Preface

Recent studies on food quality for zooplankton have developed two new hypotheses, the importance of the dietary carbon to phosphorus (C : P) ratio and of certain ‘essential’ fatty acids. These new hypotheses, the controversies they have provoked and the general lack of field studies prompted us to organize a workshop. Thus, the first international Plankton Ecology Group (PEG) workshop on The Role of Food Quality for Zooplankton was held from 17 to 21 March 1996 at the Centre of Limnology in Nieuwersluis, the Netherlands. The main objectives were to take stock of the state-of-the-art in food quality research, to address the factors that determine food quality, and to integrate the available information into a coherent and consistent view of food quality for the zooplankton. In contrast to most PEG meetings, we restricted the meeting to about 35 participants. However, international participation was broader than usual and included colleagues from North America and Asia, as well as Europe. The workshop’s organizers, Ramesh Gulati, a senior scientist, and Paul M. M. Weers, a post-doc at the Centre Of Limnology, approached potential speakers, in some cases suggesting topics for presentation. Each of five sessions was introduced with a review paper or a modelling paper.This special volume of Freshwater Biology contains twenty-six papers, including a summary by R. Gulati & B. DeMott. This paper briefly describes the findings of the workshop, and discusses them in light of the literature, discussions at the workshop and written comments from participants. Each of the workshop papers was peer reviewed, usually by two but in some cases by three referees. We learned that fatty acids and phosphorus are not mutually exclusive alternatives; the content of polyunsaturated acids of certain algae, especially eicosapentaenoic acid (EPA), is markedly affected by the algal P content. Thus, elemental limitation and fatty acid limitation may have additive or interacting effects. Although further research might modify this view, the accumulating evidence for P limitation seems stronger than for fatty acids limitation. However, despite laboratory studies that have clarified the potential mechanisms of food quality, the paucity of field studies prevents a consensus on the importance of specific mechanisms in nature. We also learned much from aquaculture studies, which have developed techniques for enriching zooplankton food with fatty acids and essential nutrients. We believe that the workshop proceedings will both contribute to our understanding and stimulate further research on food quality for zooplankton. Lastly, we take cognizance that cyanobacterial toxins are an important area of food quality research, which for practical reasons, did not receive our attention. Several presentations, however, revealed that cyanobacteria are also potentially poor food due to both higher C:P ratios and lower fatty acid content than many other algal species.Paul Weers’ efforts in organizing the workshop were very helpful. We acknowledge the facilities, funds and encouragement provided by staff and administration of the Centre of Limnology. We are also grateful to the participants for their cooperation and to the reviewers, many of whom were workshop participants.RAMESH D. GULATI BILL DE MOTT     

Oligotrophication of a large,deep lake alters food quantity and quality constraints at the primary producer–consumer interface

To assess nutritional consequences associated with lake oligotrophication for aquatic consumers, we analyzed the elemental and biochemical composition of natural seston and concomitantly conducted laboratory growth experiments in which the freshwater key herbivore Daphnia was raised on natural seston of the nowadays (2008) oligotrophic Lake Constance throughout an annual cycle. Food quality mediated constraints on Daphnia performance were assessed by comparing somatic growth rates with seston characteristics (multiple regression analysis) and by manipulating the elemental and biochemical composition of natural seston experimentally (nutrient supplementation). Results were compared to similar experiments carried out previously (1997) during a mesotrophic phase of the lake. In the oligotrophic phase, particulate carbon and phosphorus concentrations were lower, fatty acid concentrations were higher, and the taxonomic composition of phytoplankton was less diverse, with a more diatom‐ and cryptophytes‐dominated community, compared to the previous mesotrophic phase. Multiple regression analysis indicated a shift from a simultaneous limitation by food quantity (in terms of carbon) and quality (i.e. α‐linolenic acid) during the mesotrophic phase to a complex multiple nutrient limitation mediated by food quantity, phosphorus, and omega‐3 fatty acids in the following oligotrophic phase. The concomitant supplementation experiments also revealed seasonal changes in multiple resource limitations, i.e. the prevalent limitation by food quantity was accompanied by a simultaneous limitation by either phosphorus or omega‐3 fatty acids, and thus confirmed and complemented the multiple regression approach. Our results indicate that seasonal and annual changes in nutrient availabilities can create complex co‐limitation scenarios consumers have to cope with, which consequently may also affect the efficiency of energy transfer in food webs.     

Highly unsaturated fatty acids in nature: what we know and what we need to learn

The supply and demand of omega‐3 highly unsaturated fatty acids (ω‐3 HUFA) in natural ecosystems may lead to resource limitation in a diverse array of animal taxa. Here, we review why food quality in terms of ω‐3 HUFAs is important, particularly for neural tissue, across a diversity of animal taxa ranging from invertebrate zooplankton to vertebrates (including humans). Our review is focused on ω‐3 HUFAs rather than other unsaturated fatty acids because these compounds are especially important biochemically, but scarce in nature. We discuss the dichotomy between ω‐3 HUFA availability between aquatic primary producers, which are often rich in these compounds, and terrestrial primary producers, which are contain little to none of them. We describe the use of fatty acids as qualitative and quantitative tracers for reconstructing animal diets in natural ecosystems. Next, we discuss both direct and indirect ecological implications of ω‐3 HUFA limitation at the individual, population, food web, and ecosystem scales, which include: changes in behavior, species composition, secondary production rates, trophic transfer efficiency and cross‐ecosystem subsidies. We finish by highlighting future research priorities including a need for more research on ω‐3 HUFAs in terrestrial systems, more research their importance for higher order consumers, and more research on the food web and ecosystem‐scale effects of ω‐3 HUFA limitation. Synthesis Mismatches between the supply of and demand for omega‐3 highly unsaturated fatty acids (ω‐3 HUFA) in natural ecosystems have the potential to result in resource limitation across a diverse array of ecosystems. We combined perspectives from ecology and nutritional science to develop a unified synthesis of ω‐3 HUFA ecology. We reviewed the importance of ω‐3 HUFAs for animals, the striking differences in ω‐3 HUFA availability at the base of terrestrial versus aquatic food webs, and the implications of ω‐3 HUFA limitation for food webs. We finished by highlighting research priorities in the field including more research on ω‐3 HUFAs in terrestrial systems, on higher order consumers, and at the food web and ecosystem‐scales.     

The role of food quality for zooplankton: remarks on the state-of-the-art, perspectives and priorities

1. This paper summarizes the salient features of the contributions to the workshop on The Role of Food Quality for Zooplankton. In this paper we attempt critically to evaluate our present knowledge in the light of new studies. 2. For the growth and reproduction of zooplankton, the existing literature considers two main limiting factors in the diet, i.e. phosphorus (homeostasis theory) and fatty acids. Nevertheless, interpretations and opinions regarding the importance of these two factors are the subject of controversy in the literature. No attempts have been made to link these two potentially limiting factors, let alone give a coherent view based on the mechanisms behind limitation. Aquaculture studies provide some direct evidence of the importance of the long-chained poly unsaturated fatty acids (PUFA) for zooplankton. The presence of PUFA in phytoplankton is reported to affect the growth rates of zooplankton significantly. 3. Field data on carbon and phosphorus indicate a greater constancy of the C : P ratios of zooplankton than of their food. Empirical data and modelling studies suggest that zooplankton, especially Daphnia spp., may maintain nutrient homeostasis by incorporating a greater proportion of the limiting nutrients ingested and releasing more of nutrients in excess supply. The need for conserving nutrients in short supply increases with the increase in growth rates. 4. Phosphorus certainly influences zooplankton food directly. Direct supplementation of the P-insufficient algal diet with PO₄-P alone discernibly improves the growth in daphnids. It is highly plausible that P limitation and fatty acid limitation are not mutually exclusive alternatives. The two, separately or in conjunction, can control growth of at least some lake zooplankters, especially daphnids. 5. Besides a shortage of nutrient (P), other environmental factors (irradiance, UV-radiation, temperature) can also adversely affect the zooplankton diet, including its digestibility and assimilation efficiency. 6. It is not yet clear if PUFA deficiency in the diet is in some way related to or caused by P deficiency. It is, however, now known that the EPA (eicosapentaenoic acid, 20 : 5ω3) content of certain algae is markedly reduced under P-limitation and that it differs significantly among the different taxonomic groups of phytoplankton. Diatoms and flagellates are generally considered as good-quality foods because of their high EPA content. On the contrary, cyanobacteria are low-quality food, having both low EPA and P content. 7. Recent experiments reveal that the relative importance of fatty acids for daphnids increases with a decreasing C : P ratio in the food, i.e. if P is no longer limiting, and vice versa. For daphnids, there is possibly a switch between P-limitation and PUFA limitation at intermediate C : P ratios. At higher C : P ratios, P is more important but at lower ratios PUFA are more crucial for growth and reproduction. 8. Lastly, the accumulating evidence for P limitation is stronger than that for fatty acid limitation.     

Interactions between P-limitation and different C conditions on the fatty acid composition of an extremophile microalga

The extremophilic microalga Chlamydomonas acidophila inhabits very acidic waters (pH 2-3.5), where its growth is often limited by phosphorus (P) or colimited by P and inorganic carbon (CO(2)). Because this alga is a major food source for predators in acidic habitats, we studied its fatty acid content, which reflects their quality as food, grown under a combination of P-limited and different carbon conditions (either mixotrophically with light + glucose or at high or low CO(2), both without glucose). The fatty acid composition largely depended on the cellular P content: stringent P-limited cells had a higher total fatty acid concentration and had a lower percentage of polyunsaturated fatty acids. An additional limitation for CO(2) inhibited this decrease, especially reflected in enhanced concentrations of 18:3(9,12,15) and 16:4(3,7,10,13), resulting in cells relatively rich in polyunsaturated fatty acids under colimiting growth conditions. The percentage of polyunsaturated to total fatty acid content was positively related with maximum photosynthesis under all conditions applied. The two factors, P and CO(2), thus interact in their effect on the fatty acid composition in C. acidophila, and colimited cells P-limited algae can be considered a superior food source for herbivores because of the high total fatty acid content and relative richness in polyunsaturated fatty acids.     

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

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

Light,nutrients and the fatty acid composition of stream periphyton

1. While the balance of light and nutrients is known to influence the food quality of herbivores by altering algal phosphorus and nitrogen content, the combined effects of light and nutrients on fatty acid synthesis in freshwater periphyton are relatively unknown. In this study, we manipulated light and phosphorus concentration in large, flow‐through experimental streams to examine their effects on both elemental stoichiometry and fatty acid content in periphyton. 2. Two levels of phosphorus (4 and 80 μg L^?1) and three of light (17, 40, 110 μmol photons m^?2 s^?1) were applied in a factorial design in two separate experiments. Diatoms dominated periphyton communities in both experiments, comprising >95% of algal biovolume. Periphyton growth in the streams was simultaneously affected by both resources, even at low rates of supply. 3. Periphyton C/P and C/N ratios increased with light augmentation and decreased with phosphorus enrichment, and consistent with the light : nutrient hypothesis (LNH). Light effects were strongest in streams with low phosphorus concentrations. 4. Periphyton fatty acids reflected the dominance of diatoms : palmitic (16 : 0), palmitoleic (16 : 1ω7) and eicosapentanoic (20 : 5ω3) were the principal saturated (SAFA), monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA), respectively. Linoleic (18 : 2ω6) and linolenic (18 : 3ω3) acids, characteristic of chlorophytes and cyanophytes, were rare, comprising <2% of total fatty acids. 5. Periphyton fatty acid profiles were highly sensitive to light and phosphorus. The proportion of fatty acids comprised by SAFA and MUFA increased with light augmentation and decreased with phosphorus enrichment, whereas PUFA decreased with light and increased with phosphorus. Light effects on fatty acid composition were strongest in phosphorus‐poor streams. PUFA declined with increasing light/phosphorus ratios in the streams, whereas ‘energy’ fatty acids (16 : 0 and 16 : 1) increased. The ratio of SAFA/PUFA was strongly and positively correlated with C/P and C/N ratios. SAFA and MUFA, normalised to dry mass, increased two‐ to threefold with increasing light, while PUFA normalised to dry mass was not significantly affected by light. 6. Similarities in the responses of fatty acids and elemental stoichiometry to light and phosphorus treatments suggested that they were influenced by a common mechanism. Both components of food quality appeared to be sensitive to light‐regulated rates of carbon fixation which, when coupled with insufficient supplies of phosphorus, caused diatom cells to store surplus carbon in SAFA, MUFA and other carbon‐rich compounds that diluted both essential fatty acids and mineral nutrients.     

Essential fatty acid content and the phosphorus to carbon ratio in cultured algae as indicators of food quality for Daphnia

1. A series of experiments examined growth of Daphnia magna on three algal diets ( Rhodomonas minuta , Scenedesmus acutus and Synechococcus sp.) at varying physiological states [nitrogen and phosphorus (P) limitations] to test whether variation in algal fatty acid and/or elemental composition can predict Daphnia growth.
2. These algae differed widely in their essential fatty acid (EFA) composition while phosphorus (P) or nitrogen limitation had only a small influence on their ω3-polyunsaturated fatty acid (PUFA) content.
3. Individually, algal ω3-PUFA content explained 69% of the variation in the normalised growth of D. magna, while algal phosphorus content explained 11% of the variation. Quantitative models for D. magna growth used both algal ω3-PUFA content and algal C : P ratio as food quality indices. Together, algal ω3-PUFA content and C : P ratio explained 70% of the variation in the normalised growth rate of D. magna .
4. Our results indicate that EFA influenced algal food quality much more strongly than P. The EFA and mineral P impacts appear to be independent.     

Essential fatty acids and phosphorus in seston from lakes with contrasting terrestrial dissolved organic carbon content

1. It is often assumed that lakes highly influenced by terrestrial organic matter (TOM) have low zooplankton food quality because of elemental and/or biochemical deficiencies of the major particulate organic carbon pools. We used the biochemical [polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA) – 20:5ω3] and elemental (C : P ratio) composition of particulate matter (PM) as qualitative measures of potential zooplankton food in two categories of lakes of similar primary productivity, but with contrasting TOM influence (clear water versus humic lakes). 2. C : P ratios (atomic ratio) in PM were similar between lake categories and were above 400. The concentration (μg L⁻¹) and relative content (μg mg C⁻¹) of EPA, as well as the particulate organic carbon concentration, were higher in the humic lakes than in the clear‐water lakes. 3. Our results show high fatty acid quality of PM in the humic lakes. The differences in the biochemical quality of the potential zooplankton food between lake categories can be attributed to the differences in their phytoplankton communities. 4. High biochemical quality of the food can result in high efficiency of energy transfer in the food chain and stimulate production at higher trophic levels, assuming that zooplankton are able to ingest and digest the resource available.     

Algal Polyunsaturated Fatty Acids: Food,Pheromones and Foul Odour

Algal polyunsaturated fatty acids (PUFAs) are known to be high‐energy, often‐essential resources to freshwater aquatic food webs. On the other hand, high PUFA cell content in some algal taxa has been linked with the production of derivatives that may act as pheromones, allelogens or toxins. It has been known for some time that these compounds function in marine chemical ecology, but recent evidence indicates that they may play similar roles in freshwater ecosystems. This paper presents field and laboratory studies of planktonic and periphytic communities from freshwater systems, and shows that these communities are rich sources of certain PUFAs and derivatives which may function in both positive and negative foodweb interactions. This also has important implications for surface water quality, as these compounds are potent sources of rancid fishy odours.     

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

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

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

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

两种水华藻——球形棕囊藻和铜绿微囊藻的脂肪酸组成特征与水华形成机制

浮游植物所含的不饱和脂肪酸是测定其作为食物质量的指标,并在浮游植物向浮游动物及其它动物能量转化过程中起着关键的作用,必需不饱和脂肪酸的缺乏有利于水华的形成。球形棕囊藻(Phaeocystis globosa)和铜绿微囊藻(Microcystis aeruginosa)分别是常见的海洋和淡水水华藻类,该文分析了它们在不同生长期的脂肪酸组成,探讨了这两种藻类的脂肪酸组成特征。球形棕囊藻和铜绿微囊藻的脂肪酸碳链长为14~20个碳原子,脂肪酸种类组成都比较简单,以饱和脂肪酸为主,未检测到二十碳五烯酸(Eicosapentaenoic acid,EPA)和二十二碳六烯酸 (Docosahexaenoic acid,DHA)等动物的必需脂肪酸。球形棕囊藻的总脂肪酸含量在247.294~735.44 μg·g^-1干重之间,在对数期和延滞期含量最高的脂肪酸分别是C14:0和C16:0;而两株铜绿微囊藻的总脂肪酸在1 405.095~6 087.617μg·g^-1干重之间,以C16:0含量最高。两株铜绿微囊藻的脂肪酸含量在对数期和延滞期差异明显(p<0.05),但球形棕囊藻的脂肪酸含量在不同生长期差别不大。由于缺乏必需脂肪酸EPA和DHA,球形棕囊藻和铜绿微囊藻不能为高营养级的生物提供必需的不饱和脂肪酸,不是浮游动物等生物的良好食物。因此球形棕囊藻和铜绿微囊藻作为浮游动物的食物质量较低,浮游动物对它们的捕食压力也较小,可能是这两种藻容易暴发水华的重要原因。     

Physiological responses to stoichiometric constraints: nutrient limitation and compensatory feeding in a freshwater snail

Nitrogen (N) and phosphorus (P) are considered to be essential nutrients that control secondary production in various ecosystems; insufficient availability of N and P can limit herbivore growth. Here, data are presented from field samplings and from a laboratory experiment on the potential of primary producers low in P, N, or P and N to constrain growth of the freshwater gastropod Radix ovata . The filamentous green alga Ulothrix fimbriata was cultured under different nutrient regimes, resulting in algae with different C:N:P ratios. The pure algae were fed in high and low quantities to juvenile R. ovata . Low availability of N and especially P in the algae strongly constrained the biomass accrual of the herbivore. In accordance with theoretical predictions, these food quality differences were highly dependent on the food quantity. The snails' growth rate was significantly related to their body C:P ratio, thereby supporting the growth rate hypothesis. R. ovata displayed a pronounced compensatory feeding response to low-nutrient food that could partly dampen but not fully compensate the food quality effects on snail growth. Increased feeding of gastropods at low P and/or N availability leads to depletion of periphyton biomass; hence compensatory feeding would shift the benthic herbivore community from a P or N limitation to a C limitation and thus have whole-ecosystem effects.     

Effects of changes in ambient PAR and UV radiation on the nutritional quality of an Arctic diatom (Thalassiosira antarctica var. borealis)

Polyunsaturated fatty acids (PUFAs) are essential macromolecules that are synthesized by phytoplankton during spring bloom, and they play a key role in the Arctic food web. They are, however, considered to be sensitive to oxidation by UV radiation (280-400 nm). Changes in the food quality of primary producers may affect the transport of biomass and energy in the whole ecosystem. Using a common Arctic diatom, we looked at the effect of ambient and increased UV radiation on its nutritional quality, specifically, the fatty acid composition and elemental ratios. In May 2004, in the archipelago of Svalbard (79° N), a unialgal culture of Thalassiosira antarctica var. borealis was subjected to a 17-day experiment in outdoor aquaria. The diatoms were kept in semi-continuous culture (40 1) and exposed to three treatments with different levels of UV radiation: none (UV-shielded), ambient, and enhanced. Fatty acid composition, C:N:P ratios, photosynthetic pigment composition, optimum quantum yield of PSII, and cell numbers were analysed over the experimental period. An initial increase in PAR (photosynthetically active radiation, 400-700 nm) intensities profoundly affected the fatty acid composition and substantially inhibited the synthesis of PUFAs, but the relative amounts of PUFAs were not reduced by UV radiation. Enhanced UV radiation did, however, cause a significant reduction in optimum quantum yield of PSII and affected some fatty acids, mainly 18:0 and 16:1 n-7, during the first week of the experiment. Both ambient and enhanced UV radiation caused significantly lower C:P and N:P ratios. At the same time, these treatments elicited a higher relative content of the photoprotective pigments diadinoxanthin and diatoxanthin. After acclimation to the new light levels these effects faded off. Thus, brief periods with high light exposure may cause significant changes in photosynthetic activity and food quality, but the capacity for photo-acclimation seems high. The impact of UV radiation seems to be less important for food quality than that of PAR during a sudden rise in total light intensity.     

Lipid-ovary indices in food-limited Daphnia

The amount of lipid carried by a zooplankton individual is highlyvariable. Various lines of evidence have suggested that thenutritional condition of animals could be ascertained via thedetermination of lipid content. However, few studies have beenconducted where the lipid levels of animals of known conditionhave been measured. Here, we report on the lipid index and combinedlipid-ovary index values of Daphnia obrusa reared on foods madeup of varying quantity and quality of a unialgal (Scenedesmusacutus) diet. Lipid levels differed in different food types.However, in no cases were these indices positively correlatedto growth rate, such as would be necessary if one were to usethem to diagnose food limitation in the field. Thus, we urgecaution in the utilization of either the lipid or lipid-ovaryindex until more is known about the role of fatty acids in thediets of these animals.     

Effects of fatty acid and phosphorus content of food on the growth, survival and reproduction ofDaphnia

1. Mixtures of microencapsulated lipids and Scenedesmus quadricauda grown at different degrees of P limitation were used as food for Daphnia galeata in two growth experiments. Thereby, food quality in terms of ω3-fatty acid (ω3-FA) or phosphorus (P) content could be assessed without interference from other factors.
2. ω3-highly unsaturated fatty acids (ω3-HUFA), given to Daphnia as fish oil or eicosapentaenoic acid (EPA) together with non P-limited algae, decreased the time to first reproduction. When fed fish oil, somatic growth and survival were also enhanced. Linolenic acid also decreased the time to first reproduction but to a lesser extent than EPA.
3. Food quality depended to a large extent on the degree of P limitation of Scenedesmus , which is consistent with P limitation of Daphnia. The overall impact of P was always larger than the effect of ω3-FA. Growth, survival and reproduction were elevated when Daphnia were fed non P-limited Scenedesmus compared to treatments with P-limited algae.
4. The relative importance of ω3-HUFA and P content in the food changed over a C : P gradient, with stronger effects of ω3-HUFA at low C : P ratios.     

Rapid adaptation of herbivore consumers to nutrient limitation: eco‐evolutionary feedbacks to population demography and resource control

Humans alter biogeochemical cycles of essential elements such as phosphorus (P). Prediction of ecosystem consequences of altered elemental cycles requires integration of ecology, evolutionary biology and the framework of ecological stoichiometry. We studied micro‐evolutionary responses of a herbivorous rotifer to P‐limited food and the potential consequences for its population demography and for ecosystem properties. We subjected field‐derived, replicate rotifer populations to P‐deficient and P‐replete algal food, and studied adaptation in common garden transplant experiments after 103 and 209 days of selection. When fed P‐limited food, populations with a P‐limitation selection history suffered 37% lower mortality, reached twice the steady state biomass, and reduced algae by 40% compared to populations with a P‐replete selection history. Adaptation involved no change in rotifer elemental composition but reduced investment in sex. This study demonstrates potentially strong eco‐evolutionary feedbacks from shifting elemental balances to ecosystem properties, including grazing pressure and the ratio of grazer:producer biomass.     

Survival,growth and reproduction of Daphnia galeata feeding on single and mixed Pseudomonas and Rhodomonas diets

1. Bacteria can be an important resource for zooplankton production in aquatic food webs, although the degree to which bacteria sustain zooplankton growth and reproduction is not clear. We performed a growth experiment with Daphnia galeata feeding on different ratios of P‐replete Pseudomonas and Rhodomonas, ranging from a 100% bacterial to a 100% algal diet. 2. A pure bacterial diet did not support survival, growth or reproduction of D. galeata. While a 20% share of Rhodomonas in the food allowed survival of daphniids, the occurrence of offspring on a 50% algal diet indicated that the threshold for successful reproduction was between those two proportions of algal food. Increasing the proportion of the alga further increased growth and reproductive output, indicating that Rhodomonas was a higher‐quality food than Pseudomonas. 3. A subsequent labelling experiment demonstrated that D. galeata incorporated phosphorus from Pseudomonas and Rhodomonas with similar efficiency, whereas carbon was incorporated more efficiently from Pseudomonas than from Rhodomonas. 4. we hypothesise that inadequate levels of essential biochemicals in pure bacterial diets led to decreased Daphnia performance. Concentrations of fatty acids in general, and especially of polyunsaturated fatty acids, were much lower in Pseudomonas than in Rhodomonas. This difference could explain the different growth and reproduction responses, although limitation by other essential biochemicals (e.g. sterols) cannot be ruled out. 5. Hence, where they dominate, bacteria may provide a significant part of the elemental flux to species feeding higher in the food web on the short term. However, the performance of consumers may be constrained by essential biochemicals.     

Complementary information from fatty acid and nutrient stoichiometry data improve stream food web analyses

Hydrobiologia - Our study characterized the food web of a shaded stream to investigate consumer food choices and food source quality. We measured fatty acids (FA) and elemental carbon (C), nitrogen...