Phytoplankton food quality control of planktonic food web processes

We developed a mechanistic model of nutrient, phytoplankton, zooplankton and fish interactions to test the effects of phytoplankton food quality for herbivorous zooplankton on planktonic food web processes. When phytoplankton food quality is high strong trophic cascades suppress phytoplankton biomass, the zooplankton can withstand intense zooplanktivory, and energy is efficiently transferred through the food web sustaining higher trophic level production. Low food quality results in trophic decoupling at the plant-animal interface, with phytoplankton biomass determined primarily by nutrient availability, zooplankton easily eliminated by fish predation, and poor energy transfer through the food web. At a given nutrient availability, food quality and zooplanktivory interact to determine zooplankton biomass which in turn determines algal biomass. High food quality resulted in intense zooplankton grazing which favored fast-growing phytoplankton taxa, whereas fish predation favored slow-growing phytoplankton. These results suggest algal food quality for herbivorous zooplankton can strongly influence the nature of aquatic food web dynamics, and can have profound effects on water quality and fisheries production. Handling editor: D. Hamilton     

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     

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     

武汉东湖颗粒悬浮物的结构与元素组成

本研究于1989-1990对武汉东湖营养水平不同的二个湖区的颗粒悬浮物的干物质结构和元素组成进行了分析。综合平均值表明,浮游动物的现存量约为浮游植物的1/4,浮游动物群落以小型的原生动物和轮虫占优势。从年平均值来看,浮游生物的干重占颗粒悬浮物干物质的2.5-7.6%,浮游生物碳量占颗粒悬浮物碳量的4.0-9.8%;颗粒悬浮物的碳/氮比与一般浮游植物的比值相似,但明显大于多数浮游动物;颗粒悬浮物的碳与干物重之比约为一般浮游生物的3/4;颗粒悬浮物的灰分含量约为45%,显着高于除硅藻以外的其它浮游生物。从数量上来看,有机碎屑是东湖生态系统颗粒悬浮物最重要的组成部分,而活体浮游生物只占颗粒悬浮物很小的一部分(<10%);这意味着在东湖来自以浮游植物为核心的食物网的有机碎屑的形成速率显着大于有机碎屑的矿化速率。       

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.

     

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.     

Nitrogen effects on the pelagic food web are modified by dissolved organic carbon

Global environmental change has altered the nitrogen (N) cycle and enhanced terrestrial dissolved organic carbon (DOC) loadings to northern boreal lakes. However, it is still unclear how enhanced N availability affects pelagic food web efficiency (FWE) and crustacean zooplankton growth in N limited boreal lakes. Here, we performed in situ mesocosm experiments in six unproductive boreal Swedish lakes, paired across a DOC gradient, with one lake in each pair fertilized with N (2011: reference year; 2012, 2013: impact years). We assessed how zooplankton growth and FWE were affected by changes in pelagic energy mobilization (PEM), food chain length (phytoplankton versus bacterial production based food chain, i.e. PP:BP), and food quality (seston stoichiometry) in response to N fertilization. Although PP, PEM and PP:BP increased in low and medium DOC lakes after N fertilization, consumer growth and FWE were reduced, especially at low DOC—potentially due to reduced phytoplankton food quality [increased C: phosphorus (P); N:P]. At high DOC, N fertilization caused modest increases in PP and PEM, with marginal changes in PP:BP and phytoplankton food quality, which, combined, led to a slight increase in zooplankton growth and FWE. Consequently, at low DOC (<12 mg L^?1), increased N availability lowers FWE due to mismatches in food quality demand and supply, whereas at high DOC this mismatch does not occur, and zooplankton production and FWE may increase. We conclude that the lake DOC level is critical for predicting the effects of enhanced inorganic N availability on pelagic productivity in boreal lakes.     

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

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

Spatial patterns of seston concentration and biochemical composition between nearshore and offshore waters of a Great Lake

1. The concentration and composition of seston were measured in the east basin of Lake Erie (U.S.A.–Canada) to determine whether the classical patterns of nearshore versus offshore zonation considered typical of large lakes have been altered by decades of change in nutrient loading and food web structure, and to test the possibility that food quality for seston consumers, as reflected in nutrient and lipid composition, may currently be diminished. Chlorophyll‐a (chl‐a), particulate organic carbon (POC), particulate P, total lipid (TL), lipid classes, fatty acids and major phytoplankton groups were compared between the nearshore (<20 m) and offshore zones in spring and early summer. 2. Particulate organic carbon, chl‐a and TL concentrations were lower nearshore than offshore, which is a reversal of the classical large lake pattern but consistent with other evidence from the Great Lakes that colonization by filter‐feeding dreissenid mussels can cause seston depletion in relatively shallow waters. 3. Terrestrial and bacterial fatty acid markers were slightly higher in the nearshore than the offshore, but there was little difference in nutrient, taxonomic or biochemical composition between zones otherwise. This shows a small differential influence of terrestrial and detrital inputs, and a slight shift away from phytoplankton dominance of seston composition in the nearshore, but little difference in potential food quality of seston. 4. Microscope counts, phytoplankton fluorescence, and fatty acid composition indicated succession from diatom dominance in May to dominance by cryptophyte and chromophyte flagellates in June and July. Depletion of dissolved P and Si, increased contribution of storage lipids, and changes in fatty acid composition suggested a mild degree of nutrient deficiency during this spring to summer transition. 5. Total lipids were dominated by classes typical of pigments and membranes with only moderate amounts of neutral (storage) lipids, polyunsaturated fatty acids (PUFA) were abundant (especially the long chain ω‐3 acids) and the POC : chl‐a and POC : particulate P ratios indicated P sufficiency or slight P‐deficiency. Despite concerns that some important consumers in the Great Lakes food web may be suffering from diminished food quality, these results point to a generally high seston food quality even with highly abundant mussels and incipient P deficiency in the east basin of Lake Erie.     

Modelling the role of highly unsaturated fatty acids in planktonic food web processes: Sensitivity analysis and examination of contemporary hypotheses

Aquatic food web models typically treat the constituent trophic levels as static elements interacting with one another and the environment. Dynamic biological stoichiometry has relaxed this assumption and considers evolutionary responses in said elements. The incorporation of organismal response in food web models holds promise for a more realistic portrayal of ecosystem dynamics. Recent advances in aquatic ecology pinpoint the importance of highly unsaturated fatty acids (HUFAs) on food web interactions and ecosystem resilience. In this study, we utilized a HUFA explicit submodel in conjunction with a limiting nutrient–phytoplankton–zooplankton–detritus (NPZD) mathematical system to incorporate elements of the physiology of individual animals into the context of plankton dynamics. Our HUFA-augmented plankton model provided a realistic platform to examine functional properties and physiological strategies that modulate resource procurement in different trophic environments and to effectively link variability at the organismal level with ecosystem-scale patterns. First, we were able to illustrate the implications of the filling-cup hypothesis, in which species’ fitness stems from dynamic HUFA turnover rates in response to bottom-up stresses. We then examined an evolutionary hypothesis of consumer fitness dependence on HUFA quota management strategies, whereby adaptive individuals with low HUFA minimum and optimum requirements gain competitive advantage. Several studies have reported higher HUFA concentrations in consumers than producers, and our results suggest that this pattern could be driven by a combination of conservative turnover and elevated bioconversion rates. Oligotrophic settings showed strong reliance upon exogenous phosphorus subsidies and frequently yielded inverted food web biomass distributions. With the prevalence of eutrophic conditions, consumer growth is primarily controlled by HUFA availability, and the associated biochemical limitation can ultimately result in patterns of algal accumulation. Finally, our study discusses directions to improve the representation of the producer–grazer interactions and thus advance our understanding of the factors that determine the flow of nutrients and energy to the higher trophic levels.     

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.

     

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。     

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.     

Grazing effects of a freshwater bivalve (Corbicula leana Prime) and large zooplankton on phytoplankton communities in two Korean lakes

This study examined the effects of a freshwater filter feeding bivalve (Corbicula leana Prime) and large zooplankton (>200 μm, mostly cladocerans and copepods) on the phytoplankton communities in two lakes with contrasting trophic conditions. A controlled experiment was conducted with four treatments (control, zooplankton addition, mussel addition, and both zooplankton and mussel addition), and each established in duplicate 10-l chambers. In both lakes there were significant effects of mussel grazing on phytoplankton density and biomass. The effects were greater in mesotrophic Lake Soyang than in hypertrophic Lake Ilgam. Effects of zooplankton grazing did not differ between these lakes, and zooplankton effects on phytoplankton were much less than the effects of mussels. Although mussels exerted a varying effect on phytoplankton according to their size, mussels reduced densities of almost all phytoplankton taxa. Total mean filtering rate (FR) of mussels in Lake Soyang was significantly greater than that in Lake Ilgam (p=0.002, n=5). Carbon fluxes from phytoplankton to mussels (977–2,379 μgC l^?1d^?1) and to zooplankton (76–264 μgC l^?1 d^?1) were always greater in Lake Ilgam due to the greater phytoplankton biomass (p<0.01, n=6). Based on the C-flux to biomass ratios, the mussels consumed 170–754% (avg. 412%) of phytoplankton standing stock in Lake Soyang, and 38–164% (avg. 106%) in Lake Ilgam per day. The C-flux to biomass ratio for mussels within each lake was much greater than for large zooplankton. Mussels reduced total phosphorus concentration by 5–34%, while increasing phosphate by 30–55% relative to the control. Total nitrogen also was reduced (by 9–25%), but there was no noticeable change in nitrate among treatments. The high consumption rate of phytoplankton by Corbicula leana even in a very eutrophic lake suggests that this mussel could affect planktonic and benthic food web structure and function by preferential feeding on small seston and by nutrient recycling. Control of mussel biomass therefore might be an effective tool for management of water quality in shallow eutrophic lakes and reservoirs in Korea.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Evolution of nutrient uptake reveals a trade-off in the ecological stoichiometry of plant-herbivore interactions

Nutrient limitation determines the primary production and species composition of many ecosystems. Here we apply an adaptive dynamics approach to investigate evolution of the ecological stoichiometry of primary producers and its implications for plant-herbivore interactions. The model predicts a trade-off between the competitive ability and grazing susceptibility of primary producers, driven by changes in their nutrient uptake rates. High nutrient uptake rates enhance the competitiveness of primary producers but also increase their nutritional quality for herbivores. This trade-off enables coexistence of nutrient exploiters and grazing avoiders. If herbivores are not selective, evolution favors runaway selection toward high nutrient uptake rates of the primary producers. However, if herbivores select nutritious food, the model predicts an evolutionarily stable strategy with lower nutrient uptake rates. When the model is parameterized for phytoplankton and zooplankton, the evolutionary dynamics result in plant-herbivore oscillations at ecological timescales, especially in environments with high nutrient availability and low selectivity of the herbivores. High herbivore selectivity stabilizes the community dynamics. These model predictions show that evolution permits nonequilibrium dynamics in plant-herbivore communities and shed new light on the evolutionary forces that shape the ecological stoichiometry of primary producers.     

Polyunsaturated fatty acids in zooplankton: variation due to taxonomy and trophic position

1. Food quality has major effects on the transfer of energy and matter in food webs, and essential long‐chained polyunsaturated fatty acids (PUFAs) can affect the quality of phytoplankton as food. In a study of oligotrophic lakes in north‐western Sweden, we investigated the fatty acid composition of four planktonic cladocerans and two calanoid copepods, representing herbivorous and carnivorous species. We also collected seston samples. 2. The proportions of long‐chain PUFAs in the organisms increased with their increasing trophic position. Thus, both their quality as food for other organisms, as well as their requirement for fatty acids (FAs), differed among taxa and depended on their trophic position. 3. We found taxon‐specific differences in the FA composition of zooplankton that were not related to sestonic FA composition. This implies that the variation in zooplankton FA composition is constrained by phylogenetic origin, life history characteristics, or both. 4. The cladoceran taxa contained 12–23% eicosapentaenoic acid (EPA) but only 0.9–2.1% docosahexaenoic acid (DHA) of the total FA content. In contrast, the calanoid copepods contained 7–11% EPA and 14–21% DHA. Thus, our results show that differences in the PUFA content among zooplankton species could have repercussions for both food web structure and function.     

Assimilation of Diazotrophic Nitrogen into Pelagic Food Webs

The fate of diazotrophic nitrogen (N_D) fixed by planktonic cyanobacteria in pelagic food webs remains unresolved, particularly for toxic cyanophytes that are selectively avoided by most herbivorous zooplankton. Current theory suggests that N_D fixed during cyanobacterial blooms can enter planktonic food webs contemporaneously with peak bloom biomass via direct grazing of zooplankton on cyanobacteria or via the uptake of bioavailable N_D (exuded from viable cyanobacterial cells) by palatable phytoplankton or microbial consortia. Alternatively, N_D can enter planktonic food webs post-bloom following the remineralization of bloom detritus. Although the relative contribution of these processes to planktonic nutrient cycles is unknown, we hypothesized that assimilation of bioavailable N_D (e.g., nitrate, ammonium) by palatable phytoplankton and subsequent grazing by zooplankton (either during or after the cyanobacterial bloom) would be the primary pathway by which N_D was incorporated into the planktonic food web. Instead, in situ stable isotope measurements and grazing experiments clearly documented that the assimilation of N_D by zooplankton outpaced assimilation by palatable phytoplankton during a bloom of toxic Nodularia spumigena Mertens. We identified two distinct temporal phases in the trophic transfer of N_D from N. spumigena to the plankton community. The first phase was a highly dynamic transfer of N_D to zooplankton with rates that covaried with bloom biomass while bypassing other phytoplankton taxa; a trophic transfer that we infer was routed through bloom-associated bacteria. The second phase was a slowly accelerating assimilation of the dissolved-N_D pool by phytoplankton that was decoupled from contemporaneous variability in N. spumigena concentrations. These findings provide empirical evidence that N_D can be assimilated and transferred rapidly throughout natural plankton communities and yield insights into the specific processes underlying the propagation of N_D through pelagic food webs.     

Phosphorus limitation and excess carbon in zooplankton

Organisms rely on a series of chemical reactions, which are constrained by the availability of key chemical elements, such as carbon (C), nitrogen (N), and phosphorus (P). Ecological stoichiometry provides a tool for analyzing how the balance of elements required by organisms affects food-web dynamics. Ecological stoichiometric theory suggests that the balance between supply and demand of elements is determined by the conversion efficiency from resources to organisms.Autotrophs and heterotrophs commonly face unequal access to and uptake of elements. The stoichiometric variability of autotrophs is based on their ability to maintain the balance of elements required for growth. This creates a challenge for their grazers. Phytoplankton can adjust their P content to ambient nutrient concentrations, while zooplankton cannot store excess nutrients. Ecological stoichiometric theory thus suggests that zooplankton have relatively fixed stoichiometry compared with phytoplankton.Nutrient limitation is common in aquatic systems. Stoichiometric imbalances between phytoplankton and zooplankton mean that zooplankton rarely find optimal food sources, and phytoplankton production is in excess. P availability potentially limits zooplankton growth, because of the high C:P ratio in phytoplankton relative to zooplankton demand. Based on the Liebig minimum principle, organisms are normally limited by a single nutrient, while everything else is in excess. Under P deficiency, excess C cannot be allocated to zooplankton somatic growth, and the net intake of C must balance the C:P ratio of zooplankton. Thus, when zooplankton encounter nutritionally imbalanced foods the elements in excess are released in order to maintain homeostasis. Excess C, released by zooplankton results in two biochemical challenges: (1) to sequester the limiting element and (2) to either store or dispose of the element in surplus.Zooplankton must resort to various physiological solutions to cope with these challenges. As a first option, zooplankton can reduce their C assimilation efficiency but maintain their P assimilation efficiency. Alternatively, after assimilation, excess C may be stored in C-rich compounds. Finally, assimilated excess C could also be disposed of through respiration or extracellular release. Excess C released by zooplankton reduces C transfer efficiency and sequestration in aquatic ecosystems.In aquatic ecosystems, C sequestration largely depends on the balance between uptake and demand for key nutrient elements. These feedback mechanisms have arisen only because organisms must obey stoichiometric rules at the cell and body levels, which greatly constrain the range of element values in ecosystems. Thus, the fate of C in ecosystems is determined by the absolute and relative demands for N and P of each organism. Limiting elements are utilized for growth and transferred in food chains with high efficiency, while non-limiting elements must be disposed of. Therefore, low C:P phytoplankton communities subject to high turnover rates and high productivity are selectively channeled into zooplankton. When zooplankton face high C:P foods, excess C is returned to the environment. Hence, nutrient-deficient phytoplankton constitute poor food, influencing the entire food web and adversely affecting secondary production at all levels.Excess C processed by zooplankton has far-reaching implications for ecosystem food-web functioning and C sequestration. Studies of the fate of excess C in zooplankton would increase the understanding of energy flow and material cycling in aquatic ecosystems. This paper reviews the reasons for P limitation and excess C in zooplankton, principal routes for the disposal of excess C, and the ecological effects of this. In addition, the paper aims to provide insight and a theoretical foundation for related studies in China.