic and Arctic biomes. Our study provides novel insights in that global change affects pelagic GPP in northern lakes not only by modifying the organic carbon cycle and light and nutrient conditions, but also through modifications of inorganic carbon supply and temperature. Considering the large-scale impacts and similarities of global warming, browning and recovery from acidification of lakes at higher latitudes throughout the northern hemisphere, these changes are likely to operate on a global scale.     

Browning reduces the availability—but not the transfer—of essential fatty acids in temperate lakes

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Nutrient limitation of bacterioplankton and phytoplankton in humic lakes in northern Sweden

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L^–1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), respectively. A third lake was unfertilized and served as a control. In addition to this lake fertilization experiment, data from different regional surveys were used to assess the role of different limiting factors.
2. The P fertilization had no effects on bacterioplankton or phytoplankton, while phytoplankton were significantly stimulated by N fertilization. Inorganic nutrient limitation of bacterioplankton was a function of DOC concentration in water of the investigated region and nutrient-limited bacteria were found only in lakes with DOC concentrations less than around 15 mg L^–1
3. The fertilization experiments demonstrated that the DOC-rich experimental lakes contained a bioavailable pool of P that was not utilized to its full potential under natural conditions. The overall mobilization of energy (bacterioplankton plus phytoplankton) in the experimental lakes was restricted by lack of inorganic N.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Dissolved organic carbon in a humic lake: effects on bacterial production and respiration

Allochthonous matter was the main source of carbon for pelagic bacteria in a humic lake, accounting for almost 90% of the carbon required to support observed bacterial growth. The estimated contribution from zooplankton excretion was of the same magnitude as direct phytoplankton release, both accounting for 5–7% of bacterial demands for dissolved carbon. Bacteria were an important source of carbon both for heterotrophic phytoplankton and for filter feeding zooplankton species, further stressing the role of humus DOC in overall lake productivity. The high contribution of allochthonous DOC implies a stoichiometry of dissolved nutrients with a surplus of C relative to P. The high P cell quota of bacteria suggest that under such conditions they are P-limited and act like net consumers of P. Excess C will be disposed of, and bacterial respiration rate will increase following a transition from carbon-limited bacterial growth towards mineral-nutrient-limited growth. Thus the high community respiration and frequent CO₂-supersaturation in humic lakes may be caused not only by the absolute supply of organic C, but also by the stoichiometry of the dissolved nutrient pool.     

Primary production under hypertrophic conditions and its relationship with bacterial production

In shallow hypertrophic lakes where light availability restricts the growth of macrophytes and benthic phytoplankton, pelagic phytoplankton modulates importantly ecosystem production and the energy transfer to heterotrophic bacteria. Diel and seasonal variations in primary production (PP) were studied in the hypertrophic Albufera de Valencia (Spain). Additionally, the relationship between PP and heterotrophic bacterial production (BP) was assessed. PP was extremely high, exceeding most values reported for hypertrophic lakes to date. PP displayed marked diurnal variations defined by the solar radiation curve. Likewise, PP changed importantly across seasons. Minimum PP coincided with maximum water transparency and short water residence times in winter, whereas maximum PP was observed in late spring associated with high chlorophyll a. The spring PP maximum contrasted with the summer maximum often observed in hypertrophic lakes. When compared to spring PP values, summer PP values were lower as a result of strong nitrogen limitation. In contrast to PP, BP remained fairly constant across seasons. Nonetheless, there was a joint diminution during increased water transparency followed by an increase in early spring. Phytoplankton was always the most relevant input to particulate carbon production, but the BP/PP ratio showed clear seasonal variations. The BP/PP ratio was minimum in spring, low in summer and highest in winter. The extracellular dissolved organic carbon released by phytoplankton was sufficient to meet bacterial carbon demand in all experimental dates, suggesting that allochthonous carbon sources play a minor role in sustaining BP, though they cannot be excluded. However, we hypothesize that high availability of dissolved organic carbon might explain the lack of coupling observed between BP and PP.     

Algal picoplankton production and contribution to food-webs in oligotrophic British Columbia lakes

The pelagic communities of two contrasting oligotrophic lakes in British Columbia were studied to determine why an interior, dimictic lake (Quesnel) supports a greater biomass of zooplankton and produces larger planktivorous sockeye salmon (Oncorhynchus nerka) than a coastal warm-monomictic lake (Sproat). The ultra-oligotrophic status and differing planktivore densities in Sproat Lake increased the relative importance of algal picoplankton, diminished the abundance of large zooplankton, and increased the significance of rotifers and other small-bodied zooplankton. These picoplankton based food webs result in longer, indirect and less efficient pathways of carbon flow from phytoplankton to fish. In contrast, Quesnel Lake is a more productive oligotrophic lake and its pelagic food webs are based more on nanoplankton and small microphytoplankton that support larger-bodied zooplankton (Daphnia, Diaptomus), and a more direct and efficient two-step transfer to fish. The greater variability of the annual recruitment of sockeye fry in interior lakes may keep zooplankton communities in a non-steady state, this in turn may perpetuate the occurrence of quadrennial cyclic dominance in adult salmon returning to these systems.     

A Framework for Understanding Variation in Pelagic Gross Primary Production of Lake Ecosystems

Light and nutrient availability are key physiological constraints for primary production. Widespread environmental changes are causing variability in loads of terrestrial dissolved organic carbon (DOC) and nutrients from watersheds to lakes, contributing to simultaneous changes in both light and nutrient supply. Experimental evidence highlights the potential for these watershed loads to create complex and context-dependent responses of within-lake primary production; however, the field lacks a predictive model to investigate these responses. We embedded a well-established physiological model of phytoplankton growth within an ecosystem model of nutrient and DOC supply to assess how simultaneous changes in DOC and nutrient loads could impact pelagic primary production in lakes. The model generated a unimodal relationship between GPP and DOC concentration when loads of DOC and nutrients were tightly correlated across space or time. In this unimodal relationship, the magnitude of the peak GPP was primarily determined by the DOC-to-nutrient ratio of the load, and the location of the peak along the DOC axis was primarily determined by lake area. Greater nutrient supply relative to DOC load contributed to greater productivity, and larger lake area increased light limitation for primary producers at a given DOC concentration, owing to the positive relationship between lake area and epilimnion depth. When loads of DOC and nutrients were not tightly correlated in space or time, the model generated a wedge-shaped pattern between GPP and DOC, consistent with spatial surveys from a global set of lakes. Our model is thus capable of unifying the diversity of empirically observed spatial and temporal responses of lake productivity to DOC and mineral nutrient supply presented in the literature, and provides qualitative predictions for how lake pelagic primary productivity may respond to widespread environmental changes.     

Performance trade-off across a natural resource gradient

An important environmental factor determining both phytoplankton and zooplankton community composition is lake depth and thermal stratification. However, there is little information on how the interaction between zooplankton grazers and their phytoplankton food changes along an environmental gradient of lake depth. We contrasted resource availability for daphniid zooplankton populations living in two shallow, unstratified lakes and in two deep, stratified lakes using a novel growth bioassay. Stratified lakes had consistently lower resource richness than shallow unstratified lakes. To test whether resources were important in explaining differences in daphniid composition of shallow and deep lakes, we performed reciprocal transplant experiments. We raised daphniids typical of shallow (Ceriodaphnia reticulata) and deep (Daphnia dentifera) lakes in the resources from replicate shallow and deep lakes and monitored survival and reproduction. The two species exhibited a performance trade-off, measured by life table r and R ₀, across a gradient in natural resource richness. D. dentifera had higher relative fitness than C. reticulata when raised in the poorest resource environment from a deep lake. However, under richer resource conditions typical of shallow lakes, C. reticulata outperformed D. dentifera. We further created a gradient in natural resource quantity (by dilution) to test whether this trade-off in species relative fitness involved aspects of resource quality. No trade-off in species performance was evident across the dilution gradient, indicating that resource quality was important to the trade-off. We conclude that shifts in daphniid species composition along a gradient of lake depth involve an adaptive trade-off in ability to exploit rich versus poor resource quality. Received: 11 May 1998 / Accepted: 15 January 1999     

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.     

Effects of a Whole-lake,Experimental Fertilization on Lake Trout in a Small Oligotrophic Arctic Lake

We tested whether increased phosphorus and nitrogen concentrations would affect a lake trout (Salvelinus namaycush) population in a small oligotrophic lake with a benthically dominated food web. From 1990 to 1994, nitrogen and phosphorus were added to Lake N1 (4.4 ha) at the arctic Long-Term Ecological Research site in Alaska. We used mark/recapture methods to determine the lake trout population size, size structure, recruitment, and individual growth from 1987 to 1999. Data were also collected on water chemistry and food availability. Fertilization resulted in increased pelagic primary productivity, chlorophyll a, turbidity, snail density, and hypoxia in summer and winter. Lake trout density was not affected by the manipulation however growth and average size increased. Recruitment was high initially, but declined throughout the fertilization. These results suggest that lake trout were affected through increased food availability and changes to the physical characteristics of the lake. During fertilization, hypoxia near the sediments may have killed over-wintering embryos and decreased habitat availability. Although lake trout responded strongly to increased nutrients, loss of recruitment might jeopardize lake trout persistence if arctic lakes undergo eutrophication.     

Light and nutrient control phytoplankton biomass responses to global change in northern lakes

Global change affects terrestrial loadings of colored dissolved organic carbon (DOC) and nutrients to northern lakes. Still, little is known about how phytoplankton respond to changes in light and nutrient availability across gradients in lake DOC. In this study, we used results from whole‐lake studies in northern Sweden to show that annual mean phytoplankton biomass expressed unimodal curved relationships across lake DOC gradients, peaking at threshold DOC levels of around 11 mg/L. Whole‐lake single nutrient enrichment in selected lakes caused elevated biomass, with most pronounced effect at the threshold DOC level. These patterns give support to the suggested dual control by DOC on phytoplankton via nutrient (positively) and light (negatively) availability and imply that the lakes' location along the DOC axis is critical in determining to what extent phytoplankton respond to changes in DOC and/or nutrient loadings. By using data from the large Swedish Lake Monitoring Survey, we further estimated that 80% of northern Swedish lakes are below the DOC threshold, potentially experiencing increased phytoplankton biomass with browning alone, and/or combined with nutrient enrichment. The results support the previous model results on effects of browning and eutrophication on lake phytoplankton, and provide important understanding of how northern lakes may respond to future global changes.     

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     

Accounting for littoral primary production by periphyton shifts a highly humic boreal lake towards net autotrophy

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Climate-driven changes on phytoplankton–zooplankton coupling and nutrient availability in high mountain lakes of Southern Europe

1. The effect of climate variability on phytoplankton and zooplankton dynamics and nutrient availability was studied in two high mountain fishless lakes (La Caldera and Río Seco) of contrasting morphology, hydrology and dissolved inorganic nitrogen : soluble reactive phosphate (DIN : SRP) ratios during 1986 and after a 10‐year‐long drought in 1996 and 1997. 2. Thaw was delayed and water temperatures were lower in both lakes in 1996 than in 1986 and 1997. However, the lake‐specific DIN : SRP ratio was maintained in the 3 years studied, reflecting its local control. 3. On other hand, the presumptive limiting nutrient in each lake, P in La Caldera and N in Río Seco, showed higher concentrations in 1996 versus 1986 and 1997. Significant positive correlations between temperature and chlorophyll a were found in both lakes in 1996 but these relationships were negative or not significant in 1986 and 1997. Zooplankton biomass showed lower values in 1996 than in 1986 or 1997. 4. These findings can be explained by a decoupling of the phytoplankton–zooplankton interaction because of a constraint on zooplankton growth by low temperatures in the coldest year studied. This observation furnishes evidence that regional climatic control on the phytoplankton–zooplankton link can modulate the overall demand for nutrients.     

Effects of water-column mixing on bacteria,phytoplankton, and rotifers under different levels of herbivory in a shallow eutrophic lake

Water-column mixing is known to have a decisive impact on plankton communities. The underlying mechanisms depend on the size and depth of the water body, nutrient status and the plankton community structure, and they are well understood for shallow polymictic and deep stratified lakes. Two consecutive mixing events of similar intensity under different levels of herbivory were performed in enclosures in a shallow, but periodically stratified, eutrophic lake, in order to investigate the effects of water-column mixing on bacteria abundance, phytoplankton abundance and diversity, and rotifer abundance and fecundity. When herbivory by filter-feeding zooplankton was low, water-column mixing that provoked a substantial nutrient input into the euphotic zone led to a strong net increase of bacteria and phytoplankton biomass. Phytoplankton diversity was lower in the mixed enclosures than in the undisturbed ones because of the greater contribution of a few fast-growing species. After the second mixing event, at a high biomass of filter-feeding crustaceans, the increase of phytoplankton biomass was lower than after the first mixing, and diversity remained unchanged because enhanced growth of small fast-growing phytoplankton was prevented by zooplankton grazing. Bacterial abundance did not increase after the second mixing, when cladoceran biomass was high. Changes in rotifer fecundity indicated a transmission of the phytoplankton response to the next trophic level. Our results suggest that water-column mixing in shallow eutrophic lakes with periodic stratification has a strong effect on the plankton community via enhanced nutrient availability rather than resuspension or reduced light availability. This fuels the basis of the classic and microbial food chain via enhanced phytoplankton and bacterial growth, but the effects on biomass may be damped by high levels of herbivory. Received: 3 May 1999 / Accepted: 13 April 2000     

Links between Terrestrial Primary Production and Bacterial Production and Respiration in Lakes in a Climate Gradient in Subarctic Sweden

We compared terrestrial net primary production (NPP) and terrestrial export of dissolved organic carbon (DOC) with lake water heterotrophic bacterial activity in 12 headwater lake catchments along an altitude gradient in subarctic Sweden. Modelled NPP declined strongly with altitude and annual air temperature decreases along the altitude gradient (6°C between the warmest and the coldest catchment). Estimated terrestrial DOC export to the lakes was closely correlated to NPP. Heterotrophic bacterial production (BP) and respiration (BR) were mainly based on terrestrial organic carbon and strongly correlated with the terrestrial DOC export. Excess respiration over PP of the pelagic system was similar to net emission of CO₂ in the lakes. BR and CO₂ emission made up considerably higher shares of the terrestrial DOC input in warm lakes than in cold lakes, implying that respiration and the degree of net heterotrophy in the lakes were dependant not only on terrestrial export of DOC, but also on characteristics in the lakes which changed along the gradient and affected the bacterial metabolization of allochthonous DOC. The study showed close links between terrestrial primary production, terrestrial DOC export and bacterial activity in lakes and how these relationships were dependant on air temperature. Increases in air temperature in high latitude unproductive systems might have considerable consequences for lake water productivity and release of CO₂ to the atmosphere, which are ultimately determined by terrestrial primary production.